Immigrant Tortricidae: Holarctic versus Introduced Species in North America

Simple Summary The family Tortricidae includes approximately 11,500 species of small moths, many of which are economically important pests worldwide. A large number of tortricid species have been inadvertently introduced into North America from Eurasia, and many have the potential to inflict considerable negative economic and ecological impacts. Because native species behave differently than introduced species, it is critical to distinguish between the two. Unfortunately, this can be a difficult task. In the past, many tortricids discovered in North America were assumed to be the same as their Eurasian counterparts, i.e., Holarctic. Using DNA sequence data, morphological characters, food plants, and historical records, we analyzed the origin of 151 species of Tortricidae present in North America. The results indicate that the number of Holarctic species has been overestimated by at least 20%. We also determined that the number of introduced tortricids in North America is unexpectedly high compared other families, with tortricids accounting for approximately 23–30% of the total number of moth and butterfly species introduced to North America. This suggests that introduced tortricids have a greater potential of becoming economically important pests than moths in other families, and why distinguishing Holarctic from introduced species is critical to American agriculture. Abstract In support of a comprehensive update to the checklist of the moths of North America, we attempt to determine the status of 151 species of Tortricidae present in North America that may be Holarctic, introduced, or sibling species of their European counterparts. Discovering the natural distributions of these taxa is often difficult, if not impossible, but several criteria can be applied to determine if a species that is present in both Europe and North America is natively Holarctic, introduced, or represented by different but closely related species on each continent. We use DNA barcodes (when available), morphology, host plants, and historical records (literature and museum specimens) to make these assessments and propose several taxonomic changes, as well as future areas of research. The following taxa are raised from synonymy to species status: Acleris ferrumixtana (Benander, 1934), stat. rev.; Acleris viburnana (Clemens, 1860), stat. rev.; Acleris pulverosana (Walker, 1863), stat. rev.; Acleris placidana (Robinson, 1869), stat. rev.; Lobesia spiraeae (McDunnough, 1938), stat. rev.; and Epiblema arctica Miller, 1985, stat. rev. Cydia saltitans (Westwood, 1858), stat. rev., is determined to be the valid name for the “jumping bean moth,” and Phiaris glaciana (Möschler, 1860), comb. n., is placed in a new genus. We determine that the number of Holarctic species has been overestimated by at least 20% in the past, and that the overall number of introduced species in North America is unexpectedly high, with Tortricidae accounting for approximately 23–30% of the total number of Lepidoptera species introduced to North America.


Introduction
Globalization, the international movement of commodities and people among different nations, has dramatically increased the spread of plant and animal species around the world, e.g., [1][2][3]. Among these "introduced" or "exotic" species, insects are the most pervasive, representing 87% of the approximately 2500 nonnative terrestrial invertebrates in Europe [4]. According to Lovett et al. [5], nonnative insect species have accumulated in United States forests at a rate of approximately 2.5 per year over the last 150 years, with the gypsy moth (Lymantria dispar (Linnaeus, 1758), Erebidae) being among the most notorious. Examples of important lepidopteran pests that have been introduced recently to new regions around the globe include the fall armyworm, Spodoptera frugiperda (J. E. Smith) (Noctuidae), a native of the New World that has spread to much of Africa and Asia [6,7] and was recently discovered in Australia [8]; the Old World bollworm, Helicoverpa armigera (Hübner) (Noctuidae), which was first reported in the New World in Brazil and has spread to much of South America and the Caribbean [9][10][11]; the light brown apple moth, Epiphyas postvittana (Walker) (Tortricidae), a native of Australia that was documented from California in 2006 and has now spread throughout much of the state [12]; and the European grapevine moth, Lobesia botrana (Denis and Schiffermüller) ([Tortricidae]), a native of Europe that was inadvertently introduced to the wine-growing regions of Argentina, Chile, and California [13]. While these contemporary or recent arrivals to new regions are well documented, for many other species, there is considerable ambiguity regarding their origin and/or native distributions. For example, in North America, the discovery of a "European" species could indicate that the taxon was recently introduced, that it is natively Holarctic in distribution and previously undiscovered, or that it represents an unrecognized sibling species of its European counterpart. In the absence of direct evidence of an introduction, the taxonomist is left to speculate among these alternative explanations. In many cases, a compelling explanation is further complicated by the fact that many apparently European species are discovered along the northwestern or northeastern coasts of North America, regions where one would expect to find Holarctic elements or introductions owing to the proximity to major U.S. and Canadian ports of entry.
In support of a comprehensive update to the checklist of the moths of North America [14], we determine the status of 151 species of Tortricidae present in North America that have been previously assumed to be Holarctic, or introduced, or whose status in the Nearctic was questionable. We use DNA barcodes (when available), morphology, host plants, and historical records (literature and museum specimens) to make these assessments, and we propose appropriate taxonomic changes where necessary.

Biogeographic Framework
The Holarctic. The Holarctic is defined as the biogeographical region comprised of the Nearctic (North America) and the Palearctic (Eurasia and northern Africa). These continents have been variably linked since the breakup of Pangea approximately 180 mya. The following geographic history of the region is summarized from Hopkins [15], Enghoff [16], and Sanmartín et al. [17]. Following the breakup of Pangea, North America and Eurasia comprised the northern supercontinent Laurasia, which was split into two smaller paleocontinents, Euramerica (Europe and eastern North America) and Asiamerica (Asia and western North America) around 100-80 mya. The western Palearctic was separated from the eastern Palearctic by the Turgai Sea until ca. 30 mya, and the western Nearctic was separated from the eastern Nearctic until ca. 60 mya by the Mid-Continental Seaway. Europe and eastern North America were linked by several North Atlantic land bridges until sometime in the late Eocene (ca. 39 mya). Asia and western North America were connected by the Bering land bridge from the mid-Cretaceous (100 mya) continuously until the Late Pliocene (3.5 mya), but this connection was established again intermittently throughout the Pleistocene (1.5-0.011 mya) before it was submerged completely ca. 11,000 years ago.
These Trans-Beringian land bridges are assumed to have played an important role in dispersal across the Holarctic. Beringia underwent three distinct phases with associated changes in climate and vegetation [17]. Beringian Bridge I: A continuous belt of boreotropical forest extended over the entire Northern Hemisphere by the Early Eocene (ca. 50 mya), and the climate was much warmer and more humid than today. Beringian Bridge II: As the climate cooled and became drier, the forests transitioned to mixed deciduous hardwoods and conifers, and eventually only to conifer forests that were split into eastern and western portions. The marine transgression of the Bering Strait in the Late Pliocene (3.5 mya) permanently separated the Palearctic and Nearctic forests. Beringian Bridge III: As glaciation commenced in the Pleistocene (1.5-1.0 mya), the Beringian land bridge was once again established. Originally assumed to be a continuous "mammoth steppe" or "steppe tundra," Elias et al. [18] found that the vegetation was dominated by birch-heath-graminoid tundra with little or no evidence of steppe elements. This land connection persisted intermittently during glaciation events until it was permanently interrupted at the end of the Pleistocene.
It is generally assumed that trans-Beringian dispersal between the eastern Palearctic and western Nearctic is more frequent than trans-Atlantic dispersal events. Sanmartín et al. [17] found that in the Early Tertiary (70-45 mya), the North Atlantic connection was more important than the Beringian connection, but that faunal exchange was more frequent across the second Beringian Bridge (20-3.5 mya), and much more frequent across the third Beringian Bridge (1.5-0.011 mya). This is similar to what has been proposed for plants, where Beringia was more important than the North Atlantic bridges since the Middle Eocene (45 mya) [19]. During full-glacial periods in the Pleistocene, Beringia was essentially part of Siberia, with the unglaciated parts of Alaska and the Yukon separated from the rest of North America, which was largely covered in ice sheets. As glaciers receded across North America, Palearctic or Beringian insects had the opportunity to disperse from former refugia, events that have occurred relatively recently. When examining ant species, Schär et al. [20] determined that all three of the naturally occurring Holarctic ant species had dispersed from the Palearctic no more than 2 mya. Studies of Noctuidae have found that most naturally occurring (e.g., not introduced) Holarctic species are associated with the tundra zone, and all truly Holarctic noctuid species are present somewhere in Beringia [21].
Holarctic or Introduced? Determination of whether a species currently found in both Eurasia and North America is Holarctic versus introduced may, in many cases, represent an unsolvable puzzle. Although the association of a species with Beringia lends support to a Holarctic distribution, other factors require consideration in determining a species' natural distribution. These factors include the organism's current range, its host plant use, available historical records, morphological and/or molecular variation, etc.
Factors that would be expected for a naturally occurring Holarctic species include the following: (1) no direct evidence of introduction; (2) association with native occurring host plants; (3) first reported in noncoastal, inland areas; (4) no evidence of range expansion in recent times; and (5) presence in the Arctic biogeographical region.
Factors associated with introductions, as would be expected for taxa that were transported by man between or among continents, include the following: (1) direct evidence of introduction; (2) association with nonnative or frequently imported host plants; (3) first documented in a coastal area; (4) evidence of range expansion in recent times; and (5) absence from the Arctic biogeographical region.
Although the above criteria can be applied to hypothesize a taxon's natural distribution, the most important factor may lie in an initial determination of whether more than one species is involved. Early taxonomists in North America often sought the opinions of their European counterparts. For example, in Tortricidae, the opinions of Lord Walsingham and P. C. Zeller, European authorities in the latter half of the nineteenth and early twentieth centuries, strongly influenced the taxonomy of New World species and genera. Superficial similarity between a Nearctic and Palearctic species was sometimes enough to assign them to the same taxon, particularly before the discovery of the value of genitalia for identification. One of the most striking examples of a North American taxon hiding under a European name is the Nearctic Paralobesia viteana (Clemens) and the Palearctic Lobesia botrana. Both moths are pests of grape and have similar wing patterns, but their genitalia are very different, and a recent phylogenetic study has shown that Lobesia and Paralobesia are indeed separate genera [22]. In a paper on the North American species viteana, Zeller [23] determined that it was the same as the European botrana and synonymized them, with botrana as the senior name. What was far from understood at the time was that the early concept of P. viteana included numerous other species of Paralobesia, all with similar wing patterns. A recent revision of this genus resulted in the identification of 40 different species in North America [22]; thus Zeller had inadvertently synonymized up to 40 different Nearctic taxa under a single European name, none of which were actually Holarctic. The advent of studying genitalia helped to resolve some of the more obvious problems with taxon assignment in Tortricidae; however, genitalia were not always a panacea, as the amount of morphological variation in these structures was not well studied, a problem that remains today. Thus, even Heinrich [24,25], in his two monographs on North American Olethreutinae in which he examined genitalia for nearly every species, incorrectly identified some endemic Nearctic taxa as similar European species.

Taxa Evaluated
The list of tortricid taxa examined for this study was obtained primarily from the Checklist of the Moths of North America [14] and the World Catalogue of Tortricidae [26], with updates from the web-based catalogue [27], the preliminary "P3" updated checklist of the Lepidoptera of North America [28], and unpublished notes and partial manuscripts prepared by the late William E. Miller. Numerous other publications were referenced for each individual taxon; these are listed in the species accounts. Determination of the "previous status" for each species was obtained primarily from the checklist of Pohl et al. [28], with some minor modifications based on our knowledge of particular taxa or obvious mistakes in the "P3" list. Taxon order follows the "P3" list and the updated North American checklist that is in preparation. The single institutional abbreviation used in the text is USNM = U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Molecular Methods
Although the use of genitalia in lepidopteran systematics was a giant step forward, the use of molecular data represents an even greater advancement. While new DNA sequencing technologies continue to make the sequencing entire genomes more affordable, even simple DNA barcodes can be highly informative. DNA barcodes [29] represent an excellent source of molecular data, complementary to morphology, because sequences are easy to obtain, even from older museum specimens, and a large database of reference sequences has been assembled over the last decade (Barcode of Life Database; BOLD). Patterns in DNA sequences can be used as evidence regarding the status of morphologically similar populations in Europe and North America, either directly or through the discovery of morphological characters that corroborate similarities in DNA sequences. Within Tortricidae, several recent studies have used DNA barcoding to determine if two or more entities represent a single Holarctic taxon or are similar species occurring on different continents, e.g., [30][31][32][33].
As with genital morphology, DNA barcodes are not without shortcomings with regards to their ability to identify species boundaries. Early studies (e.g., [29,34]) proposed fixed distance thresholds between species (e.g., 2%) and "barcoding gaps" between different taxa. Although these early concepts have been largely discarded (see summary in [35]), DNA barcodes remain an extremely informative source of evidence of genetic similarity among closely related species.
DNA barcode sequences were obtained from BOLD. All records for a particular taxon, usually at the genus level, were selected with the option "include Public" records. Records were filtered to exclude contaminants, records with stop codons, and records flagged as misidentifications or errors. Only sequences ≥400 bp were used because of random alignment issues with shorter sequences. Sequences were aligned in BOLD using the BOLD amino acid-based HMM Aligner, and neighbor-joining trees were constructed in BOLD using the Kimura 2 Parameter (K2P) distance model. Sequences of A. holmiana form a single BIN, with specimens from Europe (Germany, Norway, Finland, Italy) and North America (Washington, British Columbia). There is little sequence variation, and many specimens share identical barcodes. The first records of A. holmiana in North America are from the Lower Fraser Valley, British Columbia in 1977 [38,39]. Miller (unpublished) estimated the actual introduction to be three years earlier (1974).
Specimens of A. notana in BOLD are placed in a single BIN (BOLD:AAB4490) with representatives from Europe (Austria, Finland, Germany, Norway, United Kingdom). These form a distinct cluster close to sequences of A. ferrugana from Europe (under which A. notana was previously synonymized). Razowski [47] states that A. notana was introduced into North America. Pohl et al. [46] state that records of A. notana from North America may refer to A. arcticana (see [48]), and that Winn [49] used the name A. ferrugana for records of A. notana from Canada. The DNA data are not sufficient to support or refute any of these statements because there are no specimens from North America of this taxon in BOLD.
Sequences identified as A. schalleriana cluster into two BINs: BOLD:AAG9608 contains specimens from Europe (Germany, Norway, Bulgaria, and Finland) and BOLD:AAB2825 contains specimens from Canada and the United States. The two BINs are separated by 2.9% (p-dist) and together form a single cluster.
Razowski [47] treats A. schalleriana as Holarctic with a different subspecies in Europe and North America (i.e., nominotypic schalleriana in Europe, viburnana in North America). Based on the sequence data, it is likely that the Palearctic and Nearctic populations represent different species. As such, we resurrect A. viburnana (Clemens, 1860), stat. rev., to species status and assign the specimens represented by BOLD:AAB2825 to this name. Acleris schalleriana is represented by BOLD:AAG9608 and restricted to the Palearctic.  [44] states that "A. capizziana Obr. is very difficult to determine when similarly coloured as A. lipsiana (Schiff. and Den.)," and also noted that the male genitalia of these two species are very similar.

Acleris variegana ([Denis and
Sequences from specimens identified as either A. effractana or A. emargana fall into three BINs: BOLD:AAB3524 contains specimens from Europe (Austria, Germany, Finland, France, Italy, Norway); BOLD:AAB3525 contains specimens from North America (British Columbia, Quebec, Washington, Yukon Territory) and Europe (Finland, Norway); and BOLD:AAA6999 contains specimens from Finland and Norway.
Karsholt et al. [54] resolved the taxonomy of these two species. They determined that A. emargana and E. effractana could be reliably separated by morphological characters (length of the socii in the males and length of the ductus bursae in the females). Acleris emargana is Palearctic and distributed throughout Europe and Asia, and A. effractana is Holarctic and distributed throughout North Europe, northern Central Europe, Russia, Japan, Canada, and the northwestern United States [47].
DNA data support the findings of Karsholt et al. [54]. BOLD:AAB3524 represents E. emargana from Europe. BOLD:AAB3525 represents E. effractana and forms two groups within a single cluster, one containing specimens from Europe and the other containing specimens from North America. There is, however, a third BIN that likely represents a different species. BOLD:AAA6999 forms a separate cluster that is sufficiently distinct (2.75% and 3.98% p-dist) from the other two BINs to indicate it is a potential cryptic species from Finland and Norway. Specimens in BOLD included in this BIN are identified as either A. emargana, A. effractana, or Acleris sp. Sequences of C. longana are represented in a single BIN, BOLD:AAC7742. These include specimens from North America (California, Washington) and Europe (Austria, Denmark, Germany, Norway, United Kingdom). Many of the specimens from Europe are identified as C. genitalana (and a few as C. longana). These two species can be separated by genitalia, and a few of the European specimens are dissected, indicating that at least some of the identifications in BOLD are correct. Thus, it appears that barcodes of C. longana and C. genitalana are nearly identical and cannot be used to reliably separate these two taxa.
Cnephasia longana was first reported from California by Pritchard et al. [55] and from British Columbia by Cram and Tonks [56]. Powell [57] [58] provided a history of C. stephensiana and C. asseclana (as C. interjectana) in North America. This species was first discovered in North America when it was collected in Nova Scotia in 1954, and it was found in several locations in eastern Canada in the late 1970s [58]. Cnephasia stephensiana is currently distributed throughout the northeastern U.S. and eastern Canada. BOLD:AAA8425 is divided into two distinct clusters: the first includes specimens from Canada (Alberta, British Columbia, Yukon Territory) and Washington; the second includes specimens from Europe (Austria, Finland, Germany, Italy, Macedonia). The mean variation within each of these clusters is 0.14% (p-dist), but they are separated by approximately 1% (p-dist) and treated as separate entities in BOLD's cluster analysis. The other BIN, BOLD:ACE7529, is >3.3% (p-dist) from the main BIN and almost certainly contains misidentified specimens or an undescribed species. Based on the DNA data, it appears that E. argentana is restricted to the Palearctic, and specimens identified as this taxon in North America represent one or more cryptic species that have yet to be described. This is not surprising, given that many of these specimens are found at higher elevations and in isolated mountain habitats. There are approximately 200 sequences representing specimens identified as E. osseana in BOLD. These specimens are placed in four BINs: BOLD:AAA6265 contains specimens from Europe (Austria, Finland, Germany, Italy, Norway, Slovenia, United Kingdom); BOLD:AAA6264 contains specimens from Canada (Alberta, British Columbia, Newfoundland, Nova Scotia) and the United States (Alaska and California); BOLD:ABZ7263 contains specimens from Alberta and Yukon Territory; and BOLD:AAA4129 contains specimens from Canada (British Columbia, Newfoundland, Manitoba, Alberta, and Yukon Territory), the United States (Alaska and Utah), and Russia.
The BIN from Europe, BOLD:AAA6265, forms a distinct cluster that is >3.30% (p-dist) divergent from the other E. osseana BINs. Its nearest neighbor is actually E. argentana. Based on the geographic separation of the other BINs, it is likely BOLD:AAA6265 represents "true" E. osseana, which was described from Slovenia. This would indicate that E. osseana is Palearctic in distribution and not Holarctic.
The remaining three BINs (BOLD:AAA6264; BOLD:ABZ7263; BOLD:AAA4129) form a large cluster that also contains BOLD:ACE7529 (identified as E. argentana) and several other Eana specimens that are not placed in BINs. It is likely that this cluster represents several species. The largest group, BOLD:AAA4129, represents a taxon that is possibly Beringian, given that it is distributed across Northwest Canada and Alaska to Russia (the sequences from Russia are high quality and full length, and thus appear to be placed correctly). BOLD:AAA6264 and BOLD:ABZ7263 seem to be restricted to North America and also possibly represent new taxa.
There are more than 20 junior synonyms placed under E. osseana currently. Thus, it is possible that at least some of these names could be applied to these other BINs. Pohl et al. [46] state that the subspecies niveosana "applies to all North American populations, and may warrant full status." It is likely that this name could be applied to one of the North American groups, but without further research, it is not clear to which BIN it should be assigned. Similar to E. argentana, many of these specimens are found at higher elevations and genitalic variation is high in this group. Further research will be necessary to resolve these Eana species complexes. Specimens of A. deutschiana are placed in several BINs: five BINs contain specimens from North America and a single BIN (BOLD:AAB4420) contains specimens from Europe. The five BINS form two distinct clusters, with one representing Europe and the other North America. Thus, it appears that A. deutschiana is Palearctic and not present in North America. The large amount of genetic diversity in the North American cluster suggests the possibility of several cryptic species. There are two junior synonyms of A. deutschiana described from Labrador that could potentially be applied to one or more of the North American BINs. Sequences of A. rutilana form a single BIN (BOLD:AAC1488), with shallow splits between North American representatives (i.e., British Columbia and Quebec) and two European clusters that overlap in geographic distribution. Razowski [60] described the subspecies Aethes rutilana canadiana from specimens from Quebec, Ontario, Manitoba, and Alberta, and though he did not explicitly state so, it seems as though he intended the name to represent a North American subspecies. Aethes rutilana was introduced into British Columbia sometime prior to 1965 [41]. All sequences of A. smeathmanniana reside in a single BIN (BOLD:AAB1945), with specimens from Europe and North America, indicating it is likely Holarctic, as concluded by Razowski [60]. In the Nearctic region, this species is recorded from Canada (Alberta, British Columbia, Quebec, Yukon Territory) and the U.S. (California, Maine, Minnesota, Montana, Ohio, New Jersey). In the Palearctic region, it ranges across the northern latitudes (e.g., Finland, Norway, Sweden) and higher elevations (e.g., the alps of Italy and Switzerland). Sequences of A. zoegana are placed in six BINs in BOLD. There is a significant amount of variation between BINs (averaging 3% p-dist), but all six BINs form a single cluster. It is not known if this variation is indicative of different geographic populations of the same species, or if one or more cryptic species are involved. Regardless, the taxon currently recognized as A. zoegana has a well-documented history of being released in North America as a biocontrol agent.
According to Story [61] and Story et al. [62], specimens of A. zoegana from Austria and Hungary were introduced near Stevensville, Montana in July 1984 for the biological control of star-thistles (Centaurea sp.; Asteraceae), which had become a weedy pest in rangelands. By 1991, A. zoegana was well established in Montana, and according to Wilson and Randall [63], it is now present "in most western states." We have verified records from Montana, Washington, and British Columbia. Sequences of C. subroseana form a single BIN (BOLD:AAD2823) that includes specimens from Italy, Austria, Finland, Norway, and South Korea; there are no sequence data from North American specimens. Although Pogue [64] considered our single North American representative to be an undescribed species, Razowski [60] reported it from Ontario and Alberta as Cochylidia subroseana.
We examined specimens from Colorado that appear to be conspecific with C. subroseana. Hence, we suspect that it is indeed Holarctic. In the Palearctic, this species is widely distributed from the United Kingdom and Norway across northern and central Europe to China. Sequences of C. atricapitana are represented by four BINs in BOLD, which include specimens from Europe (Denmark, The Netherlands, Norway, Russia, United Kingdom). There are no sequence data for any North American specimens. Each of the BINs contains only a few specimens, and they all form a single cluster; we do not know if this indicates a single variable species or multiple taxa. According to De Clerck-Floate and Cárcamo [65], this species was introduced into British Columbia from Europe in 1990 for biocontrol of tansy ragwort (Jacobaea vulgaris; Asteraceae). Sequences of N. dubitana form in a single BIN (BOLD:AAC3780) with a shallow split between North American representatives (i.e., British Columbia and Washington) and European specimens (i.e., Austria, Finland, Germany, Norway, Italy, United Kingdom); however, specimens from Ontario, Newfoundland, Labrador, and Nova Scotia appear more similar to European specimens than to other North American specimens.

Neocochylisdubitana (Hübner, 1799) Previous status: Holarctic
Neocochylis was proposed by Razowski [66] as a subgenus of Cochylis, with the type species Conchylis calavrytana Rebel, 1906. It later was synonymized with Cochylis by Razowski [67], and recently redefined and elevated to generic status by Brown et al. [68], who included dubitana in the genus.
Neocochylis dubitana is assumed to be Holarctic in distribution [50,60], ranging throughout much of Europe to Asia (China) and throughout much of northern North America. Razowski [60] reported it from Quebec, Alberta, and British Columbia in Canada, and we have examined specimens from Maine in the Northeast, and from Oregon, California, Colorado, Montana, South Dakota, and Wyoming in the West.

BIN: N/A
Eugnosta argyroplaca is not represented in BOLD. Brown [69] provided the following summary of the status of this species. Euxanthis argyroplaca Meyrick, 1931 was described from a single male from "Usgent, Arizona," U.S.A., but it has not been collected since. Because of its U.S. type locality, the species has long been cited in checklists and catalogs of North American Tortricidae. The male holotype is deposited in the Natural History Museum, London, and the adult and its genitalia were illustrated by Clarke [70].
The species resided in Euxanthis until Powell [71] transferred it to Carolella Busck. Pogue [64] subsequently transferred it to Eugnosta Hübner in his unpublished Ph.D. dissertation. Because Eugnosta and Carolella are now considered synonyms [68], with Eugnosta being the senior synonym, the generic assignment of the species has stabilized. Nonetheless, in a recent list of the Cochylini (Cochylina) of the U.S. and Canada, Metzler and Brown [72] incorrectly returned it to "Cochylini unplaced" without comment. Based on the male genitalia of the holotype, the species is unquestionably assigned correctly to Eugnosta. However, there is some doubt regarding the geographic origin of the holotype. In facies, it is similar to several Palaearctic species of Eugnosta (e.g., E. lathoniana, E. hydragyrana, E. medvedevi, and E. magnificana), with a forewing pattern that features extensive areas of silvery white (e.g., [67]). This forewing pattern is not known to occur in New World Eugnosta. As mentioned previously, the holotype is the only North American record of the species, and the type-locality of "Usgent" could not be located in any map or gazetteer. Although it is possible that "Usgent" is the collector, in all comparable descriptions in Meyrick's Exotic Microlepidoptera, collectors are given in italics and in parentheses. These lines of evidence seem to suggest that E. argyroplaca is a Palearctic species with incorrect collecting information associated with the type. There are no barcode data for this species. This species was described from Durango, Mexico, far south of the U.S. border. Among the large number of specimens of Eugnosta we have examined, no U.S. or Canadian specimens match the description and/or illustrations provided by Razowski [73].

Phtheochroa vulneratana (Zetterstedt, 1839) Previous status: Holarctic
Revised status: Phtheochroa vulneratana (Palearctic); North American cryptic species (Nearctic) Supported by DNA in BOLD: Yes BINs: BOLD:AAD9720 (Europe); BOLD:AAD8609 (North America) Sequences of P. vulneratana are represented by two BINs, one of which (BOLD:AAD9720) includes only Palearctic specimens (Finland, Italy, Norway, Switzerland) and the other (BOLD:AAD8609) only Nearctic specimens (Canada and U.S.). Powell [74] first proposed that P. vulneratana is a Holarctic boreal species, reporting it from Alaska and Colorado; perhaps the earliest record is in the USNM from Colorado, identified as P. vulneratana by Jack Clarke in 1967. However, the two BINs containing P. vulneratana in BOLD are separated geographically and by >2% (p-dist). Hence, barcode data strongly support the hypothesis that Nearctic and Palearctic specimens represent two different species.
According to Razowski [50,67], P. vulneratana ranges from Scandinavia south through the highest mountains of central and southern Europe (e.g., Switzerland, Austria, Italy) and east through the Balkan, Pamir, and Altai mountains, through Siberia, Mongolia, and Amur Territory, to Japan. We have seen North American records identifying this species from British Columbia and Alberta south through the Rocky Mountains to Colorado, at elevations ranging from about 4500 to 12,000 feet; however, it is likely these represent a different taxon.

Rolandylis maiana (Kearfott, 1907) Previous status: Holarctic?
Revised status: Nearctic Supported by DNA in BOLD: Yes BINs: BOLD:AAM0777 The only sequences of R. maiana in BOLD are from specimens collected in Ontario and are placed in a single BIN (BOLD:AAM0777), along with specimens identified as R. fusca, with the latter likely being a misidentification.
Gibeaux [75] described the genus Rolandylis for the species R. catalonica Gibeaux, 1985. However, there is no question that catalonica is conspecific with the North American species maiana Kearfott, 1907. Hence, Gibeaux's specimen (from France) was either mislabeled or represents an inadvertent introduction of this species to Europe. The former seems more likely. Pohl et al. [28] listed Spinipogon thes Razowski and Becker, 1983 as present in North America. This record is based on a specimen in the USNM collected in Cameron Co., Texas and identified as S. thes (type locality Santa Catarina, Brazil). Upon further examination, the specimen in the USNM from Texas is almost certainly S. harmozones (type locality Nuevo Leon, Mexico). This is the first record of this species from the U.S. There are no DNA barcode data in BOLD for this taxon (the Texas specimen was submitted but sequencing failed). Sequences of T. nana are placed in two BINs in BOLD: BOLD:ABY5998 contains specimens from Europe (Austria, Finland, Germany, Norway, United Kingdom) and Canada (Nova Scotia, Quebec), and BOLD:AAB3573 contains specimens from North America (Canada and U.S.). These BINs form a single cluster and likely represent a single taxon. In the Palearctic, this species is known from Europe, Asia Minor, Siberia, and the Russian Far East [50]. In North America, this species is found in the northeastern U.S., south to Tennessee, and west to Minnesota. This species has traditionally been treated as Holarctic; however, because it is not recorded near Beringia or anywhere in western North America, it may be introduced. All of the sequences for E. ministrana fall into a single BIN: BOLD:AAA7315. There is a substantial amount of sequence variation, with sequences from several localities forming distinct subclusters, but there is no clear separation between North American and European specimens. This appears to be a single, although genetically variable, Holarctic species.

Eulia ministrana (Linnaeus, 1758) Previous status: Holarctic
As currently defined, Eulia is monotypic and Holarctic in distribution, ranging from Great Britain across northern Europe and northern Asia to Siberia, and south to Japan [76][77][78]. In North America, it has been recorded from Canada (Alberta, British Columbia, Ontario, Quebec, New Brunswick, Northwest Territories, and Nova Scotia) and the northern United States (Alaska, Washington, Oregon, Idaho, Utah, Pennsylvania, New York, New Hampshire, Maine), ranging south in the eastern U.S. to Virginia, Tennessee, and North Carolina. Sequences of P. cerasana form a single BIN (BOLD:AAA3660), with specimens from North America (British Columbia) sharing nearly identical sequences with those from Europe (Austria, Belgium, the Czech Republic, Finland, Germany, Italy, The Netherlands, and the United Kingdom). The earliest record of P. cerasana from North America is from Victoria, British Columbia in 1964 [79]. Mutuura [80] was apparently unaware of Evans' report (using the junior synonym P. ribeana), and stated that the earliest introduction of P. cerasana to British Columbia was in 1965. This and the next taxon are common, widespread pest species.

Pandemis heparana ([Denis and Schiffermüller], 1775) Previous status: Introduced from Eurasia
Revised status: No change Supported by DNA in BOLD: Yes BINs: BOLD:AAA9254 (Europe, North America); BOLD:ACM3812 (Japan) Similar to P. cerasana, most sequences of P. heparana form a single BIN: BOLD:AAA9254, with little sequence variation. Europe specimens are from Austria, the Czech Republic, Finland, Germany, Italy, The Netherlands, Norway, and the United Kingdom. These specimens share identical sequences with those from British Columbia and Washington. There are three specimens identified as P. heparana from Japan that are assigned to a separate BIN (BOLD:ACM3812). At least one of these appears to be correctly identified, and P. heparana has been recorded from Japan. The two BINs are only 1.65% different (p-dist); thus, this could simply represent geographic variation. More sequence data from Asia are needed to evaluate the status of populations in this region.
Pandemis heparana was first discovered in British Columbia in 1978 [80]. This species was reared from larvae collected on apple, crabapple, pear, plum, and species of Prunus, Crataegus, Lonicera, Rubus, Vaccinium, and Spiraea from several locations in the Lower Fraser Valley, British Columbia [81]. Sequences of specimens identified as A. franciscana (and the junior synonym A. citrana) fall into two BINs, BOLD:AAA9216 and BOLD:ABZ7843, separated by a distance of 1.69% (p-dist). The two BINs form a single cluster and do not appear to have any taxonomic-level significance. Landry et al. [82] performed a phylogenetic analysis of A. franciscana, A. citrana, A. franciscana insulana, and A. isolatissima using a larger segment of COI and determined that these taxa form a single monophyletic clade that should be considered a single species. They did, however, find a relatively high amount of variation (3.8% p-dist) in this group. Rubinoff and Powell [83] reanalyzed these same populations and determined that A. isolatissima was indeed a valid species separate from the A. franciscana complex.
There are several reports of a significant range (and associated host) expansion for A. citrana; these are summarized by Powell [57]. It is likely that much of these data is obscured by the uncertain taxonomy in this group, as A. citrana and A. franciscana were treated as separate species until Landry et al. [82]. For instance, Powell [57] states that A. citrana was not found in the San Francisco Bay area before 1910, yet this is the type locality in which A. franciscana which was collected by Walsingham in 1871. It does appear that the range of A. franciscana to the north and its associated "pest status" did increase at some point, with reports of A. citrana in the fruit-growing regions of Oregon and Washington as early as 1930 [57]. Reports of this species in Europe and Florida are erroneous [57,84]. Sequences of C. albaniana fall into a single BIN: BOLD:AAB3030, with representatives from Ontario, Manitoba, British Columbia, and Finland. The specimens from Finland cluster separately from the Canadian specimens, but overall genetic variation is low. Hence, it appears that this is a single Holarctic species with some minor geographic variability. Sequences from specimens of A. oporana from Europe (Austria, Finland, Germany, Italy, The Netherlands, Norway), Asia (Japan, South Korea), and North America (Canada) are represented in BOLD. All sequences are placed in BIN BOLD:AAD6710, with little to no variation; the majority of specimens share identical barcode sequences. One sequence from Japan is labeled as A. audax, but this is likely a misidentification. Freeman [85] indicates that this species was collected from two different locations in Canada: Vancouver, British Columbia in 1937, and Font Hill, Ontario. Pohl et al. [46] were unable to locate these specimens for verification and suggest that if this species was found in Canada at one time, it did not establish. However, BOLD contains sequences from three specimens collected in Ontario from 2013-2017, indicating this species is still present in Canada. Archips rosana is represented in BOLD by specimens from Europe (Austria, Finland, Germany, Italy, and Norway) and North America (British Columbia and Washington). All sequences form a single BIN, BOLD:AAB9404, with several specimens labeled as A. grisea, which are assumed to be misidentifications. This species was first reported in North America, likely arriving in 1889, by Comstock and Slingerland [87] from Albany, New York. It was found in Canada for the first time in 1919, both in Victoria, British Columbia, and Nova Scotia [41]. It has since spread to other locations in the East, including Connecticut, New Brunswick, Nova Scotia, Ontario, Pennsylvania, Prince Edward Island, and Quebec [46,85,88], and other locations in the West, including Alaska, Alberta, Oregon, and Washington [46,88]. The majority of sequences of A. podana are from European specimens, which form a single BIN, BOLD:AAB5839, with little genetic variation. However, two specimens from Canada (British Columbia) share identical barcode sequences with several of the European specimens. Freeman [85] reported that A. podana was first recorded in North America from British Columbia in 1937; however, these records were apparently confused with those of A. oporana [89]. The first confirmed record of A. podana in North America is from the lower Fraser River Valley in British Columbia in 1988 [42]. LaGasa et al. [89] provided details of surveys for this species in Whatcom County, Washington in the early 2000s, representing the first records for the U.S. The report of A. podana from Ontario by Belton [42] is an error [46]. The only specimens of A. fuscocupreanus represented in BOLD are from Washington, which fall into a single BIN, BOLD:AAD6614, and form two clusters with little genetic variation. Archips fuscocupreanus was first discovered in North America in Connecticut, where it has been present since at least 1982 [90]. Maier [91] determined that this species was widespread in the northeastern U.S. (Connecticut, Massachusetts, New Jersey, New York, and Rhode Island). He also discovered two specimens in the collection of the USNM from Washington collected in 1995 [91]. Sequences from specimens of C. pronubana are resolved into two separate BINs: BOLD:AAD3477 contains specimens from Europe (France, Germany, The Netherlands, Norway, United Kingdom) and BOLD:AAL5782 contains specimens from Spain. These two BINs are separated by >3.0% (p-dist). There are no phenotypical differences in the specimens that would provide evidence that two species are involved, although it is possible that specimens from Spain represent a cryptic species. No specimens from North America are represented in BOLD, although some have been sequenced in other facilities.

Archips fuscocupreanus Walsingham, 1900 Previous status: Introduced from Eurasia
Cacoecimorpha pronubana was found in Oregon in 1964 [92]. It has since spread to Washington, with the first record in 1974; surveys in the late 1990s revealed that it was widespread in western Washington [93]. In 2011, C. pronubana larvae were found in California feeding on Daphne (Thymelaeaceae), and in 2013 an infestation of C. pronubana was found in the Denver Zoo in Denver, Colorado [94]. Sequences of D. histrionana fall into two BINs: BOLD:AAC2478 includes specimens from Europe (Austria, Denmark, Italy) and Canada; and BOLD:ACF5563 contains specimens from Europe (Austria, Finland, Germany, Norway). The two BINs are separated by only 1% (p-dist), and both cluster together, suggesting that they are the same species. DeWaard et al. [95] first discovered this species in North America while conducting a DNA barcoding study of moth species found in Stanley Park, Vancouver, Canada. Sequences of C. spectrana are placed in a single BIN in BOLD (BOLD:AAC1795), with specimens from Europe (Finland, Germany, Italy, The Netherlands, Norway, United Kingdom) and Canada (British Columbia, Quebec). The earliest record in North America is a single specimen collected in British Columbia in 1950 [96]. This species was rediscovered feeding on raspberry, currant, spruce, and cedar in the early 1990s in British Columbia, and the first U.S. record was reported from Washington in 1997 [97]. Razowski [98] records the Palearctic distribution for this species as Central, Northern, and East Europe. This species is not represented in BOLD. Clepsis danilevskyi is present in Alaska and the Yukon [46] in the Nearctic, and in the Altai and Polar Ural Mountains in the Palearctic [50]. Clepsis firthana, described by Mutuura [99] from Canada, is considered a junior synonym of C. danilevskyi. Specimens of C. consimilana are placed in a single BIN (BOLD:AAC4212) in BOLD with representatives from Europe (Austria, Denmark, Germany, Italy, United Kingdom) and North America (British Columbia, Washington). Zlatkov and Huemer [100] revised a group of European Clepsis, including this species, and determined the following regarding BOLD:AAC4212: "The intraspecific average of the barcode region is 0.34%, the maximum distance 1.08% (p-dist) (n = 29). The minimum distance to the nearest neighbour, Clepsis eatoniana, is 2.25%."

Clepsis consimilana (Hübner, 1822) Previous status: Introduced from Eurasia
Several other single specimens labeled as C. consimilana in other BINs are likely misidentified. This species was first reported from North America by Klots [36] (as C. unifasciana), who identified two specimens from Long Island, New York collected in 1939. Powell [101] reported it from Oregon, and Dang et al. [96] reported the first records from British Columbia. The distribution of C. consimilana in the Palearctic includes Europe, Asia Minor, Syria, European Russia, and West Africa to Lebanon; it had been reported as introduced to Madagascar [100], but that population may represent a different species.  [50]. In North America, this species is recorded from Labrador, across Canada to Alaska [46], and from a few unverified records in other U.S. states. Sequences of C. illustrana in BOLD form two separate BINs: BOLD:AAF4488 contains three specimens from Finland, and BOLD:ACL6434 contains three specimens from Canada (Manitoba, Ontario, Yukon Territory). The two BINs cluster together with less than 2% (p-dist) overall variation; hence, the two BINs could represent closely related species or variation between two isolated populations of the same species: one in the Palaearctic and one in the Nearctic. Ferris et al. [102] first recorded this species from North America. We treat it here as tentatively Holarctic pending the sequencing and examination of additional specimens. Nearly 1600 sequences of E. postvittana are present in BOLD, and all are assigned to a single BIN (BOLD:AAB0679). This species is native to Australia and has been introduced to Tasmania, New Zealand, Hawaii, the United Kingdom, and the Azores, with isolated records from other countries in Europe, including The Netherlands. This species was first detected in California in 2006 [12], and it has since become established and common along most of the coastal counties from Mendocino south to San Diego. Genetic studies have determined that there were likely two separate introductions of this species into California, with Australia being the most likely origin [103]. The only specimens of S. rubicundana in BOLD are from Europe (Finland and Norway) and are placed in a single BIN (BOLD:AAF4352). Powell [71] synonymized S. hudsoniana Freeman, 1940, from Manitoba with the Palearctic S. rubicundana. Powell and Brown [104] followed this treatment and recorded additional specimens from Alaska and Nunavut. Pohl et al. [46] added the Northwest Territories and Ontario to the distribution for this species. In the absence of DNA evidence to the contrary, we assume that this is a Holarctic species. Specimens identified as P. stultana in BOLD are placed in two BINs that are separated by >3% (p-dist): BOLD:AAB6120 contains specimens from California and Florida, and BOLD:AAC5877 contains specimens collected near San Diego, California. It is possible that the San Diego population represents a cryptic species.

Ditula angustiorana (Haworth, [1811]) Previous status: Introduced from Europe
Powell and Brown [104] provide a detailed account for this species. A native of Mexico and possibly the southwestern U.S., P. stultana was introduced into California sometime around 1898. It greatly expanded its geographic range northwards in the 1960s. At the same time, it expanded its host range onto many nonnative plants [105], including important crops, and has now become an important pest of grapes and greenhouse plants [94]. It was reported from Florida in the 1940s, where it is apparently established [106]. Platynota stultana has been recorded from California, Arizona, Hawaii, Texas, Florida, Mexico, and there are sporadic records from other U.S. states, although it is probably not established elsewhere. This species was reported on peppers in Spain in 2009, and Groenen and Baixeras [107] documented its presence in that country as early as 2005. Sequences of E. gentianaeana are placed in a single BIN, BOLD:ACE6355, with specimens from Austria, Finland, Germany, and the United Kingdom. This species has been proposed as a biocontrol agent for teasel [109]. Beebe [110] reported that it was present in Michigan; however, Miller [108] determined that the specimens to which Beebe referred were misidentified E. hebesana. Sequences of B. furfurana are placed in three BINs in BOLD: BOLD:AAA7927 contains specimens from Canada and the U.S.; BOLD:ACE7672 contains specimens from Washington; and BOLD:ABZ2466 contains specimens from Finland, Norway, and the United Kingdom. Although it is possible that these different BINs represent different species, all three are resolved in a single cluster, and there is no further evidence that more than one taxon is represented in the samples. This is a common and widespread species. Identification of Apotomis to species is difficult, even with genitalic dissection [116]. As such, most of the identifications in BOLD require confirmation before any conclusions can be made based on sequence clustering, BIN assignment, etc. Some BINs, such as BOLD:ABZ6958, contain approximately 250 specimens identified as 10+ different species of Apotomis. It is also possible that DNA barcodes cannot convincingly separate the species in this genus. Specimens identified as A. infida in BOLD are placed into two BINs: BOLD:ACF3687 contains specimens identified as A. infida and A. moestana from Europe and Yukon Territory, and BOLD:AAA4846 contains specimens from Canada identified as A. infida and a variety of other Apotomis species. There is currently insufficient data in BOLD to make any definitive conclusions regarding this species. See the previous taxon for more details. Sequences of C. hartigiana in BOLD fall into two BINs: BOLD:AAE3063 contains specimens from Austria, Germany, and New York), and BOLD:ACF2297 contains specimens from Finland and Norway. Both BINs form a single cluster and are separated by 1.65% (p-dist). We assume that these BINs represent geographical variation and not separate taxa.

Cymolomia hartigiana (Ratzeburg, 1840) Previous status: Introduced from Europe
McGuinness and Brown [117] reported There is considerable confusion regarding the identity of specimens in these BINs, which are sufficiently separated to assume that they all represent distinct taxa. Orthotaenia undulana was described from Austria, and thus, it is likely that BOLD:AAB4021 represents this species. This BIN consists of two main clusters: one with specimens from Alaska, The Netherlands, and Italy, and the other with specimens from across Europe. There is no sequence variation in the first cluster, and minor variation in the second; i.e., the overall variation in the BIN is only 0.62% (p-dist). It is possible that identical sequences in Europe and Alaska could indicate that O. undulana is introduced; however, without any direct evidence of such, and with low overall sequence variation, the simplest explanation is that this BIN represents a single Holarctic taxon.
The majority of specimens from Canada and the U.S. that are identified as O. undulana in BOLD fall into a separate BIN, BOLD:AAA8540. We noticed subtle differences in both the male and female genitalia between specimens traditionally identified as O. undulana from Europe and North America. As such, it is possible that BOLD:AAA8540 represents a cryptic species present only in North America. If so, there are two current junior synonyms of O. undulana described from North America that could apply. The oldest is Sericoris campestrana Zeller, 1875, described from Maine or Massachusetts; the other species is Sericoris dilutifuscana Walsingham, 1879, described from southern Oregon.
The third BIN, BOLD:AAB4022, contains specimens that are clearly not O. undulana based on photos of the wing pattern and genitalic dissections. Specimens in this BIN were tentatively identified as the next taxon, Syricoris lacunana, based on dissections of specimens from Washington and Oregon. However, Steve Nanz, Editor in Chief of the Moth Photographers Group, has brought to our attention that the male genitalia for specimens in BOLD:AAB4022 are more similar to those of Olethreutes deprecatoria Heinrich, 1926, than to S. lacunana. This BIN is also well-separated from BOLD:AAC3531 (next taxon), which we believe to represent the true S. lacunana. In addition, given the similarity in genitalia of O. deprecatoria to S. lacunana, it is likely that O. deprecatoria would be more properly placed in Syricoris. Syricoris lacunana has not been previously recorded from North America. DNA barcodes indicate that this species is present in both eastern and western Canada. Based on its northern distribution in North America, we believe it is likely Holarctic instead of introduced. This species is similar in wing pattern to other taxa in North America (Olethreutes, Celypha, Orthotaenia), so it is not surprising that it could have remained undetected until now. NOTE: Heinrich [25] attempted to resolve the taxonomy of the North American species of "Olethreutes" by synonymizing Argyroploce, Phiaris, Celypha, Orthotaenia, Selenodes, and Mixodia with Olethreutes, and recognizing Exartema as a distinct, closely related genus. Exartema was subsequently synonymized with Olethreutes, resulting in essentially a single large genus for all North American species formerly included in this array of genera [71]. In Europe, these proposed changes were either short-lived or simply ignored, and the European definition of Olethreutes includes only the type (O. arcuella) and two additional species. Although Brown [26] made some progress toward resolving discrepancies in the generic assignments between the North American and European faunas: nearly all of the North American species remaining in Olethreutes would be assigned to different genera in European treatments, and all of the species listed below under Olethreutes are currently assigned to either Phiaris or Argyroploce in the most recent European checklist [118]. We generally agree with the European treatment, but hesitate to make extensive changes in generic assignment to species that occur in North America until there is a comprehensive revision of the group that includes all of these taxa. Nedoshivina [119] provides the most recent revision of generic concepts in this group, but her work is not particularly relevant to the North American fauna. Miller [120] synonymized O. metallicana with murina Packard, 1866 (described from Canada) and major Walsingham, 1895 (described from Colorado). He did note a slight difference in the female genitalia, but did not have a large enough sample size to determine if this was significant. Based on the DNA data, it appears that the European and North American populations are potentially different species. However, Nedoshivina [119] includes many records of this species from the Russian Far East, indicating that it may indeed be Holarctic. Additional study is necessary to resolve the status of this species. Miller and Jalava [121] determined that records of O. obsoletana from northeast Siberia were actually misidentified O. nordeggana. They state that the two species are very similar but can be separated by differences in the structure and size of the male cornuti. It is not known if the specimens of Phiaris obsoletana in BOLD that cluster with O. nordeggana are also misidentified, if these are actually the same taxa, or if DNA barcodes are unable to separate these species. In either case, it seems that O. nordeggana is likely Holarctic. In Europe, this taxon is currently treated under Phiaris [118]. inquietana; and BOLD:ACF5701 contains specimens from Europe and Yukon Territory identified as P. schulziana, P. palustrana, P. septentrionana, P. turfosana, and Phiaris sp. As stated above, it is difficult to know if many of these specimens are simply misidentified or if they cannot be separated by DNA barcodes. Therefore, the DNA data is currently of limited use for these taxa.

Olethreutes heinrichana (McDunnough, 1927) Previous status: Holarctic
Olethreutes schulziana was listed as Holarctic by Lafontaine and Wood [122]. It is not known when it was first discovered in North America, but Pohl et al. [46] list records from most of Canada and Alaska.  [111] formally synonymized Mixodia ? intermistana Clemens, 1865, described from Labrador, with O. turfosana, resulting in a Holarctic taxon. This species is currently recorded from Alaska, Maine, and much of Canada. Nedoshivina [119] records this species from much of Russia, including the Far East. In Europe, this taxon is currently treated under Phiaris [118].  [25] lists the type locality of this species as "Arctic American," there is nothing in the original description to indicate the origin of the types ("The box contained two specimens of this small Tortrix, which resembles a little the T. hybridana of Hübner, pl. 38, fig, 238."), and the prior species account of bentleyana describes specimens collected in England.

Olethreutes septentrionana (Curtis, 1835) Previous status: Holarctic
Regardless, there are two junior synonyms of septentrionana that are described from the Nearctic, and Jalava and Miller [111] synonymized the Palearctic schaefferana with septentrionana, resulting in a Holarctic taxon. In North America, it is found in Alaska, Maine, and parts of Canada. It is also present in the Russian Far East [119]. As with the previous three species, it is difficult to determine the distribution or even taxonomic status of this species. Heinrich [25] included several junior synonyms under inquietana that have since been placed under turfosana. Currently, O. inquietana is the senior synonym of hepialana (described from Mongolia), hoyoshi (described from Greenland), and retortimacula (described from Munku-Sardyk on the border of Mongolia and Russia). In North America, this species is found in Alaska, Maine, and parts of Canada. We assume it is Holarctic pending further analysis of the junior synonyms. In Europe, this taxon is currently treated under Phiaris [118].

Olethreutes mengelana (Fernald, 1894) Previous status: Nearctic
Revised status: Holarctic Supported by DNA in BOLD: No (no specimens from Europe in BOLD) BINs: BOLD:AAB9942 This species was described from McCormick Bay, Greenland. Miller and Jalava [121] included it in their study of boreal Olethreutini, but concluded that it was restricted to the Nearctic. Aarvik [118] states that it is present in Novaya Zemlya, an archipelago off northern Russia in the Arctic Ocean. This would indicate that it is indeed a Holarctic taxon. In Europe, this species is listed under the combination Argyroploce mengelana [118]. There are only six specimens of this species represented in BOLD, all of which were collected from Nunavut, Canada. This species is currently unrepresented in BOLD. It was known only from Russia until Jalava and Miller [111] reported a specimen from the Ogilvie Mountains in the Yukon, Canada collected in 1985. The type locality is Chukotka in the Russian Far East. According to Nedoshivina [119], this species should likely be placed in Argyroploce. with representatives from Austria, Finland, Germany, Italy, and Norway. Jalava and Miller [111] documented the first North American records for this species with specimens collected at Schrader and Galbraith Lakes in Alaska. This species is widespread in Europe and occurs in far eastern Russia [119]. In Europe, this taxon is currently treated under Phiaris [118]. This species was first reported from North America by Landry et al. [32], with a specimen from Churchill, Manitoba collected in 2007. It has been suggested in the past that glaciana and bipunctana might be synonyms. Miller [120] examined both species and determined that they were not conspecific, and Landry et al. [32] provided a more detailed list of differences between the two taxa. (Möschler, 1860) Argyroploce glaciana is represented in BOLD by more than 300 specimens in a single BIN (BOLD:AAA6005), which is shared with specimens of Phiaris bipunctana. Although both species are very similar, previous studies [32,120] concluded that it is possible to differentiate them by wing pattern and genitalia, and the DNA data for both species form distinct and separate clusters. Specimens of P. glaciana in BOLD are from Alaska, Washington, and across much of Canada.

Phiaris glaciana
Lafontaine and Wood [122] listed P. glaciana as Holarctic, but it is possible they were referring to misidentified specimens of P. bipunctana instead, or even another species. We have not found any verified records of P. glaciana from Europe (including under other generic names), and this species is not included in the most recent European faunal list [118]. As such, P. glaciana is likely not Holarctic. Landry et al. [32] suggested that the similarity of glaciana to bipunctana meant that they should be placed in the same genus. We agree, based on both morphological and molecular data, and thus transfer glaciana to Phiaris. Only two specimens of P. siderana are present in BOLD: one from Norway and the other with no locality data. This species is widespread across Europe and parts of Asia, including Russia [119,123]. Sericoris chalybeana was described from California and is present in the Pacific Northwest and Montana [88]. This species is currently considered a junior synonym of P. siderana, and it has been treated as a subspecies in the past [71]. In Europe, this taxon is currently treated under Celypha [118]. There is a single BIN (BOLD:AAB9941) in BOLD representing A. aquilonana, with specimens from Canada (Manitoba, Northwest Territories, Nunavut, Yukon Territory), Greenland, and Finland. Jalava and Miller [111] reported this species (as Olethreutes aquilonanus) from the Ogilvie Mountains in the Yukon, which was the first North American record. They also synonymized this taxon with Olethreutes kononenkoi, extending the distribution of A. aquilonana into Russia. Nedoshivina [119] records this species from Chukotka in the Russian Far East. Specimens of A. externa are placed in a single BIN (BOLD:AAI3239) with representatives from Europe (Finland, Norway) and Canada (Alberta, Northwest Territories, Yukon Territory). Jalava and Miller [111] reported the presence of the junior synonym "Olethreutes dalecarlianus" in North America, from the Ogilvie Mountains in the Yukon and Wellington, British Columbia. This name was synonymized under the Palearctic Argyroploce externa by Nedoshivina [124], making the senior taxon Holarctic. In BOLD, sequences of H. atropunctana are placed in a single BIN (BOLD:ABZ7645) that includes specimens from Europe (Austria, Finland, Germany, Italy, The Netherlands, Norway, United Kingdom) and a single specimen from Alaska. Jalava and Miller [111] reported specimens of this species from the Ogilvie Mountains in the Yukon and the Dietrich River in Alaska, confirming that it is indeed Holarctic. Nedoshivina [119] lists many records of this species from Asia. Alipanah and Baixeras [125] reviewed the status of this genus and resurrected the combination Hedya atropunctana (which was previously placed in Metendothenia).

Hedya atropunctana (Zetterstedt, 1840) Previous status: Holarctic
Enarmonia formosana sequences are placed in a single BIN in BOLD (BOLD:AAC5227), which contains specimens from the U.S., Canada, and Europe. Enarmonia formosana is widely distributed from Western Europe and northern Africa to Asia Minor, Russia, and Siberia [123]. The first North American records are from British Columbia in 1989 [128]; it was subsequently found in western Washington in 1991 and in Oregon in 2000 [94]. The "cherry bark tortrix" is a common pest of numerous species of fruit trees [130]. Rhyacionia buoliana was first recorded in North America in 1913 in New York [41]. It was reported from British Columbia in 1927, and although this initial population was eradicated, it was reported again from Vancouver in 1938 [41]. The "European pine shoot moth" is one of the most common pests of conifers. It is widely distributed and variable both in wing pattern and genitalia, resulting in a long list of synonyms described from Europe [27]. Sequences of S. ocellana in BOLD are placed in two BINs: BOLD:ABZ4399, with specimens from Europe and North America, and BOLD:AAA6641, with specimens from Canada (British Columbia, Ontario, Quebec). It is not known if this second BIN represents a possible cryptic species or genetic variation.

Spilonota ocellana ([Denis and Schiffermüller], 1775) Previous status: Introduced from Eurasia
Spilonota ocellana was introduced to the Nearctic likely before 1840, with the first report from Massachusetts in 1841 [41,131]. It became widely distributed in the northeastern U.S. and Canada, and was possibly introduced separately to the Pacific Coast, where it was common in Vancouver by 1912 [41,88]. This is a common orchard pest that is present in all apple-growing regions of the northern hemisphere [94].  1% (p-dist).

Spilonota laricana (Heinemann, 1863) Previous status: Introduced from Europe
Historical records of this species in North America likely refer to S. ocellana [46]; thus it is difficult to determine when this species might have been introduced. It was reported from the northeastern U.S. by the Moth Photographers Group around 2010, and the DNA data in BOLD has confirmed that it is present and possibly widespread in eastern Canada. The second BIN (BOLD:AAA7739), with specimens from Finland, possibly represents a cryptic species in northern Europe. Sequences of E. hohenwartiana in BOLD are placed in a single BIN (BOLD:AAB4295), with all of the specimens from Europe. Ferris et al. [102] recorded it from Northwest Alaska, adding it to the list of Holarctic tortricids. Wright and Gilligan [132] were unable to confirm this record, and there is no DNA evidence in BOLD to support or refute this conclusion.

Pelochrista adamantana (Guenée, 1845) Previous status: Holarctic
Revised status: Nearctic Supported by DNA in BOLD: Yes BINs: BOLD:AAF2211 The three specimens of P. adamantana with sequence data in BOLD are from the U.S., and are placed in a single BIN (BOLD:AAF2211). This species has been listed as potentially Holarctic in some checklists because of the statement in the original description that it was described from "ex Lapponia (?)," which could refer to Lapland in Finland. However, it has been excluded from European checklists since Rebel [133] determined that "it does not seem to be an inhabitant of this territory [the Palearctic]." Heinrich [24] appended "North America" to the location of the type specimen in an attempt to clarify the issue. Wright and Gilligan [134] were unable to locate the type, but they found no evidence of this species occurring in Europe. In BOLD, two specimens labeled as P. medullana are placed in a single BIN (BOLD:AAE7175), along with specimens identified mostly as P. caecimaculana and a few other Pelochrista species from Europe. It is likely that the mixed species assignments to this BIN are the result of misidentifications. Pelochrista medullana was introduced into western North America for the biological control of Centaurea species (knapweed). This species was released in Idaho, Montana, Oregon, and British Columbia, but there is no evidence that it established in these locations [135,136]. Sequences of E. simploniana and its former junior synonym, E. arctica, are placed in five different BINs in BOLD: BOLD:ACO4599 contains specimens identified as E. arctica from the Ogilvie Mountains in the Yukon Territory, Canada; BOLD:ACO4600 contains specimens identified as E. arctica from the British Mountains in northwestern Yukon; BOLD:AAF2136 contains specimens identified as E. simploniana from Finland and Norway; BOLD:AAF2135 contains specimens labeled as E. simploniana from Finland; and BOLD:ACJ7207 contains specimens labeled as E. simploniana from Austria and Italy.

Pelochrista medullana (Staudinger, 1879) Previous status: Introduced from Europe
Miller [137] described E. arctica from specimens collected in Alaska. He later [138] determined that E. arctica was conspecific with E. simploniana, a Palearctic species recorded from the mountains of Central Europe, Scandinavia, western Russia, Siberia, and Mongolia [123]. DNA barcodes for this group segregate into several BINs that are separated geographically, with E. arctica in the Nearctic and E. simploniana in the Palearctic. In addition, we found what appear to be subtle but consistent morphological differences between E. arctica and E. simploniana that could be used to separate the two taxa (unpublished). Further research is necessary to determine why there are two separate BINs for E. arctica and three separate BINs for E. simploniana; this could indicate the possibility of one or more cryptic species in this group. Regardless, there is sufficient evidence to elevate E. arctica Miller, 1985, stat. rev., back to species status and restrict the distribution of E. simploniana to the Palearctic. There is little genetic variation in this BIN, and many specimens share nearly identical DNA barcodes. A single specimen from the United Kingdom is placed in a separate BIN by itself (BOLD:AAC1135), but the DNA sequence is short and likely accounts for placement of this specimen.
There has been a long history of attempting to determine if the European N. rosaecolana, N. trimaculana, or both species were introduced to North America. The earliest record is from Smith [139], who reported Eucosma suffusana (a junior synonym of N. trimaculana) from New Jersey. Various other publications refer to N. trimaculana or N. rosaecolana as being the introduced taxon. Miller et al. [140] resolved the issue by discovering a morphological character that could be used to consistently separate the two taxa. They determined that N. rosaecolana is the only species of the two present in North America, and that prior literature records referring to N. trimaculana were likely incorrect. Mutuura [141] documented the first record of N. cynosbatella in North America from specimens collected near Vancouver, British Columbia in 1978. Pohl et al. [46] list the distribution as Alberta, British Columbia, Newfoundland, and Ontario. Notocelia cynosbatella is very similar to N. illotana, which is native to North America. Although Mutuura [141] provides a list of characters that can be used to separate the two species, the differences appear subtle, and genitalic dissection may be necessary for a positive identification. It is possible that the BIN containing only specimens from Canada could represent N. illotana, or that DNA barcodes in BOLD are not sufficient to separate these two species. It is also possible that there are cryptic species in this group. The BIN BOLD:AAB1285 contains specimens identified as several different species of Gypsonoma, including G. parryana, G. nitidulana, G. fasciolana, G. salicicolana, and G. nebulosana. The sequences cluster into several groups, but it is not clear if many of the specimens in this BIN are misidentified or if these DNA data are not sufficient to separate out species. Pohl et al. [46] list this species from Alaska, northwestern Canada, Manitoba, and Quebec. It is also recorded from Russia, in Novaya Zemlya and the Russian Far East [123]. It is not known if many of these are misidentifications or if DNA barcodes are not able to separate species in this group.
Gypsonoma nitidulana was reported from the Yukon by Lafontaine and Wood [122]; however, Pohl et al. [46] could not locate any voucher material, and declared the record to be erroneous. There is currently no evidence in the DNA data or collections to suggest G. nitidulana is present in North America. Sequences of G. aceriana in BOLD are placed in a single BIN (BOLD:AAB0379) with specimens from Europe (Austria, Denmark, Germany, Italy, Norway, United Kingdom) and Canada. This species was first reported from North America in western Washington by Miller and LaGasa [142]. Several years later, Humble et al. [30] reported specimens of G. aceriana that were collected in southwestern British Columbia as early as 1980. Subsequent collecting by LaGasa and Humble et al. [30] confirmed that this species is established in the Pacific Northwest. Sequences of C. plebejana in BOLD are separated geographically into several BINs. BOLD:AAA7083 contains specimens only from Australia; BOLD:ABY9664 contains specimens only from Africa; BOLD:ACE9795 contains specimens from Europe and Egypt; and BOLD:AAA7084 contains specimens from California, Texas, and Costa Rica. There is seemingly no regional overlap in the BOLD data, suggesting that each of these BINs might be a separate taxon. Distance between BINs varies from 1.4% (p-dist) between BOLD:AAA7083 and BOLD:ACE9795 to 4.3% (p-dist) between BOLD:AAA7084 and the other three BINs.
Crocidosema plebejana was originally described from Europe, and an additional 13 names, described from various parts of the world, are currently considered synonyms, including several from the New World. Some other species described from Hawaii and South America are closely related [143,144]. Although C. plebejana is currently treated as a cosmopolitan species, it has been suggested that several of the synonyms are valid species [123]. Razowski et al. [145] resurrected from the list of synonyms C. synneurota Meyrick (described from the Galapagos) as a valid species and suggested other potential changes. The DNA data in BOLD suggest that at least populations on each continent might be separate taxa. Similarly, Lincango [146] in a multi-gene phylogenetic analysis found evidence of paraphyly in a complex of species around C. plebejana, and the specimens of C. plebejana from Europe did not group with those from South America. Moreover, there is extreme variation in both male and female genitalia. A more comprehensive molecular analysis is necessary to clarify the identity of the species involved. For now, we believe C. plebejana should be restricted to the Palearctic, and that none of the taxa involved in this complex are Holarctic in distribution. The sequences of C. lantana in BOLD are placed in a single BIN (BOLD:AAH5763), with specimens from Costa Rica, Jamaica, Australia, Madagascar, and Kenya.

Crocidosema lantana
This species was introduced to Hawaii from Mexico in 1902 to control Lantana [144]. It was later introduced to Australia from Hawaii around 1914, where it became common along the East Coast and prompted the description of several new names by Turner which were synonymized by Common [147]. Crocidosema lantana has been introduced to Kenya [148], and is also present in South Africa and apparently Madagascar based on DNA records. In North America, C. lantana is found in Florida. In BOLD, the nearly 700 sequences representing this species fall into a single BIN (BOLD:AAA9812), with representatives from Canada, Europe, Japan, Pakistan, and the U.S. There is no evidence of any cryptic taxa.

Rhopobota naevana (Hübner, [1817]) Previous status: Holarctic
This species is most likely Holarctic, being recorded from North America as early as 1863 and 1869 based on junior synonyms described from Hudson Bay, Canada and Massachusetts. Gillespie and Gillespie [41]  DNA data support a Holarctic distribution for this species. Pohl et al. [46] include the unpublished manuscript name jasperana Brown, 1980 [149], described from Jasper, Alberta, as a junior synonym of E. trigonella. Aarvik [118] treats E. indecorana and E. trigonella as separate species, and Karsholt et al. [48] follow this arrangement in their list of Lepidoptera from Greenland. DNA data in BOLD appear to support this separation, with sequences of E. indecorana clustering separately from those of E. trigonella.

Epinotia solandriana (Linnaeus, 1758) Previous status: Holarctic
Revised status: Holarctic? Supported by DNA in BOLD: No (sequence clustering is ambiguous) BINs: BOLD:AAA6716 Sequences of E. solandriana in BOLD are placed a single BIN (BOLD:AAA6716). However, sequences within this BIN are divided into two distinct clusters, with one representing North America (Alaska, California, Washington, Alberta, British Columbia, Manitoba, New Brunswick, Newfoundland, Quebec, Saskatchewan) and the other representing Europe (Finland, Germany, Italy, Norway). This clustering of sequences could suggest that the European and North American populations represent different species, but more study is needed to determine the genetic variability of this species.
Epinotia solandriana is currently considered to be Holarctic [28,149]. Martineau [150] suggested that this species was introduced from Europe to British Columbia in 1909. There is no other direct evidence of such; however, the clustering of sequences by continent in BOLD could also represent a genetic bottle-neck resulting from a potential introduction. In BOLD, E. nisella is placed in three BINs: BOLD:ABZ1141 containing specimens from Austria and Germany; BOLD:AAA7530 containing specimens from Europe and Alaska; and BOLD:AAA7528 containing specimens rom Canada and Alaska. Mutanen et al. [31] provides details regarding DNA barcodes for this group.
Mutanen et al. [31] resolved the taxonomy of this and the next species (E. cinereana). Epinotia nisella has a long list of synonyms, owing to its variable forewing pattern. This species is currently distributed throughout Europe and Russia to China and Japan, and has even been recorded from Iceland [31]. In North America, it is found in the northern U.S. and across Canada north to the Yukon and Alaska. The specimens of E. cinereana represented in BOLD are placed in three BINs: BOLD:AAA7532 containing specimens labeled as E. cinereana from Europe; BOLD:AAA7529 containing specimens labeled as E. criddleana from Canada; and BOLD:ABX4876 containing specimens labeled as E. cinereana from Europe and E. criddleana from Canada. Mutanen et al. [31] provide details regarding DNA barcodes for this group.

Epinotia cinereana (Haworth, [1811]) Previous status: Holarctic
Mutanen et al. [31] resolved the taxonomy of this species, which was previously synonymized under E. nisella. They determined (based on morphology) that the North American E. criddleana is a junior synonym of E. cinereana, resulting in a Holarctic distribution. Epinotia cinereana is widely distributed across Europe, with scattered records through Russia to Japan and China [31]. In North America, it is found from Quebec and New Brunswick west to British Columbia and south to Iowa and Colorado [31]. Epinotia columbia is represented in BOLD with two BINs: BOLD:AAF0407 contains specimens from California and British Columbia, and BOLD:AAE1754 contains specimens from Canada (Alberta, Northwest Territories, Ontario, Quebec, Yukon Territory). Both are separated from the BIN containing E. crenana (former senior synonym) from Europe (BOLD:AAE1756). It is not known which BIN represents E. columbia, although it was described from British Columbia, and the historic distribution has been limited to western North America. Thus, it is likely that BOLD:AAE1754 represents an undescribed cryptic species.
Heinrich [24] synonymized columbia with E. crenana based on similarities in genitalia. In his unpublished Master's thesis, R. L. Brown [149] elevated E. columbia back to species status; this change was formalized by J. Brown [26] in the tortricid world catalogue. Assuming many of the records from Canada represent a potential cryptic species (see above), the current distribution of E. columbia includes much of Canada west to British Columbia, Arizona, Nevada, Utah, and along the Pacific Coast to southern California [88]. There is no evidence that this species occurs in Europe, and records of such prior to 2005 are likely due to the previous synonymy with E. crenana. Sequences of E. nanana fall into two BINs in BOLD. BOLD:AAA8628 contains specimens from Canada (New Brunswick, Ontario, Quebec, Prince Edward Island), the U.S. (Connecticut), and Europe (Finland, Norway). BOLD:ACF5528 contains specimens from other locations in Europe (Austria, Germany, Italy). Both BINs cluster together and are separated by only 1.09% (p-dist). As such, it is likely they represent a single taxon with some geographic structuring.
Epinotia nanana was originally described from Germany, and in the Palearctic, it is widely distributed across Europe, ranging into Siberia and Mongolia [123]. The first records in North America are based on species described by Kearfott that were later determined to be junior synonyms of E. nanana by Heinrich [24]. Eucosma domonana (Kearfott, 1907) was described from Framingham, Massachusetts, and Epinotia piceafoliana (Kearfott, 1908) was described from Essex County Park, New Jersey. The first record of E. nanana in western North America is from a single specimen collected in British Columbia at some point prior to 1965 [41]. This species is currently distributed from West Virginia and Ohio northeast to Prince Edward Island. It is not known if E. nanana established in British Columbia.
from Canada is more distant (2.2% p-dist) from the other two BINs, which are only separated by 1.2% (p-dist). It is likely that one or both of the BINs containing specimens from Europe represent the true E. cruciana.
This species was described from Sweden and has at least 15 junior synonyms, several of which are from North America [27]. Epinotia cruciana ssp. alaskae was described from Alaska; cockleana was described from western Canada; direptana was described from Hudson Bay, Canada; E. cruciana ssp. lepida was described from New Hampshire; and vilisana was described from Hudson Bay, Canada. The known distribution of E. seorsa includes western Canada (Alberta, British Columbia) and California [153]. There are no records of this species from Europe and no sequences in BOLD. Pohl et al. [28] included the unpublished manuscript name blanchardi [149] as a junior synonym, which extends the range of this species into Texas. However, we suspect this is an error because specimens of the two taxa in the USNM do not appear to be similar. Sequences of D. vancouverana are placed in a single BIN, BOLD:AAC1223, with specimens from Europe (Austria, Finland, Germany, Italy, Norway, United Kingdom) and Canada (Alberta, British Columbia). As the name suggests, D. vancouverana was described from Vancouver Island, British Columbia. Miller [156] recognized that D. vancouverana was the same species as the European D. gueneeana (Obraztsov, 1953) and synonymized the two names.

Epinotia salicicolana
The taxonomic history of this species is somewhat confusing. Denis and Schiffermüller [157] described Tortrix politana from Austria (now a junior synonym of Lathronympha strigana). Guenee [158] referred to an undescribed species of Dichrorampha (now vancouverana) in Europe as politana; however, this was clearly a misidentification. Obraztsov [159] assigned the name gueneeana as a replacement name for Guenee's misidentified politana. Because Guenee's reference to politana was simply a misidentification, it is likely that Obraztsov's gueneeana is invalid, because this was assigned as a replacement name and was not a new species description. Regardless, the species previously referred to as D. gueneeana in Europe can now be correctly referred to as the Holarctic D. vancouverana. Dichrorampha sedatana can be problematic to separate from similar species such as D. aeratana, D. acuminatana, D. plumbana, and D. dana based only on wing pattern. A genitalic dissection is necessary to confirm identity in this group, and even then, identification is difficult due to variation. The majority of Dichrorampha in BOLD are not dissected, and thus, species identifications are sometimes questionable, especially when multiple species are assigned to the same BIN. Sequences of D. sedatana in BOLD are placed in two BINs. BOLD:AAA8636 contains specimens labeled as D. sedatana from Canada and specimens labeled as D. acuminatana from Europe. BOLD:AAA8637 contains specimens labeled as D. sedatana from Europe, specimens labeled as D. aeratana from Europe and Canada, and specimens labeled as D. tarmanni from Italy. Relatively low genetic diversity in both BINs suggests that multiple names in each BIN are a result of misidentifications, or that multiple species within this group share nearly identical DNA barcodes. It is not possible to determine the status of these taxa without extensive examination of the specimens from this group represented in BOLD.
Dichrorampha sedatana is currently treated as Holarctic [123]. It has been noted in the past that D. sedatana specimens identified in Europe seem to be different from those identified in North America (K. Larsen, pers. comm.). However, the BOLD data are currently not sufficient to support this conclusion because of apparent issues with species identifications. Pohl et al. [46] noted that the use of D. plumbana (Scopoli, 1763) for North American populations is erroneous.

Dichrorampha odorata Brown and Zachariades, 2007
Previous status: Introduced? Revised status: Probably native to southern Florida/Caribbean Supported by DNA in BOLD: No (this species not present in BOLD) BINs: N/A This species was described from Jamaica in 2007, but was found in Florida in 2014 feeding on weeds. There is no evidence it is introduced, and thus, it is probably native to the region and simply not detected in Florida until recently. This species is not represented in BOLD. distributed in the eastern U.S., and there are records from Ontario [46,165]. Cranshaw et al. [166] found G. delineana to be present in hemp fields in eastern Colorado in 2017-2018, and it is likely that this species will continue to spread with expanded industrial hemp production. Sequences of Grapholita aureolana fall into a single BIN in BOLD (BOLD:AAE7252), with specimens from Europe (Austria, Finland) and Canada. Pohl et al. [46] were the first to report this species from North America based on specimens in BOLD that were confirmed with genitalic dissections. Cydia coniferana is represented in BOLD by sequences that fall into two BINs: BOLD:AAC6167 contains specimens from Europe (Austria, Finland, Germany, Norway, United Kingdom) and Canada (Ontario), and BOLD:AAC6168 contains specimens from the western U.S. (Washington, New Mexico).
This species was first reported from North America by Schaffner [167] for several adults reared from the bark of red pine in New York. It was assumed that this population did not establish. LaGasa and Passoa [168] reported that this same species was also collected in Thurston County, Washington in 2000. Additional surveys in 2005 found specimens throughout most of western Washington, from the border of British Columbia to Oregon.
Based on the DNA sequence data, it seems that a population of C. coniferana did establish in eastern North America, with one specimen collected in 2015 from Peterborough in Central Ontario represented in BOLD. It is likely that the populations in the Pacific Northwest and elsewhere in western North America (New Mexico) represent a cryptic species. There are also records of this species from the Russian Far East [123]; these and other records from the region (e.g., Siberia) should be checked carefully to ensure they are indeed C. coniferana. Sequences of C. nigricana in BOLD are placed into two separate BINs: BOLD:AAA7614 contains specimens from Europe (Austria, Finland, Germany, The Netherlands, Norway, United Kingdom) and Canada (Alberta, Manitoba, Ontario, Saskatchewan), and this BIN likely represents the true C. nigricana, which was described from Great Britain; the second BIN (BOLD:AAE7682) contains specimens from only North America (Oklahoma, Ontario, Quebec), and represents either a group of misidentified specimens or a potential cryptic species. There are two junior synonyms of C. nigricana that were described from the U.S. (dandana Kearfott, 1907 and novimundi Heinrich, 1920), and it is possible that one or both of these names could be applied to specimens in BOLD:AAE7682. Cydia nigricana was introduced from Europe, and first found in North America in eastern Canada in 1893 and in British Columbia in 1933 [46]. Sequences of C. pomonella in BOLD are placed into a single BIN (BOLD:AAA3532) that consists of specimens collected throughout the world. There is no evidence of any cryptic species. The earliest record of C. pomonella in North America is a report of wormy apples and pears in the vicinity of Boston in 1819 [164]. The "codling moth" is one of the most widespread and important pest of apple, pear, and walnuts in the world. The name for the "Mexican jumping bean" moth has traditionally been cited as Cydia deshaisiana Lucas, 1858 in most modern publications. Although this name was published by Lucas in November, 1858 [169], his account (in French) is not accompanied by anything that can be construed as a description. As such, the name deshaisiana is a nomen nudum. The behavior of this insect ("jumping seeds") had actually been described a year earlier by Westwood in 1857 [170], although he did not provide a formal description or assign a name in that paper. On June 7, 1858, at the meeting of the London Entomological Society, Westwood read a paper describing the same species (in Latin) under the name Carpocapsa saltitans [171,172].

Cydia nigricana (Fabricius, 1794) Previous status: Introduced from Europe
Determining the exact date this description was published is difficult. The Proceedings of the Entomological Society of London were published on an irregular basis, most often quarterly. Wheeler [173] listed dates of publication by volume, part, and page number. It is possible that the June, 1858 proceedings would have been published in the next volume, which was delivered to the Society on July 5, 1858. However, because the exact page number is unknown, it is also not known if Westwood's description was included in the July volume or if there was a delay in the proceedings appearing in print. We do know that Westwood's account appeared in print on page 27 of the summary volume, Proceedings of the Entomological Society of London 1858-1859 [172], which appears to have been published as part of The Transactions of the Entomological Society of London, New Series, Volume V, 1858-1861, because the index that lists Westwood's account includes the years 1858 through 1861. However, the 1858 issue of The Zoologist [171] also included a transcription of Westwood's account. The exact publication date of Volume 16 of The Zoologist is not known; however, none of the dates in the volume are past 1858, and thus, it is assumed that it was printed in late 1858 or early 1859. As such, The Zoologist [171] is the earliest publication for which we can confirm a date for Westwood's description of C. saltitans, unless we simply use the June 7, 1858 date when the paper was first read and assume it was published in the next issue of the Proceedings in July. Regardless, the correct name for the "Mexican jumping bean" moth is Cydia saltitans (Westwood, 1858), stat. rev.
The only specimens in BOLD are from Costa Rica in (BOLD:AAA0992), and there is no evidence of this species occurring north of Mexico. "Mexican jumping beans" are routinely imported into the U.S. and adult moths eventually emerge from the seeds. However, there are no records of this species being found in the wild in the U.S., and the distribution of C. saltitans appears to be restricted the northern states of Sinoloa and Sonora in Mexico. and other data indicate that a maximum of 58 of these are actually Holarctic (separating Holarctic from introduced is not possible in some cases; thus, the actual number is likely less than 58). Hence, prior assumptions regarding the Holarctic distribution of Tortricidae may be overestimated by more than 20%, which is not surprising, given other recent studies, e.g., those of Landry et al. [32] and Gilligan et al. [33]. The primary reason for this discrepancy appears to be the presence of cryptic species in the Nearctic that were incorrectly identified as Palearctic taxa by early taxonomists.  appear to be increasing, with economically important species like the European grapevine moth and light brown apple moth triggering significant quarantine actions, both domestically and abroad. As such, it is important to continue surveying for immigrant Tortricidae and to continue research on these taxa to determine whether species in question are introduced or Holarctic. Funding: This research received no external funding.