ArticlePDF Available

Different continents, same species? Resolving the taxonomy of some Holarctic Ancylis Hübner (Lepidoptera: Tortricidae)

Authors:

Abstract and Figures

Several species of Ancylis related to A. unguicella (Linnaeus) and A. geminana (Donovan) have been presumed by previous authors to be Holarctic. However, difficulty in identifying genitalic characters to define and separate these taxa has brought into question their true distribution and led to inconsistencies in their taxonomic treatment in Europe and North America. Here we use a combination of DNA barcode sequence data and morphology to resolve these taxonomic differences, determine the actual geographic range of these taxa, and describe three new species. In the A. unguicella group, only A. unguicella and A. uncella (Denis & Schiffermüller) are Holarctic in distribution. In the A. geminana group, none of the taxa are Holarctic in their distribution. Three species are described as new: A. christiandiana Huemer and Wiesmair, sp.n. (Austria, Germany); A. oregonensis Gilligan and Huemer, sp.n. (USA: Oregon); and A. saliana Gilligan and Huemer, sp.n. (USA: Florida). In addition, Ancylis carbonana Heinrich, syn.n., is synonymized with A. uncella; A. cuspidana (Treitschke), syn.rev., is synonymized with A. diminutana (Haworth); and A. diminuatana Kearfott, stat.rev., and A. subarcuana (Douglas), stat.rev., are raised from synonymy.
Content may be subject to copyright.
Accepted by J.W. Brown: 5 Sept. 2016; published: 26 Oct. 2016
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4178 (3): 347
370
http://www.mapress.com/j/zt/
Article
347
http://doi.org/10.11646/zootaxa.4178.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:70509AD2-640A-497C-83EA-85B3EBEA35A0
Different continents, same species? Resolving the taxonomy of some Holarctic
Ancylis Hübner (Lepidoptera: Tortricidae)
TODD GILLIGAN
1
, PETER HUEMER
2
& BENJAMIN WIESMAIR
2
1
Identification Technology Program, USDA-APHIS-PPQ-S&T, 2301 Research Blvd., Suite 108, Fort Collins, Colorado 80526, USA.
E-mail: todd.m.gilligan@aphis.usda.gov
2
Tiroler Landesmuseen Betriebgsges.m.b.H., Naturwissenschaftliche Sammlungen, Feldstr. 11a, A-6020
Innsbruck, Austria. E-mail: p.huemer@tiroler-landesmuseen.at; b.wiesmair@tiroler-landesmuseen.at
Abstract
Several species of Ancylis related to A. unguicella (Linnaeus) and A. geminana (Donovan) have been presumed by previ-
ous authors to be Holarctic. However, difficulty in identifying genitalic characters to define and separate these taxa has
brought into question their true distribution and led to inconsistencies in their taxonomic treatment in Europe and North
America. Here we use a combination of DNA barcode sequence data and morphology to resolve these taxonomic differ-
ences, determine the actual geographic range of these taxa, and describe three new species. In the A. unguicella group,
only A. unguicella and A. uncella (Denis & Schiffermüller) are Holarctic in distribution. In the A. geminana group, none
of the taxa are Holarctic in their distribution. Three species are described as new: A. christiandiana Huemer and Wiesmair,
sp.n. (Austria, Germany); A. oregonensis Gilligan and Huemer, sp.n. (USA: Oregon); and A. saliana Gilligan and Hue-
mer, sp.n. (USA: Florida). In addition, Ancylis carbonana Heinrich, syn.n., is synonymized with A. uncella; A. cuspidana
(Treitschke), syn.rev., is synonymized with A. diminutana (Haworth); and A. diminuatana Kearfott, stat.rev., and A. sub-
arcuana (Douglas), stat.rev., are raised from synonymy.
Key words: christiandiana, DNA barcoding, Enarmoniini, oregonensis, saliana
Introduction
Tortricid taxonomy has long been plagued by inconsistencies in classification between the Nearctic and Palearctic
faunas. Early workers in North America encountering potential new species had to determine if these taxa were
undescribed or Holarctic in distribution, thus conspecific with species already described from Europe. This task
was especially difficult prior to the use of genitalia in tortricid taxonomy, and decisions regarding conspecificity
were based on wing patterns and life histories, although the latter was often unknown. The widespread use of
genitalic characters to separate species began in the 1920’s, led by Pierce and Metcalfe (1922) in Europe and
Heinrich (1923, 1926) in North America. Examination of the genitalia allowed these researchers to make
tremendous advances in tortricid taxonomy, and reliance on the genitalia to define species and higher taxonomic
groups increased to where it has become one of the most important characters sets in the classification of the
Tortricidae (Horak 1999). Specifically, it has been the male genitalia that are believed to be of diagnostic value at
the species level (Klots 1970), and taxa are often considered to be conspecific based on similarity in the male
genitalia (Mutanen et al. 2012). Care should be taken, however, when relying on a single character set to infer
taxonomy. Traditionally, reliance on the male genitalia as a species-specific character set was justified by the lock-
and-key mechanism of genital evolution (e.g., Mikkola 1992), although sexual selection, where variation in the
male genitalia is directly responsible for fertilization success (Eberhard 1985, Arnqvist 1997, Hosken and Stockley
2004) suggests that intraspecific variation in the genitalia is expected. Intraspecific variation in tortricid genitalia
has been well documented in some groups (e.g., Mutanen et al. 2007, Gilligan and Wenzel 2008, Wright and
Gilligan 2015).
The advent of molecular systematics, including DNA barcoding (Hebert et al. 2003), has led to the
GILLIGAN ET AL.
348
·
Zootaxa 4178 (3) © 2016 Magnolia Press
development of new character sets with which to test species boundaries. Patterns in DNA sequence data can be
used to determine “important” morphological characters for defining a taxon that would otherwise be
undecipherable because of intraspecific variation or an apparent lack of interspecific variation (Mutanen et al.
2012, Brown et al. 2014, Gilligan et al. 2014b). This approach can also be used to determine if morphologically
similar populations in North American and Europe are conspecific or separate species. Several recent studies (e.g.,
Humble et al. 2009, Mutanen et al. 2012, Landry et al. 2013) have used DNA barcoding to resolve issues
surrounding Holarctic (or presumed Holarctic) Tortricidae. One group with presumed Holarctic species that are
difficult to define using traditional male genitalic characters is the genus Ancylis Hübner.
Ancylis is the largest genus in the Enarmoniini with more than 140 described species (Brown 2005, Gilligan et
al. 2014a). Members of the genus are found worldwide, with the majority of species in the Holarctic, Oriental, and
Australian regions (Horak 2006). The genus was divided between Anchylopera Stephens and Ancylis until the late
1970’s based on an atrophied uncus in Anchylopera, but this character was found too variable to be diagnostic
(Razowski 1989, 2003). The genus is currently defined by the hollow bladelike signa arising from a sclerotized
area on the corpus bursae and possibly the structure of the sterigma (Horak 2006). A falcate or “hooked” apex of
the forewing is characteristic for most species, although this character is shared by species in other genera.
In North America, approximately 35 species of Ancylis are present (Powell 1983), although the exact number
is uncertain. One reason for this uncertainty is the number of taxa in species complexes that have seemingly
identical (and variable) genitalia (Miller 1987, Gilligan et al. 2008). Another reason is the number of species that
are assumed by various authors to be Holarctic; these include A. comptana (Frölich), A. diminutana (Haworth), A.
geminana (Donovan), A. tineana (Hübner), A. uncella (Denis & Schiffermüller), and A. unguicella (Linnaeus)
(Miller 1987, Razowski 2003, Brown 2005, Gilligan et al. 2008, Gilligan et al. 2014a). The majority of these
species fall into two groups with the members of each group having similar male genitalia. In addition, each group
contains new species, three of which are described here.
The European fauna is less diverse than the North American, including 24 species (Aarvik 2013). The
taxonomic history of taxa, particularly of the Ancylis geminana group and the Ancylis unguicella group, is varied
and represented differently in the European versus North American literature. Early European checklists (Stephens
1829) recognized Anchylopera biarcuana, A. diminutana, and A. uncana, treating A. geminana as a synonym of the
latter. Wocke (1871) followed this arrangement (as Phoxopteryx) and also listed subarcuana as a variety of P.
biarcuana and P. uncella as a junior synonym of P. uncana. Phoxopteris and Phoxopteryx were synonymized under
Ancylis by Walsingham (1897). Rebel (1901) recognized A. unguicella, A. uncana (including A. uncella), A.
biarcuana, and A. diminutana, listing A. geminana as a subspecies and subarcuana as a variety of A. biarcuana. It
was nearly a century before an updated list of European Lepidoptera was published, during which time A.
biarcuana and A. uncana were relegated to junior synonyms of A. geminana and A. uncella, respectively, and A.
subarcuana was elevated to species level. Thus, Razowski (1996) recognized A. diminutana, A. geminana, A.
subarcuana, A. uncella, and A. unguicella as separate species in the European checklist.
Early North American checklists (Fernald 1882, 1903) followed the European convention, treating both A.
geminana and A. uncella as synonyms of A. uncana, and A. biarcuana as a separate species. North American
checklists also included A. goodelliana and A. pacificana, and later (Barnes and McDunnough 1917), A.
diminuatana, described as new by Kearfott in 1905 and “close to European biarcuana.” Heinrich (1923) was the
first author to deviate from the European classification of Ancylis for North America. He described A. carbonana,
stating that it was different from A. uncana (= A. uncella), and that the latter species “does not occur in our fauna.”
He relegated Kearfott’s A. diminuatana to a junior synonym, stating that it was “nothing but the European
diminutana redescribed under practically the same name.” Heinrich also speculated that “the so-called biarcuana
Stephens” was the same as A. diminutana (although he did not list it as a synonym). Powell (1983) followed
Heinrich for the most part, recognizing A. carbonana (with North American A. uncana as a synonym), A.
goodelliana, A. unguicella, and A. pacificana. He also listed A. diminutana and included A. diminuatana as a
misspelling under that name. To further complicate matters, Brown (2005), while examining material primarily in
the USNM, determined that A. diminutana does not occur in North America, and specimens determined as such
were actually A. geminana. He also synonymized A. diminuatana and A. subarcuana under A. geminana. This
arrangement was preserved in Gilligan et al. (2014a) but neglected in Europe.
While examining COI DNA barcode sequences in the Barcode of Life Data System (BOLD; Ratnasingham
and Hebert 2007), we noticed several inconsistencies in the clustering of sequences from specimens identified as A.
Zootaxa 4178 (3) © 2016 Magnolia Press
·
349
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
carbonana, A. diminutana, A. goodelliana, and A. uncella from North America. These sequences were clustering in
distinct groups separate from European specimens of supposedly the same identity, and some sequence clusters
contained as many as three separate species. Upon further examination we determined that the majority of North
America Anyclis in these groups were misidentified in BOLD, and that the clusters represented morphologically
distinct species (Fig. 1) that did not correspond with the current taxonomy. We also discovered two related
undescribed species from North America and one from Europe. Here we attempt to resolve these issues by
providing a comprehensive review of these taxa, including a revised taxonomy and descriptions of new species.
Materials and methods
We examined 559 adult specimens (415 ♂, 144 ♀) together with 77 associated genitalia preparations deposited in
the following collections: American Museum of Natural History, New York, New York, U.S.A. (AMNH);
Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Canada (CNC); Colorado State
University, Fort Collins, Colorado, U.S.A. (CSU); Cornell University Insect Collection, Cornell University, Ithaca,
New York, U.S.A. (CUIC); Essig Museum of Entomology, University of California, Berkeley, California, U.S.A.
(EME); Florida State Collection of Arthropods, Gainesville, Florida, U.S.A. (FSCA); Landesmuseum Kärnten,
Klagenfurt, Austria (LMK); Mississippi Entomological Museum, Mississippi State University, Starkville,
Mississippi, U.S.A. (MEM); Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria (TLMF); and National
Museum of Natural History, Washington, D.C., U.S.A. (USNM).
Images of adults were taken with Canon 100 mm and MP-E 65 mm macro lenses attached to a Canon 7D
digital SLR or with an Olympus E 3 digital camera attached to an Olympus SZX 10 binocular microscope. Images
of genitalia were taken with a Nikon DS-Fi1 digital microscope camera attached to a Nikon Labophot-2 compound
microscope or with an Olympus E-1 Digital Camera attached to an Olympus BH2 microscope. All images were
edited using Photoshop CS6 and some are composite stacks of many individual images created with Zerene
Stacker. Forewing length (FWL) is defined as the distance from the base to the apex including the fringe, reported
to the nearest half millimeter. Measurements were made with a stereomicroscope equipped with an ocular
micrometer or a compound microscope using a slide micrometer. The number of observations supporting a
particular statistic is indicated by “n =.” Dissection methods follow those presented in Brown and Powell (1991),
and morphological nomenclature follows Horak (2006) and Gilligan et al. (2008).
Dried legs were prepared according to predescribed standards and processed at the Canadian Centre for DNA
Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) to obtain COI DNA barcodes using the
standard high-throughput protocol (deWaard et al. 2008). DNA sequences > 200 bp from our material and public
sequences from BOLD were considered for analysis. Further details including complete voucher data and images
can be accessed in the public dataset “Lepidoptera—Ancylis” (dx.doi.org/10.5883/DS-LEANCYLIS) in the
Barcode of Life Data Systems (BOLD; Ratnasingham & Hebert 2007). Degrees of intra- and interspecific variation
of DNA barcode fragment were calculated under Kimura 2 parameter model of nucleotide substitution using
analytical tools of BOLD systems v. 3.0. (http:// www.boldsystems.org). A neighbor-joining tree of DNA barcode
data was constructed under the Kimura 2 parameter model for nucleotide substitutions.
Results and discussion
Ancylis unguicella group
The taxa treated here can be divided into two groups based primarily on the structure of the male genitalia. The first
group, the A. unguicella group, contains A. unguicella, A. pacificana, A. uncella, A. goodelliana, and A.
oregonensis. Of these, A. unguicella and A. uncella have a Holarctic distribution, the latter as a result of being
synonymized here with A. carbonana.
Species in this group can be identified using a combination of wing pattern and genitalia. Male genitalia
exhibit some intraspecific variation but are generally more diagnostic than in the A. geminana group. Female
genitalia are relatively uniform but have a few species-specific characters. Shared characters include:
GILLIGAN ET AL.
350
·
Zootaxa 4178 (3) © 2016 Magnolia Press
FIGURE 1. Neighbor-joining tree of COI DNA barcode data obtained from BOLD (K2P model). Clusters representing
morphologically distinct species are color-coded with the corresponding species name.
Zootaxa 4178 (3) © 2016 Magnolia Press
·
351
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Male genitalia. Uncus bifid, well developed in all species except A. oregonensis. Socii large, membranous,
setose pads. Valva with costal margin concave; saccular angle weakly to moderately developed; neck width
relatively uniform; cucullus densely setose, outer margin convex, apex acute to broadly rounded. Caulis long;
phallus 0.5 to equal length of valva; vesica with numerous deciduous lanceolate cornuti.
Female genitalia. Papillae anales moderately setose. Apophyses posteriores and apophyses anteriores
approximately the same length. Lamella postvaginalis membranous, undefined; lamella antevaginalis sclerotized,
projecting ventrally above ostium, often with slight median indentation, forming a funnel-shaped antrum that is
sclerotized 0.4–0.6 length of ductus bursae (except in A. oregonensis). Colliculum present as two lateral sclerites of
varying length continuous with anterior sclerotization of antrum. Ductus seminalis arising from near junction of
ductus and corpus bursae. Corpus bursae large, oval; two blade- or horn-shaped signa present.
Ancylis unguicella (Linnaeus, 1758)
Figs. 2–6, 59–60, 75
Phalaena (Tinea) unguicella Linnaeus, 1758, Systema Naturae (10th ed.): 536.
Phalaena ungvicella Clerck, 1759, Icones Insectorum Rariorum 1: pl. 12, fig. 7. [misspelling of unguicella]
Pyralis unguicana Fabricius, 1775, Systema Entomologiae: 654. [unjustified emendation]
Tortrix falcana Hübner, 1796–1799, Samml. Eur. Schmett. 7: pl. 13, fig. 78.
Tortrix vappana Hübner, 1814–1817, Samml. Eur. Schmett. 7: pl. 38, fig. 241.
Anchylopera plagosana Clemens, 1864, Proc. ent. Soc. Philad. 3: 417.
Diagnosis. Ancylis unguicella is one of few Anyclis with dark fasciate wing markings. In the Palearctic, A.
achatana has a similar wing pattern, but the median fascia is not as well defined, and the male genitalia differ with
a narrow valval neck and well–defined 90° saccular angle. The Nearctic A. pacificana is identical in wing pattern to
A. unguicella, and the two species cannot be separated without dissection. In the male, the cucullus of A. unguicella
is blunter with a nearly acute apex versus the elongate rounded cucullus in A. pacificana, and the phallus is longer
(0.75 as long as valva in A. unguicella; 0.5–0.6 as long as valva in A. pacificana). In the female, the antrum of A.
unguicella is wider posteriorly and the signa are smaller than in A. pacificana.
Redescription. Forewing. FWL ♂ 7–8.5 mm (n=102), ♀ 7–9 mm (n=8). Forewings are gray and brown, with
a brown to dark brown median fascia that is complete from costa to dorsum, white costal strigulae, and silvery
striae. Some individuals have tan or light gray-tipped scales interspersed throughout the entire wing, especially in
interfascial areas. Male genitalia. Uncus bifid to approximately half its length. Valva with shallow basal excavation
extending to middle of neck; saccular angle weakly developed with variable triangular terminal projection; neck of
uniform width from sacculus to cucullus; cucullus blunt, densely setose, outer margin rounded with several rows of
short stout setae, apex nearly acute; caulis 0.5 to 0.75 as long as phallus; phallus 0.75 as long as valva, with small
triangular tooth just proximal to apex; vesica with ca. 40–60 deciduous lanceolate cornuti. Female genitalia.
Antrum sclerotized to 0.5 length of ductus bursae, widened at ostium to near distance between apophyses
anteriores. Corpus bursae large, oval, expanding abruptly from ductus bursae; signa blade-shaped, unequal in size.
Molecular data. BIN URI: BOLD:AAB3498. The intraspecific divergence of the barcode region is moderate
with average 0.84% and maximum 2.36% (n=46). However, North American and European populations cluster
separately with a minimum distance of 0.87%. The minimum distance to the nearest neighbor A. mediofasciana is
4.5%.
Distribution. Ancylis unguicella has a Holarctic distribution. In the Palearctic, it is found from Western
Europe to Siberia, the Korean Peninsula, and Japan (Razowski 2003). In the Nearctic, it is present from Alaska and
British Columbia east to Ontario and south to Colorado.
Biology. Adults are present from May to early July. Larvae feed from July to August on various species of
Erica (Ericaceae) and on Calluna (Ericaceae). Pupation takes place in early spring after larval hibernation in the
final instar (Bradley et al. 1979; Razowski 2001). Ancylis unguicella prefers heathland and moors.
Remarks. This species is identical in wing pattern to A. pacificana. In the Pacific Northwest, where A.
pacificana is also present, species-level determinations should rely on the male genitalia. We were unable to locate
any specimens of A. unguicella from California or Oregon. Despite the weak genetic divergence of North
American populations, we treat them as conspecific due to the morphological conformity.
GILLIGAN ET AL.
352
·
Zootaxa 4178 (3) © 2016 Magnolia Press
FIGURES 2–16. Adults. 2–6, A. unguicella (2, Italy; 3–4, Germany; 5, Washington; 6, South Dakota). 7–10, A. pacificana
(California). 11–16, A. uncella (11–12, Germany; 13, Japan; 14, Pennsylvania, A. carbonana holotype; 15, Connecticut; 16,
Virginia).
Zootaxa 4178 (3) © 2016 Magnolia Press
·
353
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
FIGURES 17–31. Adults. 17–22, A. goodelliana (17, northeastern U.S.A., holotype; 18, Connecticut; 19, New York; 20–21,
North Carolina; 22, Nova Scotia). 23 – 25, A. oregonensis (23, Oregon, holotype; 24–25, Oregon). 26–31, A. geminana (26–27,
[no data]; 28–29, Germany; 30, Italy; 31, Austria).
GILLIGAN ET AL.
354
·
Zootaxa 4178 (3) © 2016 Magnolia Press
FIGURES 32–46. Adults. 32–34, A. christiandiana (Austria). 35–38, A. diminutana (Germany); 39–43, A. diminuatana (39,
New Jersey, holotype; 40, Ohio; 41, Nebraska; 42, Manitoba; 43, Washington). 44–46, A. diminuatana complex (44,
Washington; 45, Colorado; 46, Alaska).
Zootaxa 4178 (3) © 2016 Magnolia Press
·
355
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
FIGURES 47–58. Adults. 47–52, A. saliana (47, Florida, holotype; 48–52, Florida). 53–58, A. subarcuana (53, Austria; 54–
57, Germany; 58, Finland).
Ancylis pacificana (Walsingham, 1879)
Figs. 7–10, 61–62, 76
Phoxopteryx pacificana Walsingham 1879, Illust. typical Specimens Lepid. Heterocera Colln. Br. Mus. 4: 73.
Diagnosis. Ancylis pacificana is identical in wing pattern to A. unguicella, and the two species cannot be separated
without dissection. In the male, the cucullus of A. pacificana is more elongate with a rounded apex versus the
shorter, blunt cucullus in A. unguicella, and the phallus (and caulis) is shorter in A. pacificana. In the female, the
antrum of A. pacificana is narrower posteriorly and the signa are larger than in A. unguicella.
Redescription. Forewing. FWL ♂ 7.5–10 mm (n=18), ♀ 8.3 mm (n=1). Forewing description as in A.
GILLIGAN ET AL.
356
·
Zootaxa 4178 (3) © 2016 Magnolia Press
unguicella. Male genitalia. Uncus bifid to approximately half its length. Valva with shallow basal excavation
extending to just beyond sacculus; saccular angle weakly to moderately developed with variable triangular terminal
projection; neck of uniform width or slightly narrowed from sacculus to cucullus; cucullus elongate, densely
setose, outer margin convex with several rows of short setae near the anal angle, apex rounded; caulis 0.5 as long as
phallus; phallus 0.5–0.6 as long as valva, with small triangular tooth just proximal to apex; vesica with ca. 40–60
deciduous lanceolate cornuti. The length of the cucullus, the width of the valval neck, and the saccular terminal
projections all vary between individuals; we have attempted to illustrate the range of variation in Figs. 58–59.
Female genitalia. Antrum sclerotized to 0.4 length of ductus bursae, widened at ostium to 0.5 distance between
apophyses anteriores. Corpus bursae oval, expanding gradually from ductus bursae; signa large, blade- or horn-
shaped, unequal in size.
Distribution. Because of its similarity with A. unguicella, the distribution of A. pacificana is difficult to
determine from undissected museum specimens. Heinrich (1923) reported specimens from California, British
Columbia, and Colorado, whereas Powell and Opler (2009) reported a distribution limited to montane habitats in
Oregon and northern California. The specimens that we determined as A. pacificana were from California, Oregon,
and British Columbia, although we discovered one male in the EME from San Bernardo, Sonora, Mexico. This
suggests that the distribution of A. pacificana is more extensive than assumed by previous authors, although we
have not confirmed A. pacificana from any other locations (including Colorado). In any event, undissected
museum specimens identified as A. pacificana should be treated as suspect until verified using genitalic characters.
Molecular data. Unknown.
Biology. Adults are present May to July. The larval host is unknown, although Powell and Opler (2009) report
adults associated with Ceanothus.
Remarks. Heinrich (1923) stated that A. pacificana was larger and less darkly marked than A. unguicella. We
found that, whereas some specimens of A. pacificana are slightly larger, both species overlap in size. We also found
smaller specimens of A. pacificana with dark markings (e.g., Fig. 9) that were incorrectly identified as A.
unguicella. Species-level determinations of either species from the Pacific Northwest should rely on male genitalia.
Ancylis uncella (Denis and Schiffermüller, 1775)
Figs. 11–16, 63–64, 77–78
Tinea uncella Denis & Schiffermüller, 1775, Syst. Verz. Schmett. Wienergegend: 136.
Tortrix uncana Hübner, 1796–1799, Samml. Eur. Schmett. 7: pl. 13, fig. 76. [unjustified emendation]
Ancylis uncana var. subuncana Krulikowsky, 1907, Rev. Russe Ent. 7: 33.
Ancylis carbonana Heinrich, 1923, Bull. U.S. natn. Mus. 123: 248. syn.n.
Diagnosis. Most individuals of A. uncella can be diagnosed by forewing pattern: the median fascia is brown to
reddish brown, complete from costa to dorsum, and flanked on the dorsum and tornus by gray patches. Some A.
goodelliana can appear similar (Fig. 18), but the median fascia is not expressed on the costa in that species.
Redescription. Forewing. FWL ♂ 5–9 mm (n=30), ♀ 6–8 mm (n=14). Forewings are brown to reddish brown
and gray with white to gray costal strigulae. The brown to reddish-brown median fascia is complete from costa to
dorsum in most individuals; occasionally the dorsal portion is broken before reaching the dorsum (Fig. 13), but in
all cases the median fascia is expressed on the costa. Gray to light gray patches are present proximal to the median
fascia on the dorsum and distal to the median fascia on the tornus; often median fascia extends into the distal gray
patch (Fig. 12–13, 16). In some individuals the brown and gray areas of the wing are sharply contrasting (Fig. 11),
while in others the gray is darker and subdued (Fig. 14). Black streaks are sometimes weakly expressed along the
radius and cubitus (Figs. 12–14). Male genitalia. The male genitalia are identical to those of A. pacificana with the
exception of the phallus, which is 0.7 as long as the valva, and the tooth on the phallus, which is at the apex (and
difficult to see in some preparations). As with A. pacificana, the length of the cucullus, the width of the valval neck,
and the saccular terminal projections all vary between individuals; we have attempted to illustrate the range of
variation in Figs. 63–64. Female genitalia. Antrum sclerotized to 0.6 length of ductus bursae, widened at ostium to
0.5 distance between apophyses anteriores. Corpus bursae large, oval, expanding abruptly from ductus bursae;
signa large, blade- or horn-shaped, unequal in size.
Zootaxa 4178 (3) © 2016 Magnolia Press
·
357
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
FIGURES 59–66. Male genitalia. 59–60, A. unguicella (59, Colorado, TMG685; 60, Austria, TMG681). 61–62, A. pacificana
(61, California, TMG678; 62, California, TMG686). 63–64, A. uncella (63, Russia, TMG679; 64, Maine, TMG698). 65, A.
goodelliana (Connecticut, TMG705). 66, A. oregonensis (Oregon, TMG633).
GILLIGAN ET AL.
358
·
Zootaxa 4178 (3) © 2016 Magnolia Press
FIGURES 67–74. Male genitalia. 67, A. geminana ([no data], TMG618). 68, A. christiandiana (Austria, TOR 464 P. Huemer).
69, A. diminutana (Germany, TMG616). 70–71, A. diminuatana (70, Maryland, TMG688; 71, Nebraska, TMG635). 72, A.
saliana (Florida, TMG676). 73–74, A. subarcuana (73, Austria, TMG619; 74, Germany, TMG703).
Zootaxa 4178 (3) © 2016 Magnolia Press
·
359
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
FIGURES 75–80. Female genitalia (arrows denote origination of ductus seminalis). 75, A. unguicella (Germany, TMG682).
75, A. pacificana (California, TMG687). 77–78, A. uncella (74, Germany, TMG680; 75, Pennsylvania, TMG700). 79, A.
goodelliana (New York, TMG704). 80, A. oregonensis (Oregon, TMG630).
GILLIGAN ET AL.
360
·
Zootaxa 4178 (3) © 2016 Magnolia Press
FIGURES 81–86. Female genitalia (arrows denote origination of ductus seminalis). 81, A. geminana (Germany, TMG675). 82,
A. christiandiana (Austria, TOR 478 P. Huemer). 83, A. diminutana (Germany, TMG617). 84, A. diminuatana (Ohio,
TMG412). 85, A. saliana (Florida, TMG629). 86, A. subarcuana (Germany, TMG620).
Zootaxa 4178 (3) © 2016 Magnolia Press
·
361
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Molecular data. BIN URI: BOLD:AAA7191, BIN-sharing with A. goodelliana, with a minimum distance of
1.13%.
Distribution. With the synonymy of A. carbonana, A. uncella has a Holarctic distribution. In the Palearctic, it
is found from the United Kingdom and France south to Romania and east to Siberia, the Korean Peninsula, and
Japan (Razowski 2003). In the Nearctic, it is present across Canada from Ontario to Manitoba and Alberta, and in
the U.S. from Maine south to Pennsylvania and Virginia.
Biology. Adults are present from late April to June, sometimes also in a small second generation in July and
August (Razowski 2003). Larvae feed in July and August on Betula and Erica and pupate after hibernation in a
cocoon spun in a dead leaf (Bradley et al. 1979).
Remarks. Heinrich (1923) described A. carbonana as different from A. uncana (= A. uncella) based on darker
hindwings, a fainter mid-dorsal pale patch (on the forewing), a shorter phallus, and a narrower valva. We compared
the holotype and several paratypes of A. carbonana with A. uncella from Europe and found that these characters
vary to the extent that they will not reliably separate individuals from Europe and North America. Further,
individuals of A. uncella from Europe and A. carbonana from Canada (identified as A. carbonana, A. goodelliana,
or A. uncella) clustered together in the DNA barcode analysis (Fig. 1). Based on this evidence we synonymize A.
carbonana with A. uncella.
Ancylis goodelliana (Fernald, 1882)
Figs. 17–22, 65, 79
Phoxopteris goodelliana Fernald 1882, Trans. Am. ent. Soc. 10: 69.
Diagnosis. The light gray to white costal margin and black streaks along the radius and cubitus of the forewing
separate A. goodelliana from other Ancylis treated here. Some individuals of A. uncella appear similar, but the
brown to reddish-brown median fascia is always expressed on the costa in A. uncella. Lighter (or worn) individuals
of A. diminuatana may also appear similar, although the costa is usually not as pale in the distal half, and A.
diminuatana lacks the black streak along the radius.
Redescription. Forewing. FWL ♂ 6–9.5 mm (n=16), ♀ 6.5–8.5 mm (n=4). Forewings are brown to reddish
brown and gray with white costal strigulae near the apex. The costa is white to light gray from base to near apex,
and the same color is present along the dorsum. No fasciae are defined, instead a brown to reddish-brown band runs
from base to apex; the dorsal margin of this band is sinuate as a result of remnants of the median fascia extending
towards the tornus. Two black streaks are present: one along the cubitus to ca. one-third the distance to the termen,
and one along the distal two-thirds of the radius to the apex (most evident in Fig. 22). Male genitalia. Uncus bifid
to more than half its length. Valva with basal excavation nearly absent; saccular angle weakly to moderately
developed with variable triangular terminal projection; neck of uniform width or slightly narrowed from sacculus
to cucullus; cucullus elongate, tapering towards apex, densely setose, outer margin convex with several rows of
short setae near the anal angle, apex rounded. Caulis 0.5 as long as phallus; phallus 0.7 as long as valva, with small
tooth at apex (difficult to see in some preparations); vesica with ca. 40–70 deciduous lanceolate cornuti. The length
of the cucullus, the width of the valval neck, and the saccular terminal projections all vary slightly between
individuals, although genital variation is not as pronounced as in other species in this group. Female genitalia. As
in A. uncella.
Molecular data. BIN URI: BOLD:AAA7191, BIN-sharing with A. uncella, with a minimum distance of
1.13%.
Distribution. Most specimens of A. goodelliana are from eastern North America: Nova Scotia west to Ontario,
and Wisconsin south to North Carolina and Florida. Heinrich (1923) reported this species from Manitoba and
Colorado, and we examined one specimen from central Alberta in the CUIC, suggesting that it also present in the
West. Historical records are difficult to confirm because of confusion with A. diminuatana in many collections.
Biology. Adults are present from the end of May through mid-August. The larval host is unknown.
Remarks. Worn specimens of A. goodelliana are easily confused with A. diminuatana, and the two are often
mixed in collections. A genitalic dissection will easily separate A. goodelliana from all species in the A. geminana
group. Gilligan et al. (2008) incorrectly illustrated the adult and male genitalia of A. goodelliana as “A.
GILLIGAN ET AL.
362
·
Zootaxa 4178 (3) © 2016 Magnolia Press
diminutana.” Although A. goodelliana and A. uncella appear very similar based on COI barcode data (with a
minimum distance of only 1.13%), the two species can be reliably separated by forewing pattern.
Ancylis oregonensis Gilligan and Huemer, sp.n.
Figs. 23–25, 66, 80
Type material. Holotype. ♂: “Oregon, Crescent Lake, Klamath Co., 4600‘ 7.VII 1955, JFG Clarke“ (USNM).
Paratypes (9). United States: 3 ♂, 6 ♀ [same data as holotype], slides TMG630, TMG633 (USNM).
Diagnosis. Ancylis oregonensis is distinguished from all other species treated here by the discontinuous
longitudinal line of the forewing, which is broken or obscured in the middle in most specimens. In individuals
where the longitudinal line is nearly continuous (Fig. 25), the costal and dorsal halves of the forewing are not
deeply contrasting. The male genitalia are also unique in this species, with the cucullus well-defined, the phallus as
long as the valva, and the uncus reduced.
Description. Head, labial palps and thorax light brownish gray. Forewing. FWL ♂ 6.5–8.7 mm (n=4), ♀ 7.0–
7.8 mm (n=6). Forewing is falcate. The costal half of the wing is a mix of light brownish gray, dark brown, and
black. The costa is light grayish brown in the basal half becoming a mix of gray, brown, and dark brown in the
apical half with remnants of dark brown costal strigulae in some individuals. The dorsal half of the wing is light
brownish gray. The longitudinal line starts at A
1+2
, extends towards the termen, and is disrupted in the middle one-
third of the wing by brownish gray scales of the dorsal half of the wing extending to the cubitus. Beyond the
disruption the longitudinal line is usually well defined, continuing from CuA
2
straight to the apex. Cilia at the apex
dark brown to black with a white postapical strigula. Hindwing pale grayish brown. Male genitalia. Uncus bifid,
reduced, ca. 0.5 as tall as wide. Valva with shallow basal excavation extending to neck; saccular angle strongly
developed with triangular terminal projection; neck narrowed from sacculus to cucullus, ventral margin semi-
circular; cucullus well-defined, densely setose, outer margin convex with several rows of short setae, costa weakly
convex. Caulis 0.25 as long as phallus; phallus as long as valva, with small triangular tooth at apex; vesica with ca.
45 deciduous cornuti. Female genitalia. Apophyses posteriores 0.75 as long as apophyses anteriores. Lamella
postvaginalis membranous, undefined; lamella antevaginalis sclerotized, projecting ventrally above ostium; antrum
weakly developed; colliculum two small lateral sclerites. Ductus seminalis arising at junction of ductus and corpus
bursae. Corpus bursae large, oval; two horn-shaped signa present.
Molecular data. Unknown.
Distribution. Ancylis oregonensis is known only from the type locality, Crescent Lake, located on the eastern
side of the Cascades in Klamath County, Oregon.
Biology. The only collection data are from the type series, and all specimens were collected on the same night
in early July. The larval host is unknown.
Etymology. The name signifies that this species has only been recorded from Oregon.
Ancylis geminana group
The A. geminana group contains A. geminana, A. christiandiana, A. diminutana, A. diminuatana, A. subarcuana,
and A. saliana. Ancylis diminuatana and A. subarcuana are elevated from synonymy with A. geminana, and A.
christiandiana and A. saliana are described as new. As a result of these taxonomic changes, none of the species in
this group is considered to be Holarctic in distribution.
Species identification in this group is difficult and usually relies on minor differences in wing pattern. Male
genitalia are variable and useless in discriminating species; we illustrate each species to display the range of
variability in this group. Female genitalia are uniform with species-specific characters for only a couple taxa.
Shared characters include:
Forewing. Fasciae are undefined; instead the wing is divided longitudinally along the radius and/or cubitus
into a darker costal half and a lighter dorsal half, creating a two-toned appearance in some species. The border
between the costal and dorsal halves creates a sinuate “longitudinal line,” often bordered in white, which runs from
base to apex. The shape and degree of expression of the longitudinal line is often useful in diagnosing species.
Zootaxa 4178 (3) © 2016 Magnolia Press
·
363
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Male genitalia. Uncus bifid to 0.75–0.9 its total length, well developed in all species. Socii large membranous
setose pads. Valva with costal margin concave to nearly straight; saccular angle moderately to strongly developed
with variable triangular terminal projection; neck width variable, from moderate to wide; cucullus with rounded
dorsal lobe and narrow fingerlike ventral lobe, outer margin concave to straight, variably setose with rows of short
setae along dorsal lobe and outer margin. Caulis 0.5–0.7 as long as phallus; phallus 0.4–0.5 as long as valva; vesica
with > 100 deciduous lanceolate cornuti.
Female genitalia. Papillae anales moderately setose. Apophyses posteriores the same length or 0.75 as long as
apophyses anteriores. Lamella postvaginalis membranous, undefined; lamella antevaginalis sclerotized, projecting
ventrally above ostium, often with slight median indentation, forming a weakly sclerotized funnel-shaped antrum.
Colliculum present as two small lateral sclerites continuous with anterior sclerotization of antrum. Ductus
seminalis arising posterior to junction of ductus and corpus bursae. Corpus bursae large, oval; two blade- or horn-
shaped signa present.
Variation. The shape of the male valva varies greatly. The width of the neck, and corresponding degree of
development of the dorsal lobe of the cucullus, can vary extensively within species (e.g., Figs. 70–71, 73–74) or
even within the same individual (e.g., Fig. 71). The shape of the ventral lobe of the cucullus also varies within
species and individuals. Note that the appearance of valval shape is influenced by the method of slide preparation;
any tilting or twisting of the genitalia during mounting will alter the appearance of the cucullus and the sacculus.
The male genitalia figures presented here (Figs. 67–74) illustrate the range of variability found within this group;
however, none of the variability is specific to a particular species and any of the figures could represent any of the
species treated here.
Ancylis geminana (Donovan, 1806)
Figs. 26–31, 67, 81
Phalaena geminana Donovan, 1806, Nat. Hist. Br. Insects 11: 29.
Anchylopera biarcuana Stephens, 1834, Illust. Br. Ent. (Haustellata) 4: 113.
Phoxopterix crenana Duponchel, in Godart, 1835, Hist. nat. Lepid. Papillons Fr. 9: 334.
fluctigerana [uninomial] Herrich-Schäffer, 1848, Syst. Bearbeitung Schmett. Eur. 4: pl. 45, fig. 319. [nomen nudum]
Tortrix (Phoxopteryx) fluctigerana Herrich-Schäffer, 1851, Syst. Bearbeitung Schmett. Eur. 4: 286.
Diagnosis. Ancylis geminana is distinguished by the grayish brown coloration on the costal half of the forewing,
which is lighter on the costa and darker towards the longitudinal line, and the uniform curvature of the longitudinal
line. It is most similar to A. christiandiana from which it differs by the distinctly curved longitudinal line and the
more variegated light and dark wing pattern. The smaller A. diminutana is reddish brown with a longitudinal line
similar to A. christiandiana.
Molecular data. BIN URI: BOLD:AAE1223. The intraspecific divergence of the barcode region is low with
average 0.28% and maximum 0.8% (n=23). The minimum distance to the nearest neighbor A. christiandiana is
3.82%.
Redescription. Forewing. FWL ♂ 7.5–9.5 mm (n=10), ♀ 7–8 mm (n=2). The costal half of the wing is a mix
of grayish brown, dark brown, and black. The costa is lighter grayish brown with darker markings towards the
longitudinal line. The dorsal half of the wing is light brownish gray to near white. The longitudinal line is
continuous from base to just below the apex and is often bordered in white dorsally. The line starts at A
1+2
, curves
evenly up to the cubitus, back down to CuA
2
, and then up to radius, angling down slightly to reach the termen near
M
1
. Small black streaks are often present on R
5
just proximal to the termen. Male genitalia. As described for the
group. Female genitalia. As described for the group with the following modifications: apophyses posteriores and
apophyses anteriores approximately the same length; antrum weakly sclerotized to the posterior 0.15 of the ductus
bursae; and ductus seminalis arising in the anterior 0.3 of the ductus bursae.
Distribution. Ancylis geminana is distributed across much of Western Europe to Asia Minor, Mongolia,
Siberia, China, and Japan (Razowski 2003).
Biology. Adults are present from May to August with suspected occasional bivoltinism (Razowski 2001,
2003). However, our data as well as various alternative literature sources (e.g. Schütze 1931) support a single
generation of adults primarily in May and June. The reported larval host is willow (Salix spp., Salicaceae)
GILLIGAN ET AL.
364
·
Zootaxa 4178 (3) © 2016 Magnolia Press
(Razowski 2003), although this species prefers different kind of hygrophilous to mesophilous woods such those
found in riverine forests.
Remarks. Phalaena geminana was described from material collected on one occasion in Kent. Donovan
(1806: pl. 370, fig. 1, 1) figured two slightly deviating specimens, indicating that the species was described from
more than a singleton. No type material could be found in the collections of the Natural History Museum (London,
UK), but the precise color figures of the original description leave no doubt of the identity.
Prior to the release of the first world tortricid catalogue (Brown 2005), the majority of Ancylis in North
America with a wing pattern similar to species in the A. geminana group were identified as A. diminutana. Brown
(2005) realized that A. diminutana did not occur in North America, and determined that specimens in the USNM
under that name were actually A. geminana. Thus, he synonymized A. diminuatana (formally determined to be
nothing more than A. diminutana from North America by Heinrich) with A. geminana. He also included A.
subarcuana as a synonym of A. geminana.
DNA barcode data has helped greatly in solving this taxonomic chaos. Figure 1 shows that A. geminana is
clearly separate from A. subarcuana, and both are only found in the Palearctic. The remaining specimens from
North America, mostly misidentified as A. carbonana, match the type of A. diminuatana. All three species can be
reliably separated from each other using forewing pattern.
Ancylis christiandiana Huemer and Wiesmair, sp.n.
Figs. 32–34, 68, 82
Type material. Holotype. ♂: “Austria – Kärnten Griffen, Griffener See 480m 11.6.2004 – LF leg. H. Deutsch”
“BC TLMF Lep 03394” (TLMF).
Paratypes (20). Austria: 1 ♂, same data as holotype, but barcode sample ID BC TLMF Lep 03395, gen. slide
TOR 464 ♂ P. Huemer (coll. Helmut Deutsch, Bannberg, Austria); 1 ♂, ditto, but barcode sample ID BC TLMF
Lep 03396, gen. slide 07/098 ♂ H. Deutsch (coll. Helmut Deutsch, Bannberg, Austria); 1 ♂, ditto, but Griffner See,
Bruchwald, 490 m, AT-BMN 31 14,721609 E / 46,697111 N, 27.5.2016, leg. Dr. C. Wieser (LMK); 1 ♂, ditto, but
3.6.2016 (LMK); 1 ♂, Kärnten, Finkenstein, S Höfling, Finkensteiner Moor W, 536 m, 13°52´45´´E, 46°34´06´´N,
27.5.2016, leg. Huemer (TLMF); 2 ♂, ditto, but 28.5.2016 (TLMF); 1 ♂, Carinthia, Lavant Au, Aufweitung Aich,
Insel, 385 m, AT-BMN 31 14,856502 E / 46,728070 N, 31.5.2016, leg. Dr. C. Wieser (LMK); 1 ♂, Carinthia, St.
Paul/L. bei Aich, 395 m; AT-BMN 31 14,86042 E / 46,72273 N, 21.5.2014, leg. D. Wieser (LMK); 1 ♀, Carinthia,
Flughafen, Witternitz, Rückhalteb. 3, 440 m, 26.5.2008, AT-BMN 31 14,354825E / 46,642159N, leg. Dr. C.
Wieser, Kärntner Landesmuseum, gen. slide TOR 478 ♀ P. Huemer (LMK); 1 ♂, ditto, but barcode sample ID BC
TLMF Lep 06853, gen. slide TOR 462 ♂ P. Huemer (LMK); 1 ♀, ditto, but 23.6.2008 (LMK); 1 ♀, ditto, but
Rückhalteb. 1, AT-BMN 31 14,355872 E / 46,642029 N, 26.5.2008 (LMK); 2 ♂, ditto, but Rückhalteb. 4, AT-BMN
31 14,353762 E / 46,642417 N, 26.5.2008 (LMK); 1 ♂, ditto, but 24.6.2008 (LMK); 1 ♂, Carinthia, Großedlinger
Teich, 420m, 14.6.2005, AT-BMN 31 14,843800 E / 46,795742 N , leg. Dr. C. Wieser, Kärntner Landesmuseum,
barcode sample ID BC TLMF Lep 06854 (LMK); 2 ♂, Carinthia, Eberndorf, Sablatnigmoor, Lichtfalle 1, 550m,
5–8.6.1989, leg. Dr. C. Wieser, Kärntner Landesmuseum, gen. slide T390 ♂ C. Wieser (LMK); Germany: 1 ♂,
Schwaben, Dillingen a.d.D., Wertingen, Wertinger Ried, 10.6.2016, leg. Heindel, barcode sample ID BC ZSM Lep
ID 71187 (coll. Richard Heindel, Günzburg, Germany).
Diagnosis. Ancylis christiandiana is similar to A. diminutana and A. geminana, but all three species can be
distinguished by a combination of wing pattern, coloration, and size. The costal half of the forewing is dark grayish
brown in A. christiandiana versus lighter reddish brown in A. diminutana. The longitudinal line is more angulate in
A. christiandiana and reaches the termen in a diagonal shallowly curved line near the apex, whereas the
longitudinal line in A. geminana is distinctly curved, and the coloration of the forewing in A. christiandiana is
rather monotonous and less contrasting than in A. geminana. Both A. christiandiana and A. geminana are larger
than A. diminutana. The tubular shape of the antrum with a straight posterior edge of the lamella antevaginalis is
possibly diagnostic in A. christiandiana; in A. geminana the posterior part of the lamella antevaginalis is weakly
convex whereas in A. diminutana it is concave and the antrum is funnel shaped.
Description. Head, labial palpi and thorax light grayish brown. Labial palpi and head with scattered black
scales.
Zootaxa 4178 (3) © 2016 Magnolia Press
·
365
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Forewing. FWL ♂ 8.1 – 9.1 mm (n=6), ♀ 7.6 mm (n=1). Forewing is falcate. The costal half of the wing is a
mix of grayish brown, dark brown, and black. The costa is light grayish brown with darker markings towards the
apex. The dorsal half of the wing is light brownish gray to near white, with several black dots present on the
dorsum. The longitudinal line is continuous from base to apex and is bordered in whitish gray dorsally. The line
starts at A
1+2
, curves up to the cubitus, angles back down to CuA
2
and then runs in a diagonal line straight to the
apex. A row of black scales runs along the termen. The cilia at the apex are black with a white postapical strigula;
other cilia are pale on the basal half and dark on the apical half. Hindwing is gray. Male genitalia. As described for
the group. Female genitalia. As described for the group with the following modifications: apophyses posteriores
and apophyses anteriores approximately the same length; posterior edge of the lamella antevaginalis straight; and
antrum tubular.
Molecular data. BIN URI: BOLD:AAV7672. The intraspecific divergence of the barcode region is 0% (n=5).
The minimum distance to the nearest neighbor A. geminana is 3.85%.
Distribution. Ancylis christiandiana is known from a few localities in southern Austria and Germany, but it is
probably more widespread and expected to occur at additional sites.
Biology. The few adults known to date have been collected in late May and June at light. The larval host is
unknown but this species likely feeds on willow (Salix spp.) like other members of the group. Ancylis
christiandiana seems to prefer hygrophilous woodland.
Etymology. The species name christiandiana is an artificial compound form of the forenames of two
colleagues who contributed to this study, Christian Wieser and Andi [Andreas] Segerer.
Ancylis diminutana (Haworth, 1811)
Figs. 35–38, 69, 83
Tortrix diminutana Haworth, 1811, Lepid. Br. (3): 452.
Phoxopteris cuspidana Treitschke, 1830, Schmett. Eur. 8: 236. syn.rev.
Diagnosis. Ancylis diminutana is distinguished by the reddish-brown coloration on the costal half of the forewing
and the shape of the longitudinal line, which curves broadly from the cubitus down to CuA
2
, and then angles
straight to the apex. Both A. christiandiana and A. geminana are similar: A. geminana is grayish brown and the
longitudinal line is evenly curved; and A. christiandiana is larger, lacks the reddish brown color of the forewing
and the longitudinal line is more angulate.
Redescription. Forewing. FWL ♂ 6–8 mm (n=16), ♀ 6–7 mm (n=5). The costal half of the wing is reddish
brown. The costa is lighter in some specimens, but only at the base. The dorsal half of the wing is brownish gray in
the median portion, becoming lighter gray to near white at the tornus and along the termen. The longitudinal line,
which is often bordered in white dorsally, starts at A
1+2
, curves up to the cubitus, back down to CuA
2
, and then runs
in a diagonal line straight to the apex. Small black streaks are often present on R
5
just proximal to the termen. Male
genitalia. As described for the group. Female genitalia. As described for the group with the following
modifications: apophyses posteriores 0.6–0.7 as long as apophyses anteriores; antrum weakly sclerotized just
below the ostium; and ductus seminalis arising in the anterior 0.3–0.4 of the ductus bursae.
Molecular data. BIN URI: BOLD:AAB6876. The intraspecific divergence of the barcode region is low with
average 0.1% and maximum 0.64% (n=21). The minimum distance to the nearest neighbor A. geminana is 4.73%.
Distribution. Ancylis diminutana is locally distributed from the northwestern, northern and central parts of
Europe (Razowski 2003) to central Siberia (Sinev 2008).
Biology. Adults are present from May to June and from July to August (Schütze 1941). The larval host is
willow (Salix spp.) (Razowski 2003). The species prefers different kind of hygrophilous to mesophilous woods
such as those found in riverine forests.
Remarks. Tortrix diminutana was described from an unspecified number of specimens without precise locality
data from Great Britain. According to Sattler (in litt.) no type material could be found in the collections of the
Natural History Museum (London, UK). However, the original description precisely points to the comparatively
small size and the reddish brown color of the costal half.
Phoxopteris cuspidana was described from an unspecified number of specimens collected in Hungary and
GILLIGAN ET AL.
366
·
Zootaxa 4178 (3) © 2016 Magnolia Press
Germany (Saxonia). The species was previously synonymized with A. geminana. However, photographs of two
type specimens (courtesy of Laszlo Ronkay) verify that the species fully agrees with typical A. diminutana both
from the small wingspan and the reddish color of the costal half of the forewing. We therefore formally
synonymize P. cuspidana with A. diminutana. To serve stability we furthermore designate the specimen with
following labels as the lectotype: “TREITS. 3280” “cuspidana 3280” “Micropraep. upen. No. 1087 Kuznetsov,
1983“ “Lectotypus m Phoxopteris cuspidana Tr. design. Kuznetsov, 1983” (Hungarian Natural History Museum,
Budapest, Hungary).
The taxonomic confusion surrounding A. diminutana is detailed in the A. geminana species account. DNA
barcode data (Fig. 1) and consistent differences in wing pattern clearly demonstrate that the Nearctic A.
diminuatana is a separate species from A. diminutana.
Ancylis diminuatana Kearfott, 1905
Figs. 39–43, 44–46 (complex), 70–71, 84
Ancylis diminuatana Kearfott, 1905, Proc. U.S. natn. Mus. 28: 361. stat.rev.
Diagnosis. Ancylis diminuatana is distinguished by its “three-toned” forewing, which is pale tan on the costa,
reddish brown to dark brown in the remainder of the costal half, and gray in the dorsal half. In most specimens, the
shape of the longitudinal line is also distinctive, as it curves only slightly towards the cubitus before reaching CuA
2
(Figs. 39 – 41); however, in some specimens the longitudinal line curves abruptly to the cubitus and down to CuA
2
(Fig. 42). Other similar species (A. geminana, A. diminutana) appear two-toned, with contrasting costal and dorsal
halves of the forewing. Also, the longitudinal line in these other species is more sinuate. Because of the pale tan
costa, some specimens of A. goodelliana (A. unguicella group) appear similar to A. diminuatana, but the two
species have very different genitalia (Figs. 70–71, 84 versus Figs. 65, 79)
Redescription. Forewing. FWL ♂ 4.5–7.5 mm (n=44), ♀ 5.5–8 mm (n=16). The costa is pale tan in the basal
0.5–0.75 of the wing. The remainder of the costal half is brown to reddish brown, becoming darker at the
longitudinal line. The dorsal half is pale gray to brownish gray. The longitudinal line starts at A
1+2
at the base of the
wing, curves only slightly towards the cubitus before reaching CuA
2
, and then angles in a diagonal line to the apex.
In some specimens the longitudinal line curves abruptly to the cubitus and down to CuA
2
before angling to the
apex. Small black streaks are often present just proximal to where the longitudinal line intersects M
1
. Male
genitalia. As described for the group. Female genitalia. As in A. diminutana.
Molecular data. BIN URI: BOLD: AAA7190. The intraspecific divergence of the barcode region is moderate
with average 0.32% and maximum 1.28% (n=87). The minimum distance to the nearest neighbor A. subarcuana
(BIN AAB3492) is 3.53%.
Distribution. In Canada, A. diminuatana is distributed from Quebec west to Alberta and British Columbia. In
the U.S., A. diminuatana is distributed from New Hampshire and Massachusetts, south to North Carolina, west to
Nebraska, and possibly beyond. Kearfott (1905) included specimens from Colorado in his type series although we
have not located any specimens that we could confirm as A. diminuatana from that state. We did identify a single
female from California and specimens from Washington (discussed below) that we would confirm as this species.
Biology. Label data suggests that A. diminuatana is bivoltine in the East, with adults present May though early
July and again mid-August through September. Several specimens from the USNM appear to be reared from
willow (Salix spp.).
Remarks. Ancylis diminuatana has been consistently misidentified in North America, first as A. diminutana
and then as A. geminana. Specimens of A. diminuatana in BOLD consistently misidentified as A. carbonana (Fig.
1) demonstrate some of the dangers in relying on only DNA data for determinations—it is obvious from the online
photographs that none of these specimens is A. carbonana. The DNA data do clearly separate this group of
Nearctic A. diminuatana from the Palearctic A. diminutana.
We have identified several specimens from the American West that most closely match A. diminuatana but are
unlikely to be the same species. Figures 43–44 illustrate two specimens from Washington (Fig. 43, Everett; Fig. 44,
Yakima). The Everett specimen (and another from the same date and locality) appears to be a typical A.
diminuatana with slightly darker markings; however, the Yakima specimen, and a similar specimen from Walla
Zootaxa 4178 (3) © 2016 Magnolia Press
·
367
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Walla (not illustrated), appears more two-toned, and the longitudinal line is much more sinuate. Similarly, the
specimen in Fig. 45 from Silverton, Colorado, is nearly black and white with a sinuate longitudinal line. Two other
specimens (not illustrated) from Colorado match this phenotype. The specimen in Fig. 46 from Alaska is also
unusual with subdued markings and a sinuate longitudinal line. Other specimens (not illustrated) from the
Humboldt Mountains in northwest Nevada and Provo, Utah have wing patterns similar to those in Figs. 45–46.
Unfortunately, none of these specimens were collected recently enough to obtain quality DNA sequence data.
Thus, we hesitate to describe any additional species until additional diagnostic characters are discovered and
include all of these phenotypes as a potential species complex with A. diminuatana.
Ancylis saliana Gilligan and Huemer, sp.n.
Figs. 47–52, 72, 85
Type material. Holotype. ♂: “FL: Putnam Co., Palatka, At MV/UV light” “H. D. Baggett, 7-III-1992” (MEM).
Paratypes (11). United States: 2 ♂, 3 ♀ [same data as holotype], 18 February 1992 (1 ♀, MEM), 17 April 1992 (1
♂, MEM), 2 May 1992 (1 ♂, MEM), 12 May 1992 (1 ♀, MEM), 6 September 1991 (1 ♀, MEM); 1 ♂, Florida,
Collier Co., near Copeland, Fakahachee Strand, Jane’s Scenic Drive, 28 March 1986, leg. “DOW”, slide TMG676
(FSCA); 1 ♂, Florida, Collier Co., near Copeland, Fakahachee Strand, Jane’s Scenic Drive, 28 March 1986, leg. H.
D. Baggett (MEM); 2 ♀, Florida, Paradise Key, 4 March 1919, leg. Schwarz and Barber, slide TMG691 (USNM);
1 ♀, Florida, Royal Palm State Park, [date unreadable] 1930, leg. F. M. Jones, slide TMG629 (USNM); 1 ♀,
Florida, Volusia Co., Cassadaga, 29 January 1963, leg. S. V. Fuller (FSCA).
Diagnosis. Ancylis saliana is distinguished by the shape of the longitudinal line, which extends along A
1+2
before angling abruptly back towards the base of the wing, and then continuing down to CuA
2
and straight towards
the apex. This shape approximates a “wave” or “hook” in most specimens that is easy to distinguish from the
weakly sinuate line in A. diminuatana. In specimens of A. diminuatana where the longitudinal line curves more
abruptly to the costa (e.g., Fig. 42), A. saliana can be distinguished by the pale tan or brown coloration in the dorsal
half of the wing versus gray in A. diminuatana.
Description. Head, labial palpi and thorax tan. Second and third segment of labial palpi with patches of brown
scales. Forewing. FWL ♂ 5.9–6.8 mm (n=4), ♀ 6.5–7.7 mm (n=8). Forewing is falcate. The costal half of the wing
is a mix of light tan, brown, and yellowish brown. The costa is light tan becoming brown towards the apex with
remnants of costal strigulae. The dorsal half of the wing is pale tan with some brown markings. The longitudinal
line runs from the base of the wing along A
1+2
, angles abruptly back above the cubitus towards the base of the wing,
and then continues down to CuA
2
, and straight towards the apex. Small black streaks are often present on R
5
just
proximal to the termen, which has a row of dark brown scales. Some specimens (Fig. 52) have black scales
interspersed in the costal half of the wing along the margin of the longitudinal line and in the dorsal half of the
wing. Cilia at the apex dark brown with a white postapical strigula, other cilia are pale tan. Hindwing is pale
grayish brown. Male genitalia. As described for the group. Female genitalia. As described for the group with the
following modifications: apophyses posteriores 0.7 as long as apophyses anteriores; antrum sclerotized to the
posterior 0.20 of the ductus bursae; and ductus seminalis arising in the anterior 0.3 of the ductus bursae.
Molecular data. Unknown.
Distribution. Ancylis saliana is only known from few localities in Florida.
Biology. The majority of adults were collected from January to May. A single collection in September from the
type locality suggests two generations. The larval host is unknown.
Etymology. The species name is a variation of the Latin “sali” for “waves,” referring to the shape of the
longitudinal line in most specimens.
Remarks. Although it is somewhat troubling that we could find no genitalic differences from A. diminuatana,
we believe that the consistent shape of the longitudinal line and lack of gray in the dorsal half of the wing
sufficiently distinguishes A. saliana as a separate species.
GILLIGAN ET AL.
368
·
Zootaxa 4178 (3) © 2016 Magnolia Press
Ancylis subarcuana (Douglas, 1847)
Figs. 53–58, 73–74, 86
Anchylopera subarcuana Douglas, 1847, Trans. ent. Soc. Lond. 5: 21. stat.rev.
inornatana [uninomial] Herrich-Schäffer, 1848, Syst. Bearbeitung Schmett. Eur. 4: pl. 43, fig. 206. [nomen nudum]
Tortrix (Phoxopteryx) inornatana Herrich-Schäffer, 1851, Syst. Bearbeitung Schmett. Eur. 4: 287.
Diagnosis. Ancylis subarcuana is distinguished by its grayish forewing and subdued markings. The costal and
dorsal halves of the forewing are not contrasting, and the expression of the longitudinal line is obscured at the base
and apex of the wing. The longitudinal line is continuous from the base of the wing to the apex or termen in A.
geminana, A. christiandiana, and A. diminutana, and the costal and dorsal halves of the forewing are contrasting to
nearly two-toned in those species. Some individuals are dark brown with reduced markings (Fig. 58).
Redescription. Forewing. FWL ♂ 6–8.5 mm (n=15), ♀ 6–8 mm (n=10). The costal half of the wing is pale
grayish tan at the costa, becoming brown towards the longitudinal line. The dorsal half of the wing is light
brownish gray. The longitudinal line arises from below a dark brown to black mark near A
1+2
, runs up to the
cubitus, down to CuA
2
beneath another dark brown to black mark, and then up towards the apex, becoming
obscured again before reaching the termen. A small black dash is often present just proximal to where the
longitudinal line intersects M
1
. The male in Fig. 58 from northern Finland is typical of specimens from that region
that are dark brown with only a remnant of the longitudinal line. Male genitalia. As described for the group.
Female genitalia. As in A. diminutana.
Molecular data. BIN URI: BOLD:AAB3492, BOLDABX6097. The intraspecific divergence of the barcode
region is moderate when considering both BINs, with average 0.72% and maximum 2.34% (n=9). Within BIN
BOLD:AAB3492 average distance is only 0.19% with maximum 0.76%. The minimum distance of this cluster to
the nearest neighbor BIN BOLDABX6097 is 2.25%.
Distribution. Razowski (2001, 2003) reported A. subarcuana as distributed from the United Kingdom through
Northern Europe to the Baltic States and parts of central Europe, and according to Sinev (2008) also in the north-
western part of European Russia. We also examined specimens from northern Italy.
Biology. Adults are on the wing in two generations, from April to May and from July to August (Razowski
2001). The larval host is Salix repens (Razowski 2003) and, according to Wegner (2015), also Salix aurita. In
northern Europe the species prefers dunes, heathland and marshy pinewood, in southern parts it was found in
alluvial river zones.
Remarks. Anchylopera subarcuana was described from a single specimen collected on May 12, 1844 near
Wimbledon (England, GB). According to Sattler (in litt.) no type material could be found in the collections of the
Natural History Museum (London, UK). However, the original description with attached color figure leaves no
doubt of the identity.
DNA barcode data (Fig. 1) and consistent differences in wing pattern clearly separate A. subarcuana from A.
geminana. Species status is furthermore supported by unique larval characters as described already by Schütze
(1931) but largely neglected in subsequent literature (e.g. Bradley et al. 1979). The taxonomic confusion
surrounding species in this group is detailed under the A. geminana account.
Acknowledgements
We are particularly grateful to Paul Hebert and his team at the Canadian Centre for DNA Barcoding (Guelph,
Canada) who’s sequencing work was enabled by funding from the Government of Canada to Genome Canada
through the Ontario Genomics Institute. We are also grateful to the Ontario Ministry of Research and Innovation
and to NSERC for their support of the BOLD informatics platform. PH is furthermore indebted to the Promotion of
Educational Policies, University and Research Department of the Autonomous Province of Bolzano—South Tyrol
for helping to fund the project “Genetic biodiversity archive—DNA barcoding of Lepidoptera of the central Alpine
region (South, East and North Tyrol)”, and to the Austrian Federal Ministry of Science, Research and Economics
for funds received in the framework of ABOL (Austrian Barcode of Life).
We are grateful to the following persons for providing access to or loans of material under their care, and/or
access to hitherto unpublished DNA barcode data: John Brown (USNM); Helmut Deutsch (Bannberg, Austria);
Zootaxa 4178 (3) © 2016 Magnolia Press
·
369
RESOLVING TAXONOMY IN HOLARCTIC ANCYLIS
Jason Dombroskie (CIUC); David Grimaldi and Suzanne Rab Green (AMNH); James Hayden (FSCA); Jean-
François Landry (CNC); Marko Mutanen (Oulu); Paul Opler and Boris Kondratieff (CSU); Jerry Powell and Peter
Oboyski (EME); Andreas Segerer (ZSM); and Christian Wieser (LMK). Laszlo Ronkay (Budapest, Hungary)
kindly helped us with photographs of P. cuspidana and Klaus Sattler (London, UK) with information on potential
type material at the Natural History Museum. Jean-François Landry (CNC) provided the photograph illustrated in
Fig. 42. Joaquín Baixeras Almela and Richard L. Brown provided helpful review comments that greatly improved
the quality of the manuscript.
Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the products by
the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that may also
be suitable.
References
Aarvik, L.E. (2013) Fauna Europaea: Tortricidae. In: Karsholt, O. & Nieukerken, E.J. van (Eds.), Fauna Europaea:
Lepidoptera, Moths. Fauna Europaea version 2.6. Available from: http://www.fauna-eu.org (Accessed 25 Oct. 2016)
Arnqvist, G. (1997) The evolution of animal genitalia: distinguishing between hypotheses by single species studies. Biological
Journal of the Linnean Society, 60, 365–379.
http://dx.doi.org/10.1111/j.1095-8312.1997.tb01501.x
Barnes, W. & McDunnough, J. (1917) Checklist of the Lepidoptera of Boreal America. Decatur, Illinois. 200 pp.
Bradley, J.D., Tremewan, W.G. & Smith, A. (1979) British Tortricoid Moths. Tortricidae: Olethreutinae. The Ray Society,
London. Viii + 336 pp, pls 22–43.
Brown, J.W. (2005) World catalogue of insects. Volume 5: Tortricidae (Lepidoptera). Apollo Books, Stenstrup, Denmark. 741
pp.
Brown, J.W. & Powell, J.A. (1991) Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini).
University of California Publications in Entomology, 111, 87 pp.
Brown, J.W., Janzen, D., Hallwachs, W., Zahiri, R., Hajibabaei, M. & Hebert, P.N.D. (2014) Cracking complex taxonomy of
Costa Rican moths: Anacrusis Zeller (Lepidoptera: Tortricidae). Journal of the Lepidopterists’ Society, 68, 248–263.
http://dx.doi.org/10.18473/lepi.v68i4.a3
deWaard, J.R., Ivanova, N.V., Hajibabaei, M. & Hebert, P.D.N. (2008) Assembling DNA Barcodes: Analytical Protocols. In:
Cristofre, M, (Ed.), Methods in Molecular Biology: Environmental Genetics. Humana Press Inc., Totowa, USA. pp. 275–
293.
http://dx.doi.org/10.1007/978-1-59745-548-0_15
Donovan, E. (1806) The Natural History of British Insects: Explaining them in their several states, with the periods of their
transformations, their food, oeconomy, etc. Together with the history of such minute insects as require investigation by the
microscope: the whole illustrated by coloured figures, designed and executed from living specimens. Volume 11. pp. 361–
396.
Eberhard, W.G. (1985) Sexual selection and animal genitalia. Harvard University Press, Cambridge, Massachusetts. 244 pp.
http://dx.doi.org/10.4159/harvard.9780674330702
Fernald, C.H. (1882) A synonymical catalogue of the described Tortricidae of North America north of Mexico. Transactions of
the American Entomological Society, 10, 1–64.
Fernald, C.H. (1903 [1902]) Family Tortricidae. In: Dyar, H.G. (Ed.), A list of North American Lepidoptera. Bulletin of the
United States National Museum, No. 52. pp. 448–489.
Gilligan, T.M. & Wenzel, J.W. (2008) Extreme intraspecific variation in Hystrichophora (Lepidoptera: Tortricidae) genitalia —
questioning the lock-and-key hypothesis. Annales Zoologici Fennici, 45, 465–477.
http://dx.doi.org/10.5735/086.045.0601
Gilligan, T.M., Wright, D.J. & Gibson, L.D. (2008) Olethreutine moths of the midwestern United States, an identification guide.
Bulletin of the Ohio Biological Survey, Volume 16, 334 pp.
Gilligan, T.M., Baixeras, J., Brown, J.W. & Tuck, K.R. (2014a) T@RTS: Online World Catalogue of the Tortricidae (Ver. 3.0).
Available from: http://www.tortricid.net/catalogue.asp (Accessed 25 Oct. 2016)
Gilligan, T.M., Wright, D.J., Munz, J., Yakobson, K. & Simmons, M.P. (2014b) Molecular phylogeny and revised classification
of Eucosma Hübner and related genera (Lepidoptera: Tortricidae: Eucosmini). Systematic Entomology, 39, 49–67.
http://dx.doi.org/10.1111/syen.12036
Hebert, P.D.N., Cywinska, A., Ball, S.L. & de Waard, J.R. (2003) Biological identifications through DNA barcodes.
Proceedings of the Royal Society of London, B, 270, 313–321.
http://dx.doi.org/10.1098/rspb.2002.2218
Heinrich, C. (1923) Revision of the North American moths of the subfamily Eucosminae of the family Olethreutidae. Bulletin
of the United States National Museum, 123, 1–128.
Heinrich, C. (1926) Revision of the North American moths of the subfamilies Laspeyresiinae and Olethreutinae. Bulletin of the
United States National Museum, 132, 1–216.
http://dx.doi.org/10.5479/si.03629236.132.1
Horak, M. (1984) Assessment of taxonomically significant structures in the Tortricinae (Lep.: Tortricidae). Mitteilungen der
GILLIGAN ET AL.
370
·
Zootaxa 4178 (3) © 2016 Magnolia Press
schweizerischen entomologischen Gesellschaft, 57, 3–64.
Horak, M. (1999) The Tortricoidea, In: Kristensen, N.P. (Ed.), Lepidoptera: Moths and Butterflies. Volume 1: Evolution,
systematics, and biogeography. Handbook of Zoology Vol. IV, Part 35. Walter de Gruyter, Berlin and New York. pp. 199–215.
Horak, M. (2006) Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, 10, 1–
522.
Hosken, D.J. & Stockley, P. (2004) Sexual selection and genital evolution. Trends in Ecology & Evolution, 19, 87–93.
http://dx.doi.org/10.1016/j.tree.2003.11.012
Humble, L.M., deWaard, J.R. & Quinn, M. (2009) Delayed recognition of the European poplar shoot borer, Gypsonoma
aceriana (Duponchel) (Lepidoptera: Tortricidae) in Canada. Journal of the Entomological Society of British Columbia,
106, 61–70.
Kearfott, W.D. (1905) Descriptions of new species of tortricid moths from North Carolina, with notes. Proceedings of the
United States National Museum, 28, 349–364.
http://dx.doi.org/10.5479/si.00963801.1398.349
Klots, A.B. (1970) Lepidoptera. In: Tuxen, S.L. (Ed), Taxonomic Glossary of Genitalia in Insects. Munksgaard, Copenhagen.
pp. 115–130.
Landry, J.F., Nazari, V., deWaard, J.R., Mutanen, M., Lopez-Vaamonde, C., Huemer, P. & Hebert, P.D.N. (2013) Shared but
overlooked: 30 species of Holarctic Microlepidoptera revealed by DNA barcodes and morphology. Zootaxa, 3749 (1), 1–
93.
http://dx.doi.org/10.11646/zootaxa.3749.1.1
Mikkola, K. (1992) Evidence for lock-and-key mechanisms in the internal genitalia of the Apamea moths (Lepidoptera,
Noctuidae). Systematic Entomology, 17, 145–153.
http://dx.doi.org/10.1111/j.1365-3113.1992.tb00327.x
Miller, W.E. (1987) Guide to the olethreutine moths of Midland North America (Tortricidae). United States Department of
Agriculture. Forest Service Agriculture Handbook, 660. 104 pp.
Mutanen, M., Rytkonen, S., Linden, J. & Sinkkonen, J. (2007) Male genitalia variation in a moth Pammene luedersiana
(Lepidoptera: Tortricidae). European Journal of Entomology, 104, 259–265.
http://dx.doi.org/10.14411/eje.2007.040
Mutanen, M., Aarvik, L. Landry, J.-F., Segerer, A.H. & Karsholt, O. (2012) Epinotia cinereana (Haworth, 1811) bona sp., a
Holarctic tortricid distinct from E. nisella (Clerck, 1759) (Lepidoptera: Tortricidae: Eucosmini) as evidenced by DNA
barcodes, morphology and life history. Zootaxa, 3318, 1–25.
Pierce, F.N. & Metcalfe, J.W. (1922) The genitalia of the group Tortricidae of the Lepidoptera of the British Islands. Oundle,
Liverpool, England. 101 pp.
Powell, J.A. (1983) Tortricidae, In: Hodges, R.W. (Ed.), Check list of the Lepidoptera of America north of Mexico. E.W.
Classey & Wedge Entomological Research Foundation, London, pp. 31–41.
Powell, J.A. & Opler, P.A. (2009) Moths of Western North America. University of California Press. Berkeley, Los Angeles,
London. 369 pp.
http://dx.doi.org/10.1525/california/9780520251977.001.0001
Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: The Barcode of Life Data System (http://www.barcodinglife.org).
Molecular Ecology Notes, 7, 355–364.
http://dx.doi.org/10.1111/j.1471-8286.2007.01678.x
Razowski, J. (1989) The genera of Tortricidae (Lepidoptera). Part II: Palaearctic Olethreutinae. Acta Zoologica Cracoviensia,
32, 107–328.
Razowski, J. (1996) Tortricidae, In: Karsholt, O. & Razowski, J. (Eds.), The Lepidoptera of Europe. A distributional checklist.
Apollo Books, Stenstrup. pp. 130–157, 313–318.
Razowski, J. (2001) Die Tortriciden (Lepidoptera, Tortricidae) Mitteleuropas. Bestimmung – Verbreitung – Flugstandort –
Lebensweise der Raupen. František Slamka, Bratislava. 301 pp.
Razowski, J. (2003) Tortricidae of Europe, Volume 2, Olethreutinae. František Slamka, Bratislava. 301 pp.
Rebel, H. (1901) Catalog der Lepidopteren des palaearctischen Faunengebietes. II. Theil: Famil. Pyralidae-Micropterigidae.
R. Friedländer & Sohn, Berlin. 368 pp.
Schütze, K.T. (1931) Die Biologie der Kleinschmetterlinge unter besonderer Berücksichtigung ihrer Nährpflanzen und
Erscheinungszeiten. Internationaler Entomologischer Verein, Frankfurt. 235 pp.
Sinev, S.Yu. (Ed.) (2008) Catalogue of the Lepidoptera of Russia. KMK Scientific Press, St. Petersburg, Moscow. 424 pp.
Stephens, J.F. (1829) A systematic catalogue of British insects: being an attempt to arrange all the hitherto discovered
indigenous insects in accordance with their natural affinities. London: Published for the author, by Baldwin and Cradock.
852 pp.
http://dx.doi.org/10.5962/bhl.title.8987
Walsingham, L.T. de G. (1897) Revision of the West-Indian Microlepidoptera with descriptions of new species. Proceedings of
the Entomological Society of London, 1897, 54–183.
http://dx.doi.org/10.5962/bhl.title.53759
Wegner, H. (2015) Ein Beitrag zur Wickler-Fauna in Nordost-Niedersachsen und in Schleswig-Holstein (Lep., Tortricidae).
Melanargia, 27 (4), 137–154.
Wocke, M. (1871) In: Staudinger, O. & Wocke, M. (1871), Catalog der Lepidopteren des europäischen Faunengebiets.
Dresden. 426 pp.
Wright, D.J. & Gilligan, T.M. (2015) Eucosma Hübner of the Contiguous United States and Canada (Lepidoptera: Tortricidae:
Eucosmini). Wedge Entomological Research Foundation, Alamogordo, New Mexico, 256 pp.
... In accordance with the patterns described for other Lepidoptera families (Hebert et al., 2003), intraspecific DNA barcode divergences are in general lower than 2% in Tortricidae (Hulcr et al., 2007;Gilligan et al., 2016;Corley and Ferreira, 2017;Vargas-Ortiz et al., 2017), although cases of greater divergence have been described for a few widespread species (Gilligan et al., 2016). The deep divergence of the BOLD sequences of B. verutana and the results of the phylogenetic analysis, in which the haplotypes from Madagascar and Costa Rica were not clustered with those from USA, strongly suggest that more than one species is present under this name in this database. ...
... In accordance with the patterns described for other Lepidoptera families (Hebert et al., 2003), intraspecific DNA barcode divergences are in general lower than 2% in Tortricidae (Hulcr et al., 2007;Gilligan et al., 2016;Corley and Ferreira, 2017;Vargas-Ortiz et al., 2017), although cases of greater divergence have been described for a few widespread species (Gilligan et al., 2016). The deep divergence of the BOLD sequences of B. verutana and the results of the phylogenetic analysis, in which the haplotypes from Madagascar and Costa Rica were not clustered with those from USA, strongly suggest that more than one species is present under this name in this database. ...
... It is supposed that the haplotypes from North America represent the true B. verutana, as this group includes sequences sampled close to the type locality (Texas, USA). The divergence between the North American haplotypes of B. verutana and the two haplotypes of B. blepharopis (1.7-2.0%) is close to the highest values of intraspecific divergence reported for a few widespread tortricids (Gilligan et al., 2016); however, the two are currently considered valid species based on morphology (Horak, 2006;Gilligan et al., 2014). In addition, the reciprocal monophyly found in the phylogenetic analysis between the North American haplotypes of B. verutana and those of B. blepharopis reinforces their heterospecific status. ...
Article
Full-text available
The sedge-feeding moth Bactra verutana Zeller, 1875 (Lepidoptera: Tortricidae: Olethreutinae: Bactrini), described from Dallas, Texas, USA, is widespread, recorded throughout much North America, Central and South America, including the Caribbean, and Africa. The species is recorded for the first time from Chile based on specimens collected in the coastal valleys of the Atacama Desert, where its larvae feed on Cyperus corymbosus Rottb. var. subnodosus (Nees & Meyen) Kük. (Cyperaceae). A single DNA barcode haplotype, which is widespread in USA, was found in two Chilean specimens sequenced.
... 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]. ...
... 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. ...
Article
Full-text available
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.
... Molecular analysis. DNA barcodes have been widely recognized as useful tools to assess taxonomic problems as a complement to morphology in Lepidoptera, with several examples in Tortricidae , Gilligan et al. 2016, Razowski et al. 2016, Escobar-Suárez et al. 2017). According to our Bayesian analysis, S. gattii is strongly supported as a monophyletic species. ...
... The surveys should also include the Peruvian range of M. pavonis (Luebert 2004). As already reported in previous studies with Tortricidae (Gilligan et al. 2016;Razowski et al. 2016;Escobar-Suárez et al. 2017), the divergence found between the two haplotypes of S. gattii suggests that analysis of DNA barcode sequences would be useful to assess the taxonomic status of eventual new populations of this micromoth. ...
Article
The adult, larva, and pupa of Strepsicrates gattii Vargas-Ortiz & Vargas, sp. n. (Lepidoptera: Tortricidae: Olethreutinae: Eucosmini), are described and illustrated from the Atacama Desert of northern Chile. The larvae are leaf-tiers on the vulnerable native tree Morella pavonis (Myricaceae). As S. gattii was previously misidentified as S. smithiana Walsingham, morphological differences that enable the separation of the two species are highlighted. Sequences of the DNA barcode fragment of the cytochrome oxidase subunit I mitochondrial gene of the new species are provided and used in a Bayesian analysis with congeneric representatives to assess their relationships preliminarily. The divergence (K2P) with S. smithiana was 6.4–7.4%, providing additional support for separating the two species.
... Although Miller and Pogue (1984) provided a convincing case for a continuous gradient of morphological characters between E. strenuana and E. minutana, there do seem to be diagnosable differences (e.g., wing coloration) in the majority of specimens that may indicate these are different species. Molecular data, including DNA barcoding (Hebert et al. 2003), has been used successfully to resolve taxonomic issues where morphological characters are ambiguous and to determine which morphological characters are taxonomically informative (e.g., Brown et al. 2014, Gilligan et al. 2014, Gilligan et al. 2016). Here we use DNA barcoding combined with morphology and host preference to examine populations of the E. strenuana complex from North America, Australia, China, and Israel. ...
Article
Full-text available
The ragweed borer, Epiblema strenuana (Walker, 1863), has a long history of use as a biological control agent against important weed pests in the family Asteraceae. Recently, E. strenuana has been reported feeding on the invasive perennials Ambrosia confertiflora and A. tenuifolia in Israel. The geographic location of Israel has raised concern over the possibility that the moth may spread to areas such as Ethiopia where the oil-seed crop Guizotia abyssinica is cultivated, as this is a potential host for E. strenuana. However, the taxonomic status of E. strenuana and a current synonym, E. minutana (Kearfott, 1905) is unclear. These taxa have been treated as separate species in the past, and they potentially have different feeding habits and damage different parts of the plant. We analyzed DNA data and adult morphology and determined that E. minutana, stat. rev., is a valid species which we raise from synonymy with E. strenuana. Wing coloration, the shape of the female sterigma, and COI DNA barcodes are consistently different between the two species. We also determined that the species previously identified as E. strenuana in Israel is actually E. minutana. While detailed host range tests have been conducted on the E. strenuana populations released in Australia and China, the host range of E. minutana remains to be clarified. We discuss the history of biological control using E. strenuana and the implications for finding E. minutana in Israel. We also provide species redescriptions for E. strenuana and E. minutana and illustrate diagnostic characters.
... Alternative techniques, such as DNA barcoding (Hebert et al. 2003), provide an additional data set with which to test species boundaries and congruence of morphological and/or host data. Molecular data have been successfully used to solve taxonomic problems in Tortricidae in numerous studies (e.g., Mutanen et al. 2012;Brown et al. 2014;Gilligan et al. 2014Gilligan et al. , 2016. The most comprehensive DNA barcoding database is hosted by the Barcode of Life Data System (BOLD; Ratnasingham and Hebert 2007), and it currently (March 2018) contains sequence data for 282 specimens of Paralobesia representing a reputed 26 species. ...
Article
Full-text available
The genus Paralobesia Obraztsov, 1953 is found primarily in eastern North America and consists of 18 described and several undescribed species. Prior to 1900, all North American Paralobesia were assumed to be P. viteana (Clemens). However, rearing experiments by William Kearfott in the early 1900s suggested that species of Paralobesia were monophagous and could be separated by host. Recently, a species of Paralobesia was reared from showy lady’s slipper, Cypripedium reginae Walter (Orchidaceae), during a study of two populations of this orchid in eastern Ontario and southwestern Québec. Although originally assumed to be P. cypripediana (Forbes), which was described from specimens reared from Cypripedium in Manitoba, DNA barcode data and genital morphology confirmed that this was a new species similar to P. cypripediana and P. monotropana (Heinrich). Herein, we describe P. marilynae, sp. n., and provide specifics of its discovery and life history. Rearing records indicate that Paralobesia can span the range from strictly monophagous to polyphagous, even for very similar species with similar feeding habits, and that host records should be combined with morphological and molecular data when circumscribing species in this genus. This work is part of a complete systematic revision of Paralobesia currently in progress.
... Over recent years, several overlooked cryptic species have been detected (e.g. Huemer and Hausmann 2009;Huemer and Hebert 2011;Huemer et al. 2013;Huemer et al. 2014aHuemer et al. , 2014cHuemer and Timossi 2014;Buchner 2015;Huemer and Mutanen 2015;Kirichenko et al. 2015;Baldizzone and Landry 2016;Gilligan et al. 2016;Kozlov et al. 2016). In this paper we reveal a striking cryptic diversity in the taxonomically difficult genus Agrotis Ochsenheimer, 1816. ...
Article
Full-text available
An integrative taxonomic analysis of the European species of the Agrotis fatidica species-group is presented with special reference to the European sister taxa of A. fatidica (Hübner, 1824); in addition, a general overview of the entire species-group is given. The remarkable differences found in the barcodes of the Central and Western European populations of A. fatidica (sensu lato) led us to recognise isolated species of the A. fatidica complex. Two new species, A. mayrorum sp. n. (Northern Italy and the French Alps) and A. mazeli sp. n. (French Pyrenees) are described. The neotype of A. fatidica is designated. Agrotis luehri von Mentzer & Moberg, 1987 is treated as a subspecies of A. fatidica (stat. n.).
... Thus, as a first step to understand the genetic relationships between individuals of isolated locations, it is important to gain insights about the patterns of maternally inherited genetic variation, which can be performed through sequence analysis of mitochondrial genes (Sn€ all et al. 2004;Moreira et al. 2017). Besides being a helpful tool to perform accurate taxonomic identifications of micromoths (Landry and Hebert 2013;Kirichenko et al. 2015;Gilligan et al. 2016;Pereira et al. 2017), the DNA barcode fragment (sensu Hebert et al. 2003) of the cytochrome oxidase subunit I has been used successfully to assess population genetic variation in insects, either alone or in complement with other mitochondrial or nuclear markers (Velasco-Cuervo et al. 2016;Frantine-Silva et al. 2017;Gallo-Franco et al. 2017;Landvik et al. 2017;Pfeiler et al. 2017;Zhang et al. 2017), including several examples with micromoths (Shapiro et al. 2008;Valade et al. 2009;D ıaz-Montilla et al. 2013;Maia et al. 2016;Kirichenko et al. 2017). The aim of this study was to analyze DNA barcode sequences to assess for the first time the patterns of genetic variation of B. mirnae from one highland and two lowland localities of the arid northernmost part of Chile. ...
Article
Analysis of maternally inherited genes is especially helpful in population studies of host-specialized insects, as female dispersal is key to find an adequate host plant to ensure larval survival. Bucculatrix mirnae (Lepidoptera: Bucculatricidae) is a little-known Neotropical micromoth native to the arid environments of northern Chile whose hypermetamorphic larvae are miners and skeletonizers on leaves of two species of Baccharis (Asteraceae) shrubs. This micromoth has been detected in three isolated locations embracing a narrow geographic range: two from the coastal valleys of the Atacama Desert near sea level and one from the western slopes of the Andes at about 3000 m elevation. As the dispersal of B. mirnae is mostly restricted to the small adult stage, the altitudinal gradient and desert areas among the three localities could be effective barriers, triggering genetic differentiation among populations. Sequences of the DNA barcode fragment of the cytocrome oxidase subunit I mitochondrial gene were analyzed to assess for the first time the patterns of genetic variation of B. mirnae. Fifteen haplotypes, each exclusive to one locality, were found in the 71 specimens analyzed. Genetic divergence (K2P) between haplotypes of different localities was at least 2.0%. A Bayesian analysis with sequences of congeneric species grouped all the B. mirnae haplotypes in a clade, in which three well-supported locality-specific haplogroups were found. In concordance with this pattern, an analysis of molecular variance showed that the highest genetic variation was found among populations. Furthermore, all the population pairwise comparisons (FST) were significant. These results suggest that female migration between isolated populations of B. mirnae is absent. This pattern must be considered in the current scenario of habitat destruction and modification in the arid environments of northern Chile.
... DNA barcodes are useful for exploring biodiversity and taxonomy, especially in concert with other character sources , Gilligan et al. 2016, Escobar-Suárez et al. 2017, Razowski et al. 2017. Barcodes also can be used to identify immature stages of insects, including Lepidoptera, providing knowledge of their trophic interactions when rearing larvae to obtain the adults is difficult or impossible (Gossner & Hausmann 2009, Hausmann & Parra 2009, Frye & Robbins 2015. ...
Article
Full-text available
Originally described from Africa, the genus Eccopsis Zeller (Lepidoptera: Tortricidae) currently includes 25 Afrotropical and five Neotropical species. Adult morphological characters suggest that the Afrotropical and Neotropical species might not be congeneric. Here we present the first DNA sequences for Neotropical Eccopsis and use these data in a maximum likelihood (ML) analysis to evaluate the monophyly of the genus, and to examine the utility of DNA barcodes in separating the South American E. galapagana Razowski & Landry, 2008 and E. razowskii Vargas, 2011. Intraspecific and interspecific pairwise distances (K2P) were 0-0.5% and 4.9-5.2%, respectively, and each species was recovered as a distinct, well supported group of sequences (i.e., species) in the ML analysis. An analysis including barcodes of Afrotropical Eccopsis (four species), Afrotropical Paraeccopsis (one species), and Neotropical Eccopsis (two species) failed to recover Eccopsis as monophyletic. Consistent with previous suggestions based on adult morphology, this study highlights the necessity to reassess the congeneric status of Afrotropical and Neotropical species of Eccopsis.
Article
Full-text available
A new species from the Canary Islands is described in the genus Ancylis Hübner, 1825 (Lepidoptera: Tortricidae, Enarmoniini), Ancylis klimeschana spec. n. This new species differs from all other species in the genus by the structure of the wing pattern and in the female genitalia especially the shape of the sterigma. The new species is compared with Ancylis comptana (Frölich, 1828) and A. sparulana (Staudinger, 1859). A. comptana is reported from the Balearic Islands for the first time.
Book
Full-text available
A definitive species list is the foundation of biodiversity and conservation work. As we deal with massive climatic changes in the Anthropocene, knowing which species make up our diverse ecosystems will be critically important if we wish to protect and restore them. The Lepidoptera, moths and butterflies, are the fourth-largest insect order in terms of global diversity, with approximately 158,000 described species. Here we report the distributions of 5431 species that occur in Canada and Alaska, as well as 53 species that have been reported from the region but not yet verified. Additionally, 19 species are listed as interceptions or unsuccessful introductions, and 52 species are listed as probably occurring in the region. The list is based on records from taxonomic papers, historical regional checklists, and specimen data from collections and online databases. All valid species and their synonyms, and all Nearctic subspecies and synonyms are included, except for butterfly subspecies (and their synonyms) that have never been reported from the region. The list is presented in taxonomic order, with the author, date of description, and original genus provided for each name.
Article
Full-text available
Remarkably similar forewing patterns, striking sexual dimorphism, and rampant sympatry combine to present a taxonomically and morphologically bewildering complex of five species of Anacrusis tortricid moths in Central America: Anacrusis turrialbae Razowski, Anacrusis piriferana (Zeiler), Anacrusis terrimccarthijae, n. sp., Anacrusis nephrodes (Walsingham), and Anacrusis ellensatterleeae, n. sp. Morphology and DNA barcodes (i.e., the mitochondrial gene COI) corroborate the integrity of the five species, all of which have been reared from caterpillars in Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. These species are polyphagous, with larval foodplants spanning many families of flowering plants. In ACG they occupy different forest types that are correlated with elevation.
Article
Full-text available
Although much biological research depends upon species diagnoses, taxonomic expertise is collapsing. We are convinced that the sole prospect for a sustainable identification capability lies in the construction of systems that employ DNA sequences as taxon 'barcodes'. We establish that the mitochondrial gene cytochrome c oxidase I (COI) can serve as the core of a global bioidentification system for animals. First, we demonstrate that COI profiles, derived from the low-density sampling of higher taxonomic categories, ordinarily assign newly analysed taxa to the appropriate phylum or order. Second, we demonstrate that species-level assignments can be obtained by creating comprehensive COI profiles. A model COI profile, based upon the analysis of a single individual from each of 200 closely allied species of lepidopterans, was 100% successful in correctly identifying subsequent specimens. When fully developed, a COI identification system will provide a reliable, cost-effective and accessible solution to the current problem of species identification. Its assembly will also generate important new insights into the diversification of life and the rules of molecular evolution.
Book
Olethreutine moths often have fruit-boring larvae and this economically important group includes many horticultural pests such as codling moths, Oriental fruit moths and macadamia nut borers. This volume is the first reference to describe the 90 olethreutine genera present in Australia. It provides generic definitions, a key to genera, generic descriptions, and illustrations of adults, heads, venation, genitalia of both sexes and other diagnostic structures of all genera. Summaries of biology and distribution and a checklist for all named Australian species are given for each genus. Importantly, it includes a comprehensive reorganisation of olethreutine classification, based on generic revisions, with a worldwide impact. The volume contains copious illustrations (two species per genus where possible) to convey generic concepts, and to allow identification of this economically important group. Nearly all olethreutine genera present in Australia extend into Asia and beyond, so the book will be relevant to horticultural pests throughout Asia, and crucial to an understanding of olethreutine evolution worldwide. The diverse Australian olethreutine fauna is particularly rich in enarmoniine and grapholitine genera, several new to science and adding significantly to the concepts of these two tribes. Given the wealth of biological information, the book will be important for ecological work on phytophagous insects well beyond Australia.
Article
Redescribes 124 genera, with data and comments on phylogeny and distribution. -from Author
Article
Insects boast incredible diversity, and this book treats an important component of the western insect biota that has not been summarized before-moths and their plant relationships. There are about 8,000 named species of moths in our region, and although most are unnoticed by the public, many attract attention when their larvae create economic damage: eating holes in woolens, infesting stored foods, boring into apples, damaging crops and garden plants, or defoliating forests. In contrast to previous North American moth books, this volume discusses and illustrates about 25% of the species in every family, including the tiny species, making this the most comprehensive volume in its field. With this approach it provides access to microlepidoptera study for biologists as well as amateur collectors. About 2,500 species are described and illustrated, including virtually all moths of economic importance, summarizing their morphology, taxonomy, adult behavior, larval biology, and life cycles.