Zootaxa 3779 (3): 375–382 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3779.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:13A26F5C-4675-45DE-920F-53739FB1496D
Review of the Olethreutes bowmanana-magadana species group, with the description of a new species from the Amur River, Russia (Lepidoptera, Tortricidae) VLADIMIR V. DUBATOLOV1, ANNA A. BOGUNOVA (SYACHINA)2 & SVETLANA V. NEDOSHIVINA3 1 Siberian Zoological Museum, Institute of Systematics and Ecology of Animals, Siberian Branch of Russian Academy of Sciences, Frunze street 11, RF-630091 Novosibirsk, Russia. E-mail: [email protected] 2 Biology Department, Amursky Liberal-Pedagogical State University, Kirov Street, 17/2, 681000 Komsomolsk-na-Amure, Russia. E-mail: [email protected] 3 Department of Zoology, Ulyanovsk State Pedagogical University, pl. 100-letiya Lenina 4, RF-432700 Ulyanovsk, Russia. E-mail: [email protected]
Abstract We define and review the Olethreutes bowmanana-magadana species group comprised of Olethreutes bowmanana (McDunnough), O. magadana (Falkovitsh), O. kamtshadala (Falkovitsh), and O. pivanica, sp. n., the last from petrophytous sunny slopes in the Amur River. The new species differs from other species of the group by a narrow, triangular-shaped apical process from the sacculus; a round, wide, apical bulb from the cucullus; and a longer, narrow neck of the valva. Key words: Tortricidae, Olethreutes bowmanana, Olethreutes magadana, Olethreutes kamtshadala, new species, Amur River, Russia, Far East
Introduction During a study of the lepidopterous fauna of the Lower Amur Region, Russia, a new species of Olethreutes Hübner [1822] was discovered. The moths were collected on a petrophytous sunny slope of a terrace on the Amur River bank opposite to Komsomolsk-na-Amure city. The specimens belong to a group of closely related species including O. magadana (Falkovitsh 1965) and O. kamtshadala (Falkovitsh 1966), but they differ noticeably from both of them. A comparison of the specimens with types and additional material in the Zoological Institute (St. Petersburg, Russia) revealed that some specimens in the collection of Zoological Institute were misidentified. This finding prompted us to prepare a review of the species of this group, including the description of the new species.
Material and methods Dissection methods follow those of Clarke (1941) and Hardwick (1950). Terminology for morphological features of wings and genitalia follow Gilligan et al. (2008). The following depositories are abbreviated in the text: CNCO ISEA ZISP
Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Canada; Siberian Zoological Museum, Institute of Systematics and Ecology of Animals, Novosibirsk, Russia; Zoological Institute of Russian Academy of Sciences, St. Petersburg, Russia.
Accepted by J. Brown: 31 Jan. 2014; published: 17 Mar. 2014
375
Results Olethreutes Hübner [1822] Systematisch-alphabetisches Verzeichniss aller bisher bey den Fürbildungen zur Sammlung europäischer Schmetterlinge: 72. Type species: Phalaena Tortrix arcuana Linnaeus, 1761 = [Phalaena] arcuella Clerck, 1759 by subsequent designation.
Olethreutes is a large Holarctic genus with 133 described species currently assigned to it (Gilligan et al. 2012). Species included in the genus exhibit a wide range of genital morphology wing patterns, and male secondary structures, making the genus extremely heterogeneous and probably polyphyletic. However, many species within Olethreutes show remarkable similarities to presumably closely related species, and many of these could be combined into natural groups. The Olethreutes bowmanana-magadana species group, proposed here, shows clear differences from other Olethreutes and is considered a natural group supported by following characters: brown to dark gray wing colour with lighter grayish striae and blackish spots; uncus narrow, bifid distally; sacculus with a distal internal lobe covered by strong spines and a spine cluster proximally on the ventral side; cucullus with a slender neck and apical widening; and aedeagus with foldable apical spine. The assignment of the species under consideration to the genus Olethreutes is generally accepted (Brown et al. 2005), but their relationships to other Olethreutes species remain uncertain.
Olethreutes bowmanana (McDunnough 1923) (Fig. 1) Argyroploce bowmanana McDunnough, 1923. Canadian Entomologist 55: 165–166 (Type locality: Canada, Alberta, Nordegg [Holotype: CNCO]).
Material examined. 1 ♂ (holotype), No. 607, [Canada], Alta [Alberta], Nordegg, 23.vii.1922, K. Bowmann leg. [coll. CNCO]. Remarks. This species was described by McDunnough (1923) on the basis of the wing pattern and assigned to Argyroploce Hübner, [1825]. Heinrich (1926) treated Argyroploce as a junior synonym of Olethreutes Hübner, [1822], and provided only a short diagnosis, without a detailed illustration of the male genitalia. Hannemann (1961) rejected Heinrich’s (1926) opinion and treated Argyroploce as a separate genus. Falkovitsh (1965, 1966) followed Hannemann (1961) and described two species, Argyroploce magadana and A. kamtshadala, both from Russia. In his monograph on the Tortricidae of the Russian Far East, Kuznetzov (2001) also treated Argyroploce as a separate genus and noted that the Russian tundra species A. magadana is replaced in North America by the related A. bowmanana. Based on a study of the type material of O. bowmanana, we conclude that this species indeed bears a strong similarity with, and is probably related to O. magadana and O. kamtshadala. A redescription of O. bowmanana is given below with a special emphasis on the characters shared with the Palaearctic species. Description. Forewing length 8 mm. Head: Vertex and palpi with a mixture of blackish and grayish scales. Antennae dark gray, about half a long as forewing. Thorax: Concolorous with head. Forewing elongate, with costa nearly straight, termen oblique and straight, dorsal angle rounded; upperside ground color grayish brown, with distinct pattern; basal, median and postmedian fasciae well defined, with narrow dark striae between them; subbasal fascia narrow, curved in median cell, broader dorsally, with outer margin distinct; median fascia narrow; outer margin distinct, fissured, with a process beyond whitish discal spot; postmedian fascia divided by costal strigula; preterminal fascia narrow; apical spot small, round, and blackish; costal strigulae whitish gray. Cilia brownish gray, and checkered with darker spots; marginal line well defined. Hindwing elongate-ovoid, with narrowly rounded apex; grayish brown. Cilia concolorous with wing. Anal roll well developed. Abdomen: Dark gray. Male genitalia (Fig. 9) with uncus bifid, covered with long hairs. Socii digitate, laterally fused with tegumen. Valva elongate; cucullus rounded, spatulate, slightly wider than neck of valva; neck shorter than cucullus, ratio of its length to width 1.5:1; basal part of valva broad; lobular process at distal part of sacculus covered with strong spines, cluster of spines extending basally and costally to nearly basal cavity; sacculus at basal side of lobular
376 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
process with wide setose spine cluster; ventral edge of sacculus covered with row of setae. Aedeagus slightly curved downward, dorsally membranous; sclerotized ventral part projecting distally to form a narrow apical foldable process with strong spine on right side apically. Female genitalia unknown. Diagnosis. Olethreutes bowmanana is most similar to O. kamtshadala but can be distinguished by the following characters: valvae neck wider, slightly narrower than cucullus, whereas in O. kamtshadala the neck is less than half as wide as the cucullus; the lobular process of the sacculus is wide and flat, not raise above the other parts of sacculus; the setose spine cluster on the ventral margin of the sacculus is wider than that of O. kamtshadala and consists of stronger spines; and the aedeagus is longer, approximately 6 times its width, and nearly the same width throughout, whereas in O. kamtshadala the aedeagus is about 3 times as long as its maximum width and the tip about half as wide as the base. Distribution. Canada (Alberta).
FIGURES 1–6. Olethreutes spp., adults. 1–2. O. bowmanana. 1, holotype, male, CANADA, Alta., Nordegg; 2, holotype labels. 3, O. magadana, holotype, male RUSSIA, 160 km SE from Magadan, settl. Madaun. 4, O. kamtshadala, holotype, male, RUSSIA, Koryakia autonomous region, Apuka, Pakhachinsky Range. 5-6, O. pivanica. 5, holotype, male, RUSSIA, Khabarovsk Province, near Komsomolsk-na-Amure, Pivan; 6, paratype, female, the same locality.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
377
FIGURES 7–10. Olethreutes spp., male genitalia, holotypes. 7, O. kamtshadala, Russia, Kamchatka; 8, O. magadana, Russia, Magadan Province; 9, O. bowmanana, Canada, Alberta; 10, O. pivanica, Russia, Khabarovsk Province.
Olethreutes magadana (Falkovitsh 1965) (Fig. 3) Selenodes magadana Falkovitsh, 1965, Entomologicheskoe Obozrenie 44 (2): 422–423 (Type locality: Russia, 160 km SE from Magadan, settl. Madaun [Holotype: ZISP]).
Material examined. 1 ♂ (holotype), micr. prep. No. 310, [Russia], 160 km SE from Magadan, settl. Madaun, 18.vii.1964, Tsvetaev leg. [coll. ZISP]. Remarks. Descriptions of O. magadana and the next species, O. kamtshadala, were originally published in Russian. Redescriptions are given here in English for comparison of all species of the Olethreutes bowmananamagadana species group and to confirm the association of these species with the group. Description. Forewing length 9 mm. Head: Grayish brown; labial palpi relatively small, slightly broadened distally, covered externally by dark grayish brown scales and internally by whitish scales. Antennae brown, about one-half forewing length. Thorax: Grayish brown. Forewing subquadrangular, costal margin nearly straight, apex rounded; temen slightly oblique, tornus rounded; upperside ground color grayish brown; pattern typical of other members of genus, consisting of transversal light striae and dark fasciae; basal part gray, covered with dispersed darker grayish brown scales forming an oblique transversal marble pattern; median fascia grayish brown, distal margin oblique from middle part of costal margin towards tornal angle, turning acutely to dorsal margin at tornal angle, enclosing an oval grayish spot; costal strigulae dove gray; 6th–8th strigulae containing whitish scales; admixture of whitish scales reaching maximum at discal vein, forming a whitish spot; distal part of forewing
378 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
concolorous with basal part, but with one broader sinuous grayish brown preterminal fascia. Cilia gray, with well defined blackish marginal line. Hindwing elongate-ovoid, with slightly extended apex, grayish brown; costal margin with elongate basal lobe ca. 0.4 length of costa, its fore margin with elongate specialized androconial scales. Abdomen: Dark gray. Male genitalia (Fig. 8) with uncus slightly narrowed and weakly bifurcate apically, covered with long hairs. Socii separate, laterally fused with tegumen, covered with long hairs. Gnathos membranous. Valva long with broader basal part; cucullus gradually widened apically forming an ovoid club covered by long setae, its apex about three times as broad as neck of valva; neck of valve long, narrow, without setae; sacculus with wide triangular apical process, extended ventrally, narrowing towards costa, densely covered with strong setae; sacculus with a setose cluster near proximal edge of triangular process, ventral edge of process covered with row of hairs; basal side of triangular process with small setose spine cluster. Aedeagus slightly curved, nearly straight, membranous dorsally; sclerotized ventral part of aedeagus forming a narrow apical process, curved clockwise with a strong spine on top. Female genitalia with papillae anales elongate, broader apically (dorsally) and narrow ventrally. Sternite VII forming two leaf-like sclerotizations, one on each side of ostium. Ostium surrounded by two lateral semicircular sclerites. Vaginal sinus sclerotized with transverse narrowing near base. Ductus bursae moderate in length. Corpus bursae ovoid, without signum. Diagnosis. Olethreutes magadana is most similar to O. pivanica, sp. n., but can be distinguished by the following characters: cucullus ovoid, slightly elongate, whereas in O. pivanica the cucullus is wide, rounded; and apical process of the sacculus triangular, broadest ventrally, narrowing towards costa, whereas in O. pivanica the process is broader towards the costa, narrowing ventrally and forming a rounded ventral lobe. Distribution. Known only from the type series, collected at the upper part of the Kolyma river basin.
Olethreutes kamtshadala (Falkovitsh 1966) (Fig. 4) Selenodes kamtshadala Falkovitsh, 1966, Trudy Zoologicheskogo Instituta AN SSSR 37: 222–223 (Type locality: Russia, Koryakia autonomous region, Apuka, Pakhachinsky Range [Holotype: ZISP]).
Material examined. 1 ♂ (holotype), micr. prep. no. 326, [Russia], Koryakia autonomous region, Apuka, Pakhachinsky Range, mountain tundra, 300 m, 7.vii.1959, K. Gorodkov leg. [coll. ZISP]. Description. Forewing length 8 mm. Head: Light brownish gray; labial palpi small, covered externally by dark brownish gray scales. Antennae brown, slightly longer than one-half forewing length. Thorax: Light brown gray. Forewing subquadrangular, strongly elongate, with costal nearly straight and apex rounded; termen conspicuously oblique, straight, tornus rounded; upperside ground color dark brown with an admixture of whitish scales; small groups of black scales forming short black longitudinal streaks representing elements of broken subbasal, median, and postmedian fasciae; no conspicuous darker transversal fasciae; basal, mesal and distal wing parts concolorous, not separated from each other; costal strigulae white; 5th–8th strigulae fused with white ovoid postdiscal spot; ocellar region lighter at tornal part with three dark (not black) streaks; outer margin beyond apex with three white streaks. Cilia dark whitish with brown scales; marginal line distinct. Hindwing same shape as other members of species group, but apex less extended; light gray, slightly darker along outer margin, darkest at apex; costal margin with elongate basal lobe about 1/3 length of costa, its fore margin with elongate specialized androconial scales. Cilia whitish. Abdomen: Light gray. Male genitalia (Fig. 7) with uncus slightly expanded apically with distinct apical bifurcation. Socii fused laterally with tegumen. Gnathos membranous. Valva long, broadest basally; cucullus enlarged apically into an oviod bulb; neck short, ca. twice as long as width; sacculus with broadly ovoid, short, apical process, densely covered with strong setae; field of chaetae not narrowed towards costa, with a row of several strong chaetae along costa towards basal cavity; sacculus apex with small setose spine cluster. Aedeagus similar to other species of the group. Female unknown. Diagnosis. Olethreutes kamtshadala is most similar to O. magadana but can be distinguished by the following characters: neck of the valva shorter than the cucullus, ca. twice as long as wide, whereas in O. magadana the neck is nearly as long as the cucullus, ca. four times as long as wide; and the apical process of the sacculus short, broadly ovoid, whereas in O. magadana process distinctly triangular. Distribution. Northern part of Kamchatka Peninsula.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
379
Olethreutes pivanica Dubatolov, Bogunova & Nedoshivina, sp. n. (Figs. 5–6) Type material: Holotype: ♂, RUSSIA, Khabarovsk Province, near Komsomolsk-na-Amure, Pivan, 50° 34’ N 137° 03’ E, 16.vi. 2008, leg. V. V. Dubatolov. Deposited in ISEA. Paratype (allotype): female, the same locality, 14.vi 2008, leg. V. V. Dubatolov. Description. Forewing length 9 mm. Head: Dark brown; labial palpi relatively small, slightly broadened distally, covered by dark brown scales. Antennae brown, ca. one-half forewing length. Thorax: Dark brown. Forewing with costal margin slightly convex, apex rounded, termen slightly oblique, tornus rounded; upperside ground color brownish-black; pattern typical of members of genus, consisting of transversal dark brownish fasciae with narrow, lighter, dove grayish striae between them; median stria not contrasting, with sparse dark scales, extending from basal one-third of costa towards one-half of dorsal margin, outer margin of dark median fascia slightly convex, jagged, with small angle in central cell; postmedian transversal stria fused dorsally, enclosing darker pretornal patch; submarginal stria dark and indistinct; dark external field triangularly narrowed towards tornus; light pattern brighter around discal vein. Forewing cilia brownish-grey. Hind wing ovoid with slightly extended apex, dark brown with slightly lighter base; costal margin with elongate basal lobe about 1/3 of costal length, its fore margin with elongate special androconial scales (Fig. 12). Cilia lighter than hindwing. Abdomen: Dark gray. Male genitalia (Fig. 10) with uncus bifurcate apically, covered with long hairs. Socii separate, partly fused with tegumen, forming broad ovoid processes, covered with long hairs. Gnathos membranous. Valva long, with broad basal part; cucullus roughly widened apically, forming round club covered by long chaetae; cucullus apex more than four times as broad as valva neck; valva neck long, narrow, without chaetae; sacculus with narrowly triangular apical process, broader towards costa, narrower ventrally, forming rounded ventral lobe, densely covered with strong chaetae; field of chaetae expanded at costa forming a row at cucullus base, ending in two strong chaetae; sacculus with a cluster of chaetae near ventral lobe of triangular process, ventral edge of sacculus covered with row of hairs; anterior group of sparse, weaker chaetae located at distal edge of basal cavity. Aedeagus slightly curved, dorsally membranous; sclerotized ventral part of aedeagus forming a narrow apical process, curved clockwise with a strong spine on dorsum. Female genitalia (Fig. 11) with papillae anales narrow, elongate, slightly broadened apically. Sternite VII horseshoe-like, forming two broad bands laterally from ostium. Ostium surrounded by broad semicircular sclerite extended caudally. Vaginal sinus forming narrow goblet-like sclerite, slightly hollowed proximally. Ductus burase moderate in length. Corpus bursa ovoid, without signum. Diagnosis. Olethreutes pivanica is not easily distinguished from the related Palearctic species by wing pattern alone, but it is noticeably darker. The male genitalia of O. pivanica are similar to those of O. magadana (Falkovitsh 1965) from the Kolyma region in the North-East Asia and O. kamtshadala (Falkovitsh 1966) from Kamchatka. All these species have a strong process of the sacculus that is covered by dense strong chaetae; the basal part of valva is broad, and the cucullus is club shaped. Although similar, the shape of the sacculus process is different: ovoid and broadly rounded apically in O. kamtshadala, narrowly triangular in O. pivanica, and broadly triangular in O. magadana. The ratio of the length to width of this process is about 2:1 in O. pivanica and 1:1 in O. magadana. The row of chaetae on the costal edge of the process base is long in O. pivanica (nearly equal to the length of the process) and very short in O. magadana. The ratio of the length to width of the valva neck is 3:1 in O. pivanica and 4:1 in O. magadana. In addition, the club shape of the cucullus is more or less round in O. pivanica and ovoid in O. magadana. The North American species O. bowmanana (McDunnough 1923) is less similar to the new species than are the two East Asian ones. The saccular process of O. bowmanana is very broad, nearly rectangular, and expands immediately from the basal cavity, whereas in all Asian species it is located conspicuously more distal. The valval neck in O. bowmanana is short and very broad, only slightly narrower than the cucullus apex. Biology. Olethreutes pivanica was collected at the lower part of a petrophytous sunny slope of a terrace of the Amur River bank. Moths were flying over the small boulders covered with sparse vegetation. The species probably occurs in the middle part of the petrophytous slope, but the slope is too steep for collecting. The species flew with the more common Cnephasia ussurica Filipjev 1962, and these were the only tortricoid species observed at this slope. Neither was found in other places of the Region (Syachina 2008), and they are probably restricted to this biotope. Another characteristic species at this site was the Holarctic Clossiana tritonia (Boeber 1812) [Nymphalidae], represented by the topotype population of C. t. amphilochus (Ménétriès 1859).
380 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
FIGURES 11–12.Olethreutes pivanica, Khabarovsk province. 11 - female genitalia; 12 - male hindwing costal margin, underside.
Distribution. Olethreutes pivanica is probably restricted to the Lower Amur basin and the mountains of the Region. Etymology. The new species is named after the type locality place, Pivan. Remarks. The new species is most similar to O. magadana, which was reported to occur in Chukotka, Magadan Province, Koryakia, Chita Province and Taimyr (Kuznetzov 2001). We found a few specimens labeled as O. magadana that were identified incorrectly by V. I. Kuznetzov in the collection of the Zoological Institute, St. Petersburg. After dissection, specimens from Chukotka were identified as O. exaridana (Kuznetzov 1991); a female from Kolyma was not identified, but it is not similar to the O. magadana-kamtshadala species group based on its genitalia. Another female from Taimyr has the genitalia similar to the new species and belongs to an uncertain species of the same complex. So, O. magadana Flkv. is actually known only from the Kolyma basin. All other data about distribution of O. magadana Flkv. in Chukotka and Kolyma were probably based on the incorrect identification. Additional specimens from Chukotka identified by Prof. V. I. Kuznetzov as O. kamtshadala Flkv. belong to Cymolomia taigana Falkovitsh, 1966. Thus, occurrence of O. kamtshadala Flkv. in Chukotka remains also questionable.
Acknowledgments We thank Jean-François Landry (Ottawa, Canada) for his kind support during the work of the late author with the type material of North American Tortricidae; Valery A. Mutin (Komsomolsk-na-Amure, Russia) for his help in the investigation of Lepidoptera fauna of the Pivan region; Tim and Oxana Healy (Seattle, USA) for improvements of the language; and John W. Brown (USDA, National Museum of Natural History, Washington, U.S.A.) for editing the manuscript.
References Boeber, de. (1812) Continuation de la description de quelques nouvelles espèces de papillons découverts en Sibérie etc. Memoires de la Société Impériale des Naturalistes de Moscou, 3, 2–21, tab. 1. Brown, J.W., Baixeras, J., Brown, R., Horak, M., Komai, F., Metzler, E.H., Razowski, J. & Tuck, K. (2005) Tortricidae (Lepidoptera). In: Landry, B. (Ed.), World Catalogue of Insects. Vol. 5. Apollo Books, Stenstrup, 741 pp.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
381
Clarke, J.F.G. (1941) The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society, 36, 149–161. Falkovitsh, M.I. (1965) New East Asian species of leaf-rollers (Lepidoptera, Tortricidae). Entomologicheskoe Obozrenie, 44 (2), 414–437. [in Russian] Falkovitsh, M.I. (1966) New Palearctic species of leaf-rollers from subfamily Olethreutinae (Lepidoptera, Tortricidae). Trudy Zoologicheskogo Instituta AN SSSR. Leningrad, 37, 208–227. [in Russian] Filipjev, N.N. (1962) New species of Tortricinae (Lepidoptera, Tortricidae) in the fauna of the USSR. Trudy Zoologicheskogo Instituta AN SSSR. 30, 369–381. [in Russian] Gilligan, T.M., Wright, D.J. & Gibson, L.D. (2008) Olethreutine Moths of the Midwestern United States. An Identification Guide. Ohio Biological Survey Bulletin New Series, 16 (2), 334 pp. Gilligan, T.M., Baixeras, J., Brown, J.W. & Tuck, K.R. (2012) T@RTS: Online World Catalogue of the Tortricidae (Ver. 2.0). Available from: http://www.tortricid.net/catalogue.asp (accessed 19 February 2014) Hannemann, H.J. (1961) Kleinschmetterlinge oder Microlepidoptera. I. Die Wickler (s. str.) (Tortricidae). Dahl, F. (Ed.). Die Tierwelt Deutschlands. – Jena, Gustav Fischer, 48, 1–233. Hardwick, D.F. (1950) Preparation of slide mounts of lepidopterous genitalia. Canadian Entomologist, 82, 231–235. http://dx.doi.org/10.4039/ent82231-11 Heinrich, C. (1926) Revision of the North American moths of the subfamilies Laspeyresiinae and Olethreutinae. United States National Museum Bulletin, 132, 1–216. http://dx.doi.org/10.5479/si.03629236.132.1 Hübner, J. (1822) Systematisch-alphabetisches Verzeichnis aller bisher bey den Fürbildungen zur Sammlung europäischer Schmetterlinge angegebenen Gattungsbenennungen. Augsburg, VI + 81 pp. http://dx.doi.org/10.5962/bhl.title.48605 Kuznetsov, V.I. & Mikkola, K. (1991) The leaf-roller fauna of north-eastern Siberia, USSR, with descriptions of three new species (Lepidoptera, Tortricidae). Nota Lepidopterologica, 14, 194–219. Kuznetsov, V.I. (2001) Tortricidae (Olethreutidae, Cochylidae). Opredelitel’ nasekomykh Dal’nego Vostoka Rossii. Dal’nauka, Vladivostok, 5 (3), 11–473. [in Russian] Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum characteribus, differentiis, synonymis, locis. Edito Decima, Reformata. Vol. 1. Holmiae, 824 pp. McDunnough, J. (1923) New Canadian Lepidoptera. The Canadian Entomologist, 55, 163–168. http://dx.doi.org/10.4039/ent55163-7 Ménétriès, E. (1859) Lepidoptères de la Sibérie orientale et en particulier des rives de l'Amour; par M.Ménétriès. (Lu le 11 juin 1858). Bulletin de la classe physico-mathématique de l’Académie Impériale des Sciences de St.-Pétersbourg, 17 (14), 212– 221. Syachina, A.A. (2008) A review of the fauna of leaf-rollers (Lepidoptera, Tortricidae) from environs of Komsomolsk-naAmure (Khabarovskii Krai). Problemy ekologii Verkhnego Priamur’ya, Vypusk 10, Tom 2, 98–125. [in Russian]
382 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3779.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:13A26F5C-4675-45DE-920F-53739FB1496D
Review of the Olethreutes bowmanana-magadana species group, with the description of a new species from the Amur River, Russia (Lepidoptera, Tortricidae) VLADIMIR V. DUBATOLOV1, ANNA A. BOGUNOVA (SYACHINA)2 & SVETLANA V. NEDOSHIVINA3 1 Siberian Zoological Museum, Institute of Systematics and Ecology of Animals, Siberian Branch of Russian Academy of Sciences, Frunze street 11, RF-630091 Novosibirsk, Russia. E-mail: [email protected] 2 Biology Department, Amursky Liberal-Pedagogical State University, Kirov Street, 17/2, 681000 Komsomolsk-na-Amure, Russia. E-mail: [email protected] 3 Department of Zoology, Ulyanovsk State Pedagogical University, pl. 100-letiya Lenina 4, RF-432700 Ulyanovsk, Russia. E-mail: [email protected]
Abstract We define and review the Olethreutes bowmanana-magadana species group comprised of Olethreutes bowmanana (McDunnough), O. magadana (Falkovitsh), O. kamtshadala (Falkovitsh), and O. pivanica, sp. n., the last from petrophytous sunny slopes in the Amur River. The new species differs from other species of the group by a narrow, triangular-shaped apical process from the sacculus; a round, wide, apical bulb from the cucullus; and a longer, narrow neck of the valva. Key words: Tortricidae, Olethreutes bowmanana, Olethreutes magadana, Olethreutes kamtshadala, new species, Amur River, Russia, Far East
Introduction During a study of the lepidopterous fauna of the Lower Amur Region, Russia, a new species of Olethreutes Hübner [1822] was discovered. The moths were collected on a petrophytous sunny slope of a terrace on the Amur River bank opposite to Komsomolsk-na-Amure city. The specimens belong to a group of closely related species including O. magadana (Falkovitsh 1965) and O. kamtshadala (Falkovitsh 1966), but they differ noticeably from both of them. A comparison of the specimens with types and additional material in the Zoological Institute (St. Petersburg, Russia) revealed that some specimens in the collection of Zoological Institute were misidentified. This finding prompted us to prepare a review of the species of this group, including the description of the new species.
Material and methods Dissection methods follow those of Clarke (1941) and Hardwick (1950). Terminology for morphological features of wings and genitalia follow Gilligan et al. (2008). The following depositories are abbreviated in the text: CNCO ISEA ZISP
Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Canada; Siberian Zoological Museum, Institute of Systematics and Ecology of Animals, Novosibirsk, Russia; Zoological Institute of Russian Academy of Sciences, St. Petersburg, Russia.
Accepted by J. Brown: 31 Jan. 2014; published: 17 Mar. 2014
375
Results Olethreutes Hübner [1822] Systematisch-alphabetisches Verzeichniss aller bisher bey den Fürbildungen zur Sammlung europäischer Schmetterlinge: 72. Type species: Phalaena Tortrix arcuana Linnaeus, 1761 = [Phalaena] arcuella Clerck, 1759 by subsequent designation.
Olethreutes is a large Holarctic genus with 133 described species currently assigned to it (Gilligan et al. 2012). Species included in the genus exhibit a wide range of genital morphology wing patterns, and male secondary structures, making the genus extremely heterogeneous and probably polyphyletic. However, many species within Olethreutes show remarkable similarities to presumably closely related species, and many of these could be combined into natural groups. The Olethreutes bowmanana-magadana species group, proposed here, shows clear differences from other Olethreutes and is considered a natural group supported by following characters: brown to dark gray wing colour with lighter grayish striae and blackish spots; uncus narrow, bifid distally; sacculus with a distal internal lobe covered by strong spines and a spine cluster proximally on the ventral side; cucullus with a slender neck and apical widening; and aedeagus with foldable apical spine. The assignment of the species under consideration to the genus Olethreutes is generally accepted (Brown et al. 2005), but their relationships to other Olethreutes species remain uncertain.
Olethreutes bowmanana (McDunnough 1923) (Fig. 1) Argyroploce bowmanana McDunnough, 1923. Canadian Entomologist 55: 165–166 (Type locality: Canada, Alberta, Nordegg [Holotype: CNCO]).
Material examined. 1 ♂ (holotype), No. 607, [Canada], Alta [Alberta], Nordegg, 23.vii.1922, K. Bowmann leg. [coll. CNCO]. Remarks. This species was described by McDunnough (1923) on the basis of the wing pattern and assigned to Argyroploce Hübner, [1825]. Heinrich (1926) treated Argyroploce as a junior synonym of Olethreutes Hübner, [1822], and provided only a short diagnosis, without a detailed illustration of the male genitalia. Hannemann (1961) rejected Heinrich’s (1926) opinion and treated Argyroploce as a separate genus. Falkovitsh (1965, 1966) followed Hannemann (1961) and described two species, Argyroploce magadana and A. kamtshadala, both from Russia. In his monograph on the Tortricidae of the Russian Far East, Kuznetzov (2001) also treated Argyroploce as a separate genus and noted that the Russian tundra species A. magadana is replaced in North America by the related A. bowmanana. Based on a study of the type material of O. bowmanana, we conclude that this species indeed bears a strong similarity with, and is probably related to O. magadana and O. kamtshadala. A redescription of O. bowmanana is given below with a special emphasis on the characters shared with the Palaearctic species. Description. Forewing length 8 mm. Head: Vertex and palpi with a mixture of blackish and grayish scales. Antennae dark gray, about half a long as forewing. Thorax: Concolorous with head. Forewing elongate, with costa nearly straight, termen oblique and straight, dorsal angle rounded; upperside ground color grayish brown, with distinct pattern; basal, median and postmedian fasciae well defined, with narrow dark striae between them; subbasal fascia narrow, curved in median cell, broader dorsally, with outer margin distinct; median fascia narrow; outer margin distinct, fissured, with a process beyond whitish discal spot; postmedian fascia divided by costal strigula; preterminal fascia narrow; apical spot small, round, and blackish; costal strigulae whitish gray. Cilia brownish gray, and checkered with darker spots; marginal line well defined. Hindwing elongate-ovoid, with narrowly rounded apex; grayish brown. Cilia concolorous with wing. Anal roll well developed. Abdomen: Dark gray. Male genitalia (Fig. 9) with uncus bifid, covered with long hairs. Socii digitate, laterally fused with tegumen. Valva elongate; cucullus rounded, spatulate, slightly wider than neck of valva; neck shorter than cucullus, ratio of its length to width 1.5:1; basal part of valva broad; lobular process at distal part of sacculus covered with strong spines, cluster of spines extending basally and costally to nearly basal cavity; sacculus at basal side of lobular
376 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
process with wide setose spine cluster; ventral edge of sacculus covered with row of setae. Aedeagus slightly curved downward, dorsally membranous; sclerotized ventral part projecting distally to form a narrow apical foldable process with strong spine on right side apically. Female genitalia unknown. Diagnosis. Olethreutes bowmanana is most similar to O. kamtshadala but can be distinguished by the following characters: valvae neck wider, slightly narrower than cucullus, whereas in O. kamtshadala the neck is less than half as wide as the cucullus; the lobular process of the sacculus is wide and flat, not raise above the other parts of sacculus; the setose spine cluster on the ventral margin of the sacculus is wider than that of O. kamtshadala and consists of stronger spines; and the aedeagus is longer, approximately 6 times its width, and nearly the same width throughout, whereas in O. kamtshadala the aedeagus is about 3 times as long as its maximum width and the tip about half as wide as the base. Distribution. Canada (Alberta).
FIGURES 1–6. Olethreutes spp., adults. 1–2. O. bowmanana. 1, holotype, male, CANADA, Alta., Nordegg; 2, holotype labels. 3, O. magadana, holotype, male RUSSIA, 160 km SE from Magadan, settl. Madaun. 4, O. kamtshadala, holotype, male, RUSSIA, Koryakia autonomous region, Apuka, Pakhachinsky Range. 5-6, O. pivanica. 5, holotype, male, RUSSIA, Khabarovsk Province, near Komsomolsk-na-Amure, Pivan; 6, paratype, female, the same locality.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
377
FIGURES 7–10. Olethreutes spp., male genitalia, holotypes. 7, O. kamtshadala, Russia, Kamchatka; 8, O. magadana, Russia, Magadan Province; 9, O. bowmanana, Canada, Alberta; 10, O. pivanica, Russia, Khabarovsk Province.
Olethreutes magadana (Falkovitsh 1965) (Fig. 3) Selenodes magadana Falkovitsh, 1965, Entomologicheskoe Obozrenie 44 (2): 422–423 (Type locality: Russia, 160 km SE from Magadan, settl. Madaun [Holotype: ZISP]).
Material examined. 1 ♂ (holotype), micr. prep. No. 310, [Russia], 160 km SE from Magadan, settl. Madaun, 18.vii.1964, Tsvetaev leg. [coll. ZISP]. Remarks. Descriptions of O. magadana and the next species, O. kamtshadala, were originally published in Russian. Redescriptions are given here in English for comparison of all species of the Olethreutes bowmananamagadana species group and to confirm the association of these species with the group. Description. Forewing length 9 mm. Head: Grayish brown; labial palpi relatively small, slightly broadened distally, covered externally by dark grayish brown scales and internally by whitish scales. Antennae brown, about one-half forewing length. Thorax: Grayish brown. Forewing subquadrangular, costal margin nearly straight, apex rounded; temen slightly oblique, tornus rounded; upperside ground color grayish brown; pattern typical of other members of genus, consisting of transversal light striae and dark fasciae; basal part gray, covered with dispersed darker grayish brown scales forming an oblique transversal marble pattern; median fascia grayish brown, distal margin oblique from middle part of costal margin towards tornal angle, turning acutely to dorsal margin at tornal angle, enclosing an oval grayish spot; costal strigulae dove gray; 6th–8th strigulae containing whitish scales; admixture of whitish scales reaching maximum at discal vein, forming a whitish spot; distal part of forewing
378 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
concolorous with basal part, but with one broader sinuous grayish brown preterminal fascia. Cilia gray, with well defined blackish marginal line. Hindwing elongate-ovoid, with slightly extended apex, grayish brown; costal margin with elongate basal lobe ca. 0.4 length of costa, its fore margin with elongate specialized androconial scales. Abdomen: Dark gray. Male genitalia (Fig. 8) with uncus slightly narrowed and weakly bifurcate apically, covered with long hairs. Socii separate, laterally fused with tegumen, covered with long hairs. Gnathos membranous. Valva long with broader basal part; cucullus gradually widened apically forming an ovoid club covered by long setae, its apex about three times as broad as neck of valva; neck of valve long, narrow, without setae; sacculus with wide triangular apical process, extended ventrally, narrowing towards costa, densely covered with strong setae; sacculus with a setose cluster near proximal edge of triangular process, ventral edge of process covered with row of hairs; basal side of triangular process with small setose spine cluster. Aedeagus slightly curved, nearly straight, membranous dorsally; sclerotized ventral part of aedeagus forming a narrow apical process, curved clockwise with a strong spine on top. Female genitalia with papillae anales elongate, broader apically (dorsally) and narrow ventrally. Sternite VII forming two leaf-like sclerotizations, one on each side of ostium. Ostium surrounded by two lateral semicircular sclerites. Vaginal sinus sclerotized with transverse narrowing near base. Ductus bursae moderate in length. Corpus bursae ovoid, without signum. Diagnosis. Olethreutes magadana is most similar to O. pivanica, sp. n., but can be distinguished by the following characters: cucullus ovoid, slightly elongate, whereas in O. pivanica the cucullus is wide, rounded; and apical process of the sacculus triangular, broadest ventrally, narrowing towards costa, whereas in O. pivanica the process is broader towards the costa, narrowing ventrally and forming a rounded ventral lobe. Distribution. Known only from the type series, collected at the upper part of the Kolyma river basin.
Olethreutes kamtshadala (Falkovitsh 1966) (Fig. 4) Selenodes kamtshadala Falkovitsh, 1966, Trudy Zoologicheskogo Instituta AN SSSR 37: 222–223 (Type locality: Russia, Koryakia autonomous region, Apuka, Pakhachinsky Range [Holotype: ZISP]).
Material examined. 1 ♂ (holotype), micr. prep. no. 326, [Russia], Koryakia autonomous region, Apuka, Pakhachinsky Range, mountain tundra, 300 m, 7.vii.1959, K. Gorodkov leg. [coll. ZISP]. Description. Forewing length 8 mm. Head: Light brownish gray; labial palpi small, covered externally by dark brownish gray scales. Antennae brown, slightly longer than one-half forewing length. Thorax: Light brown gray. Forewing subquadrangular, strongly elongate, with costal nearly straight and apex rounded; termen conspicuously oblique, straight, tornus rounded; upperside ground color dark brown with an admixture of whitish scales; small groups of black scales forming short black longitudinal streaks representing elements of broken subbasal, median, and postmedian fasciae; no conspicuous darker transversal fasciae; basal, mesal and distal wing parts concolorous, not separated from each other; costal strigulae white; 5th–8th strigulae fused with white ovoid postdiscal spot; ocellar region lighter at tornal part with three dark (not black) streaks; outer margin beyond apex with three white streaks. Cilia dark whitish with brown scales; marginal line distinct. Hindwing same shape as other members of species group, but apex less extended; light gray, slightly darker along outer margin, darkest at apex; costal margin with elongate basal lobe about 1/3 length of costa, its fore margin with elongate specialized androconial scales. Cilia whitish. Abdomen: Light gray. Male genitalia (Fig. 7) with uncus slightly expanded apically with distinct apical bifurcation. Socii fused laterally with tegumen. Gnathos membranous. Valva long, broadest basally; cucullus enlarged apically into an oviod bulb; neck short, ca. twice as long as width; sacculus with broadly ovoid, short, apical process, densely covered with strong setae; field of chaetae not narrowed towards costa, with a row of several strong chaetae along costa towards basal cavity; sacculus apex with small setose spine cluster. Aedeagus similar to other species of the group. Female unknown. Diagnosis. Olethreutes kamtshadala is most similar to O. magadana but can be distinguished by the following characters: neck of the valva shorter than the cucullus, ca. twice as long as wide, whereas in O. magadana the neck is nearly as long as the cucullus, ca. four times as long as wide; and the apical process of the sacculus short, broadly ovoid, whereas in O. magadana process distinctly triangular. Distribution. Northern part of Kamchatka Peninsula.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
379
Olethreutes pivanica Dubatolov, Bogunova & Nedoshivina, sp. n. (Figs. 5–6) Type material: Holotype: ♂, RUSSIA, Khabarovsk Province, near Komsomolsk-na-Amure, Pivan, 50° 34’ N 137° 03’ E, 16.vi. 2008, leg. V. V. Dubatolov. Deposited in ISEA. Paratype (allotype): female, the same locality, 14.vi 2008, leg. V. V. Dubatolov. Description. Forewing length 9 mm. Head: Dark brown; labial palpi relatively small, slightly broadened distally, covered by dark brown scales. Antennae brown, ca. one-half forewing length. Thorax: Dark brown. Forewing with costal margin slightly convex, apex rounded, termen slightly oblique, tornus rounded; upperside ground color brownish-black; pattern typical of members of genus, consisting of transversal dark brownish fasciae with narrow, lighter, dove grayish striae between them; median stria not contrasting, with sparse dark scales, extending from basal one-third of costa towards one-half of dorsal margin, outer margin of dark median fascia slightly convex, jagged, with small angle in central cell; postmedian transversal stria fused dorsally, enclosing darker pretornal patch; submarginal stria dark and indistinct; dark external field triangularly narrowed towards tornus; light pattern brighter around discal vein. Forewing cilia brownish-grey. Hind wing ovoid with slightly extended apex, dark brown with slightly lighter base; costal margin with elongate basal lobe about 1/3 of costal length, its fore margin with elongate special androconial scales (Fig. 12). Cilia lighter than hindwing. Abdomen: Dark gray. Male genitalia (Fig. 10) with uncus bifurcate apically, covered with long hairs. Socii separate, partly fused with tegumen, forming broad ovoid processes, covered with long hairs. Gnathos membranous. Valva long, with broad basal part; cucullus roughly widened apically, forming round club covered by long chaetae; cucullus apex more than four times as broad as valva neck; valva neck long, narrow, without chaetae; sacculus with narrowly triangular apical process, broader towards costa, narrower ventrally, forming rounded ventral lobe, densely covered with strong chaetae; field of chaetae expanded at costa forming a row at cucullus base, ending in two strong chaetae; sacculus with a cluster of chaetae near ventral lobe of triangular process, ventral edge of sacculus covered with row of hairs; anterior group of sparse, weaker chaetae located at distal edge of basal cavity. Aedeagus slightly curved, dorsally membranous; sclerotized ventral part of aedeagus forming a narrow apical process, curved clockwise with a strong spine on dorsum. Female genitalia (Fig. 11) with papillae anales narrow, elongate, slightly broadened apically. Sternite VII horseshoe-like, forming two broad bands laterally from ostium. Ostium surrounded by broad semicircular sclerite extended caudally. Vaginal sinus forming narrow goblet-like sclerite, slightly hollowed proximally. Ductus burase moderate in length. Corpus bursa ovoid, without signum. Diagnosis. Olethreutes pivanica is not easily distinguished from the related Palearctic species by wing pattern alone, but it is noticeably darker. The male genitalia of O. pivanica are similar to those of O. magadana (Falkovitsh 1965) from the Kolyma region in the North-East Asia and O. kamtshadala (Falkovitsh 1966) from Kamchatka. All these species have a strong process of the sacculus that is covered by dense strong chaetae; the basal part of valva is broad, and the cucullus is club shaped. Although similar, the shape of the sacculus process is different: ovoid and broadly rounded apically in O. kamtshadala, narrowly triangular in O. pivanica, and broadly triangular in O. magadana. The ratio of the length to width of this process is about 2:1 in O. pivanica and 1:1 in O. magadana. The row of chaetae on the costal edge of the process base is long in O. pivanica (nearly equal to the length of the process) and very short in O. magadana. The ratio of the length to width of the valva neck is 3:1 in O. pivanica and 4:1 in O. magadana. In addition, the club shape of the cucullus is more or less round in O. pivanica and ovoid in O. magadana. The North American species O. bowmanana (McDunnough 1923) is less similar to the new species than are the two East Asian ones. The saccular process of O. bowmanana is very broad, nearly rectangular, and expands immediately from the basal cavity, whereas in all Asian species it is located conspicuously more distal. The valval neck in O. bowmanana is short and very broad, only slightly narrower than the cucullus apex. Biology. Olethreutes pivanica was collected at the lower part of a petrophytous sunny slope of a terrace of the Amur River bank. Moths were flying over the small boulders covered with sparse vegetation. The species probably occurs in the middle part of the petrophytous slope, but the slope is too steep for collecting. The species flew with the more common Cnephasia ussurica Filipjev 1962, and these were the only tortricoid species observed at this slope. Neither was found in other places of the Region (Syachina 2008), and they are probably restricted to this biotope. Another characteristic species at this site was the Holarctic Clossiana tritonia (Boeber 1812) [Nymphalidae], represented by the topotype population of C. t. amphilochus (Ménétriès 1859).
380 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
FIGURES 11–12.Olethreutes pivanica, Khabarovsk province. 11 - female genitalia; 12 - male hindwing costal margin, underside.
Distribution. Olethreutes pivanica is probably restricted to the Lower Amur basin and the mountains of the Region. Etymology. The new species is named after the type locality place, Pivan. Remarks. The new species is most similar to O. magadana, which was reported to occur in Chukotka, Magadan Province, Koryakia, Chita Province and Taimyr (Kuznetzov 2001). We found a few specimens labeled as O. magadana that were identified incorrectly by V. I. Kuznetzov in the collection of the Zoological Institute, St. Petersburg. After dissection, specimens from Chukotka were identified as O. exaridana (Kuznetzov 1991); a female from Kolyma was not identified, but it is not similar to the O. magadana-kamtshadala species group based on its genitalia. Another female from Taimyr has the genitalia similar to the new species and belongs to an uncertain species of the same complex. So, O. magadana Flkv. is actually known only from the Kolyma basin. All other data about distribution of O. magadana Flkv. in Chukotka and Kolyma were probably based on the incorrect identification. Additional specimens from Chukotka identified by Prof. V. I. Kuznetzov as O. kamtshadala Flkv. belong to Cymolomia taigana Falkovitsh, 1966. Thus, occurrence of O. kamtshadala Flkv. in Chukotka remains also questionable.
Acknowledgments We thank Jean-François Landry (Ottawa, Canada) for his kind support during the work of the late author with the type material of North American Tortricidae; Valery A. Mutin (Komsomolsk-na-Amure, Russia) for his help in the investigation of Lepidoptera fauna of the Pivan region; Tim and Oxana Healy (Seattle, USA) for improvements of the language; and John W. Brown (USDA, National Museum of Natural History, Washington, U.S.A.) for editing the manuscript.
References Boeber, de. (1812) Continuation de la description de quelques nouvelles espèces de papillons découverts en Sibérie etc. Memoires de la Société Impériale des Naturalistes de Moscou, 3, 2–21, tab. 1. Brown, J.W., Baixeras, J., Brown, R., Horak, M., Komai, F., Metzler, E.H., Razowski, J. & Tuck, K. (2005) Tortricidae (Lepidoptera). In: Landry, B. (Ed.), World Catalogue of Insects. Vol. 5. Apollo Books, Stenstrup, 741 pp.
REVIEW OF THE OLETHREUTES BOWMANANA SPECIES GROUP
Zootaxa 3779 (3) © 2014 Magnolia Press ·
381
Clarke, J.F.G. (1941) The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society, 36, 149–161. Falkovitsh, M.I. (1965) New East Asian species of leaf-rollers (Lepidoptera, Tortricidae). Entomologicheskoe Obozrenie, 44 (2), 414–437. [in Russian] Falkovitsh, M.I. (1966) New Palearctic species of leaf-rollers from subfamily Olethreutinae (Lepidoptera, Tortricidae). Trudy Zoologicheskogo Instituta AN SSSR. Leningrad, 37, 208–227. [in Russian] Filipjev, N.N. (1962) New species of Tortricinae (Lepidoptera, Tortricidae) in the fauna of the USSR. Trudy Zoologicheskogo Instituta AN SSSR. 30, 369–381. [in Russian] Gilligan, T.M., Wright, D.J. & Gibson, L.D. (2008) Olethreutine Moths of the Midwestern United States. An Identification Guide. Ohio Biological Survey Bulletin New Series, 16 (2), 334 pp. Gilligan, T.M., Baixeras, J., Brown, J.W. & Tuck, K.R. (2012) T@RTS: Online World Catalogue of the Tortricidae (Ver. 2.0). Available from: http://www.tortricid.net/catalogue.asp (accessed 19 February 2014) Hannemann, H.J. (1961) Kleinschmetterlinge oder Microlepidoptera. I. Die Wickler (s. str.) (Tortricidae). Dahl, F. (Ed.). Die Tierwelt Deutschlands. – Jena, Gustav Fischer, 48, 1–233. Hardwick, D.F. (1950) Preparation of slide mounts of lepidopterous genitalia. Canadian Entomologist, 82, 231–235. http://dx.doi.org/10.4039/ent82231-11 Heinrich, C. (1926) Revision of the North American moths of the subfamilies Laspeyresiinae and Olethreutinae. United States National Museum Bulletin, 132, 1–216. http://dx.doi.org/10.5479/si.03629236.132.1 Hübner, J. (1822) Systematisch-alphabetisches Verzeichnis aller bisher bey den Fürbildungen zur Sammlung europäischer Schmetterlinge angegebenen Gattungsbenennungen. Augsburg, VI + 81 pp. http://dx.doi.org/10.5962/bhl.title.48605 Kuznetsov, V.I. & Mikkola, K. (1991) The leaf-roller fauna of north-eastern Siberia, USSR, with descriptions of three new species (Lepidoptera, Tortricidae). Nota Lepidopterologica, 14, 194–219. Kuznetsov, V.I. (2001) Tortricidae (Olethreutidae, Cochylidae). Opredelitel’ nasekomykh Dal’nego Vostoka Rossii. Dal’nauka, Vladivostok, 5 (3), 11–473. [in Russian] Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum characteribus, differentiis, synonymis, locis. Edito Decima, Reformata. Vol. 1. Holmiae, 824 pp. McDunnough, J. (1923) New Canadian Lepidoptera. The Canadian Entomologist, 55, 163–168. http://dx.doi.org/10.4039/ent55163-7 Ménétriès, E. (1859) Lepidoptères de la Sibérie orientale et en particulier des rives de l'Amour; par M.Ménétriès. (Lu le 11 juin 1858). Bulletin de la classe physico-mathématique de l’Académie Impériale des Sciences de St.-Pétersbourg, 17 (14), 212– 221. Syachina, A.A. (2008) A review of the fauna of leaf-rollers (Lepidoptera, Tortricidae) from environs of Komsomolsk-naAmure (Khabarovskii Krai). Problemy ekologii Verkhnego Priamur’ya, Vypusk 10, Tom 2, 98–125. [in Russian]
382 · Zootaxa 3779 (3) © 2014 Magnolia Press
DUBATOLOV ET AL.
