59
Accepted by J-F. Landry: 8 Apr. 2013; published: 15 May 2013 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2013 Magnolia Press Zootaxa 3652 (1): 001059 www.mapress.com/ zootaxa/ Article 1 http://dx.doi.org/10.11646/zootaxa.3652.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:CD9BF682-A7CF-4A7B-85E7-A7302148718C Nepticulidae (Lepidoptera) of East Asia (1). Re-examination of the male genitalia of types deposited at the Russian Academy of Sciences JONAS R. STONIS* & AGN ROCIEN Division of Biosystematics Research, Department of Biology, Lithuanian University of Educational Sciences, Studentu St., 39, Vilnius LT-08106, Lithuania. E-mail: [email protected] *Corresponding author Abstract East Asia is famous for its tremendous overall biodiversity, and the Nepticulidae are no exception. The majority of the currently known fauna of Nepticulidae was described in 1984–1987 by several authors, but R. Puplesis is the author of the largest number of new species from the region. Unfortunately, the genitalia of all species from Primorskiy Kray, Sakh- alin, and the Kuril Islands were described from temporary slides in glycerol, and therefore, the drawings do not always show all details of genital armature or may be confusing. Without the baseline data providing morphology of genital ar- mature of type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae. In this paper we re- examine and document for the first time with photographs the types of 56 species described by R. Puplesis from the Rus- sian Far East. Details of the morphology is updated for most of the re-examined species. Key words: East Asia, male genitalia, Nepticulidae, taxonomy, type series Introduction A combination of features make Nepticulidae one of the most intriguing families of Lepidoptera: they are the smallest moths on the planet, phylogenetically representing very primitive heteroneurans. Furthermore, they are highly specialized: larvae live inside plant tissues and are predominantly monophagous (Diškus & Stonis 2012). Although Nepticulidae are global in distribution, there is evidence that great diversity occurs in continental East Asia and Japan. The history of description of Nepticulidae from East Asia is outlined in the next paper of this series (Rocien & Stonis 2013). R. Puplesis is responsible for the description of the largest number of new species from East Asia (see Navickait et al. 2011a): 59 new species (51 currently recognized), mostly from Primorskiy Kray (the Russian Far East), and several species from Sakhalin and the Kuril Islands (Puplesis 1984a, 1984b, 1984c, 1984d, 1985, 1987; Puplesis & Ivinskis 1985). All these species were treated by Puplesis (1994). In Nepticulidae, male genitalia usually provide the most reliable and distinctive features for species identification (diagnosis). Unfortunately, all species published by Rimantas Puplesis during 1984–1987 were described and illustrated from temporary slides in glycerol and using poor quality microscopes. In the original descriptions of the species (or even in the later review by Puplesis 1994), the indian-ink drawings of genitalia usually do not show sufficient details of the genital armature or are sometimes confusing. We made permanent mounts in Euparal and photographed genitalia slides of the holotypes of Puplesis’ species for the first time. The morphological descriptions of the re-examined species are updated. The present study is rather significant because these tiny moths have not been extensively studied and, therefore, many species still remain to be discovered and described. Without the baseline data providing detailed morphology of genital armature of the type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae.

Nepticulidae (Lepidoptera) of East Asia (1). Re-examination of ...biotaxonomy.eu/files/Publikacijos/216_ Nepticulidae of...Male genitalia of Stigmella mirabella (Puplesis) (Astigmella

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  • ZOOTAXAISSN 1175-5326 (print edition)

    ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press

    Zootaxa 3652 (1): 001–059 www.mapress.com/zootaxa/ Article

    http://dx.doi.org/10.11646/zootaxa.3652.1.1http://zoobank.org/urn:lsid:zoobank.org:pub:CD9BF682-A7CF-4A7B-85E7-A7302148718C

    Nepticulidae (Lepidoptera) of East Asia (1). Re-examination of the male genitalia of types deposited at the Russian Academy of Sciences

    JONAS R. STONIS* & AGNĖ ROCIENĖDivision of Biosystematics Research, Department of Biology, Lithuanian University of Educational Sciences, Studentu St., 39, Vilnius LT-08106, Lithuania. E-mail: [email protected]*Corresponding author

    Abstract

    East Asia is famous for its tremendous overall biodiversity, and the Nepticulidae are no exception. The majority of the currently known fauna of Nepticulidae was described in 1984–1987 by several authors, but R. Puplesis is the author of the largest number of new species from the region. Unfortunately, the genitalia of all species from Primorskiy Kray, Sakh-alin, and the Kuril Islands were described from temporary slides in glycerol, and therefore, the drawings do not always show all details of genital armature or may be confusing. Without the baseline data providing morphology of genital ar-mature of type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae. In this paper we re-examine and document for the first time with photographs the types of 56 species described by R. Puplesis from the Rus-sian Far East. Details of the morphology is updated for most of the re-examined species.

    Key words: East Asia, male genitalia, Nepticulidae, taxonomy, type series

    Introduction

    A combination of features make Nepticulidae one of the most intriguing families of Lepidoptera: they are the smallest moths on the planet, phylogenetically representing very primitive heteroneurans. Furthermore, they are highly specialized: larvae live inside plant tissues and are predominantly monophagous (Diškus & Stonis 2012). Although Nepticulidae are global in distribution, there is evidence that great diversity occurs in continental East Asia and Japan.

    The history of description of Nepticulidae from East Asia is outlined in the next paper of this series (Rocienė & Stonis 2013). R. Puplesis is responsible for the description of the largest number of new species from East Asia (see Navickaitė et al. 2011a): 59 new species (51 currently recognized), mostly from Primorskiy Kray (the Russian Far East), and several species from Sakhalin and the Kuril Islands (Puplesis 1984a, 1984b, 1984c, 1984d, 1985, 1987; Puplesis & Ivinskis 1985). All these species were treated by Puplesis (1994). In Nepticulidae, male genitalia usually provide the most reliable and distinctive features for species identification (diagnosis). Unfortunately, all species published by Rimantas Puplesis during 1984–1987 were described and illustrated from temporary slides in glycerol and using poor quality microscopes. In the original descriptions of the species (or even in the later review by Puplesis 1994), the indian-ink drawings of genitalia usually do not show sufficient details of the genital armature or are sometimes confusing.

    We made permanent mounts in Euparal and photographed genitalia slides of the holotypes of Puplesis’ species for the first time. The morphological descriptions of the re-examined species are updated. The present study is rather significant because these tiny moths have not been extensively studied and, therefore, many species still remain to be discovered and described. Without the baseline data providing detailed morphology of genital armature of the type material, it is impossible to continue studies on the East Asiatic fauna of Nepticulidae.

    Accepted by J-F. Landry: 8 Apr. 2013; published: 15 May 2013 1

  • Material and methods

    Type material of all but two species described by Puplesis from continental East Asia (Primorskiy Kray) and Sakhalin was available to the authors from the Zoological Institute of the Russian Academy of Sciences (St. Petersburg, Russia) (hereafter, ZIN).

    Permanent mounts of genitalia were prepared from temporary slides in glycerol following the method described by Robinson (1976). After taking out of glycerol, 0.3–0.5 minutes of maceration in 10% KOH and subsequent cleaning, male genital capsules were detached from the abdomen and mounted ventral side uppermost. In most cases the aedeagus was removed and mounted alongside the genital armature. Abdominal pelts were stained with Chlorazol Black (Direct Black 38/Azo Black) and mounted in Euparal. Permanent slides were photographed and studied using a Leica DFC420 digital camera and a Leica DM2500 microscope, respectively. The descriptive terminology of morphological structures follows van Nieukerken (1985, 1986), Johansson et al. (1990), Puplesis & Robinson (2000), and van Nieukerken et al. (2010).

    Photographic documentation of the types of Nepticulidae from East Asia described by R. Puplesis

    The classification of Nepticulidae follows Puplesis (1994), Puplesis & Robinson (2000), Puplesis et al. (2002), and Puplesis & Diškus (2003).

    1. Stigmella dissona (Puplesis, 1984) (Astigmella dissona Puplesis, 1984a: 112), a junior syn. of Stigmella naturnella(Klimesch, 1936).

    Morphology update (Figs 1–4): Gnathos deeply divided (Figs 1, 3, 4). System of cornuti complex (Figs 1, 3).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 28.v.1983, leg. R.

    Puplesis, genitalia slide no. AG 403 (ZIN).

    2. Stigmella mirabella (Puplesis, 1984) (Astigmella mirabella Puplesis, 1984a: 112).Morphology update (Figs 5–8): Valva without pointed apex (Fig. 6). Gnathos divided (Figs 5, 8). Vesica with 8–9

    large spines (Figs 7, 8).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 22.v.1983, leg. R.

    Puplesis, genitalia slide no. AG 404 (ZIN).

    3. Stigmella kurilensis Puplesis, 1987: 8.Recently studied and photographed by van Nieukerken & Kuroko 2005: 31–35, Figs 5–10.

    4. Stigmella sashai Puplesis, 1984b: 594–596.Morphology update (Figs 9–12): Valva with developed inner lobe (Figs 9, 10). Transverse bar of transtilla

    interrupted (Fig. 11). Cornuti spine-like; the vesica partially everted in the holotype (Fig. 12).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 29.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 401 (ZIN).

    5. Stigmella regina Puplesis, 1984b: 596.Morphology update (Figs 13–16): Aedeagus with two clusters of spine-like cornuti (Fig. 15).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 5.viii.1982, leg. R.

    Puplesis, genitalia slide no. AG 402 (ZIN).

    6. Stigmella sakhalinella Puplesis, 1984a: 115, 116.Morphology update (Figs 17, 18): Valva pointed apically (Figs 17, 19). Uncus with lateral lobes (Fig. 18). Aedeagus

    with apical cornuti (Fig. 17).Material examined: 1 ♂, neotype (designated in Puplesis 1994:75–76, fig. 177), Primorskiy Kray, 10 km S

    Slavyanka, Ryazanovka, 11.viii.1983, leg. R. Puplesis, genitalia slide no. AG 423 (ZIN); 1 ♂ (non-type), Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 14.vii.1982, leg. R. Puplesis, genitalia slide no. AG 424 (ZIN).

    7. Stigmella attenuata Puplesis 1985: 62.Morphology update (Figs 20, 21): Aedegus with apical cornuti. Uncus bilobed (Figs 20, 21).Material examined: 1 ♂, holotype, Primorskiy Kray, 14 km S Slavyanka, Ryazanovka, 11.vii.1983, leg. R. Puplesis,

    genitalia slide no. AG 417 (ZIN).

    STONIS & ROCIENE2 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • FIGURES 1–4. Male genitalia of Stigmella dissona (Puplesis) (Astigmella dissona Puplesis), holotype, slide no. AG 403 (ZIN). 1, general view; 2, ventral view; 3, dorsal view; 4, reconstruction. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 3EAST ASIA NEPTICULIDAE 1

  • FIGURES 5–8. Male genitalia of Stigmella mirabella (Puplesis) (Astigmella mirabella Puplesis), holotype, slide no. AG 404 (ZIN). 5, general view, 6, valvae; 7, dorsal view; 8, ventral view of aedeagus and gnathos. Scale bar 100 µm.

    STONIS & ROCIENE4 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • FIGURES 9–12. Male genitalia of Stigmella sashai Puplesis, holotype, slide no. AG 401 (ZIN). 9, general view (aedeagus removed); 10, gnathos and inner lobe of valvae; 11, dorsal view; 12, aedeagus. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 5EAST ASIA NEPTICULIDAE 1

  • FIGURES 13–16. Male genitalia of Stigmella regina Puplesis, holotype, slide no. AG 402 (ZIN). 13, general view; 14, valva; 15, ventral view, with two apical clusters of spine-like cornuti on vesica; 16, dorsal view. Scale bar 100 µm.

    STONIS & ROCIENE6 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • FIGURES 17–21. Male genitalia of Stigmella species. 17, S. sakhalinella Puplesis, general view, neotype, slide no. AG 423 (ZIN); 18, same, pseuduncus and gnathos; 19, valvae; 20, S. attenuata Puplesis, general view, holotype, slide no. AG 417 (ZIN); 21, same, reconstruction, slide no. AG 211 (non-type). Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 7EAST ASIA NEPTICULIDAE 1

  • 8. Stigmella ultima Puplesis, 1984a: 117.Morphology update (Figs 22, 23): Valva with papillae on inner lobe (Fig. 22). Cornuti small, mostly denticulate (Fig. 23).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Acer mono

    18.vi.1983, ex pupa 9.vii.1983, field card no. 3008, leg. R. Puplesis, genitalia slide no. AG 393 (ZIN).

    9. Stigmella tegmentosella Puplesis, 1984a: 117.Morphology update (Figs 24–26): Valva with papillae and spine-like process on inner lobe (Figs 25, 26). Cornuti

    small, mostly denticulate (Fig. 24).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Acer

    tegmentosum,13–28.vii.1983, ex pupa 24.vii.1983, field card no. 3015, leg. R. Puplesis, genitalia slide no. AG 394 (ZIN).

    FIGURES 22–26. Male genitalia of Stigmella species. 22, S. ultima Puplesis, general view (aedeagus removed), holotype, slide no. AG 393 (ZIN); 23, same, aedeagus; 24, aedeagus of S. tegmentosella Puplesis, holotype, slide no. AG 394 (ZIN); 25, same, capsule (aedeagus removed); 26, same, reconstruction. Scale bar 100 µm.

    STONIS & ROCIENE8 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • 10. Stigmella monella Puplesis, 1984a: 117, 118.Morphology update (Figs 27–31): Valva with undeveloped inner lobe and spine-like process (Figs 27–29). Transtilla

    interrupted medially. Cornuti minute, mostly denticulate (Fig. 31).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Acer

    mono11.vi.1983,ex pupa 24.vi.1983, field card no. 3001, leg. R. Puplesis, genitalia slide no. AG 392 (ZIN).

    FIGURES 27–31. Male genitalia of Stigmella monella Puplesis, holotype, slide no. AG 392 (ZIN). 27, general view (aedeagus removed); 28, dorsal view of apical part of valva; 29, ventral view of apical part of valva; 30, dorsal view of capsule; 31, aedeagus. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 9EAST ASIA NEPTICULIDAE 1

  • 11. Stigmella kozlovi Puplesis, 1984a: 118, 119.Morphology update (Figs 32–35): Valva with papillae on inner lobe (Figs 32, 33). Transtilla interrupted medially

    (Fig. 34). Cornuti minute, mostly denticulate (Fig. 35).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 22.v.1983, leg. R.

    Puplesis, genitalia slide no. AG 389 (ZIN).

    FIGURES 32–35. Male genitalia of Stigmella kozlovi Puplesis, holotype, slide no. AG 389 (ZIN). 32, general view (aedeagus removed); 33, ventral view; 34, dorsal view; 35, aedeagus. Scale bar 100 µm.

    STONIS & ROCIENE10 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • 12. Stigmella nostrata Puplesis, 1984a: 113, 114.Morphology update (Figs 36–40): Valva with large inner lobe (Figs 36, 37). Gnathos with relatively broad caudal

    processes (Fig. 38).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Pyrus

    ussuriensis 9.viii.1982, ex pupa 27.ii.1983, field card no. 2061, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2875 (ZIN); 1 ♂, paratype, same locality and host-plant, larva 24–27.vi.1983, ex pupa 14–16.vii.1983, field card no. 3012, leg. R. Puplesis, genitalia slide no. AG 400 (ZIN).

    FIGURES 36–40. Male genitalia of Stigmella nostrata Puplesis. 36, general view (aedeagus removed), holotype, slide no. E. J. van Nieukerken 2875 (ZIN); 37, same, ventral view; 38, same, dorsal view; 39, same, aedeagus; 40, aedeagus, paratype, slide no. AG 400 (ZIN). Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 11EAST ASIA NEPTICULIDAE 1

  • 13. Stigmella aurora Puplesis, 1984a: 119.Morphology update (Figs 41–44): Valva with triangular process on inner lobe (Fig. 43). Gnathos with large anterior

    processes (Figs 41–43). Cornuti small, denticulate, spine-like to irregular (Fig. 44).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Crataegus sp.

    1.vii.1982, ex pupa 18.vii.1982, field card no. 2018, leg. R. Puplesis, genitalia slide no. AG 397 (ZIN); 1 ♂ (non-type), Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Crataegus pinnatifida 18.vii.1982, ex pupa 3.iii.1983, field card no. 2040, leg. R. Puplesis, genitalia slide no. AG 396 (ZIN).

    FIGURES 41–44. Male genitalia of Stigmella aurora Puplesis, holotype, slide no. AG 397 (ZIN). 41, general view (aedeagus removed); 42, dorsal view; 43, reconstruction; 44, aedeagus. Scale bar 100 µm.

    STONIS & ROCIENE12 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • 14. Stigmella micromelis Puplesis, 1985: 59, 60.Morphology update (Figs 45–49): A more thorough examination of the male genitalia of the holotype reveals very

    little difference from that of the original description (Puplesis 1985: fig. 4 and Puplesis 1994: fig. 300).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Micromeles

    alnifolia 8–15.ix.1982, ex pupa 12.iv.1982, field card no. 2127, leg. R. Puplesis, genitalia slide no. AG 406 (ZIN).

    FIGURES 45–49. Male genitalia of Stigmella micromelis Puplesis, holotype, slide no. AG 406 (ZIN). 45, general view (aedeagus removed); 46, ventral view; 47, inner lobes of valvae; 48, aedeagus (broken); 49, same, other view to cornuti and cathrema. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 13EAST ASIA NEPTICULIDAE 1

  • 15. Stigmella crataegivora Puplesis, 1985: 60–62.Morphology update (Figs 50–53): A more thorough examination of the male genitalia of the holotype shows little

    difference from that of the original description (Puplesis 1985: fig. 3 and Puplesis 1994: fig. 297). Gnathos with larger caudal and anterior processes (Fig. 52).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Crataegus maximowiczi 17.ix.1982, ex pupa 3.iv.1983, field card no. 2149, leg. R. Puplesis, genitalia slide no. AG 407 (ZIN).

    FIGURES 50–53. Male genitalia of Stigmella crataegivora Puplesis, holotype, slide no. AG 407 (ZIN). 50, general view; 51, inner lobe of valvae; 52, gnathos; 53, dorsal view. Scale bar 100 µm.

    STONIS & ROCIENE14 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • 16. Stigmella taigae Puplesis, 1984a: 112, 113.Morphology update (Figs 54–56): Valva with papillae on inner lobe (Fig. 54). Aedeagus with larger clusters of

    spine-like cornuti (Figs 55, 56).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Rhamnus sp.

    25.vii.1983, ex pupa 5.viii.1983, field card no. 3025, leg. R. Puplesis, genitalia slide no. AG 405 (ZIN).

    FIGURES 54–59. Male genitalia of Stigmella species. 54, S. taigae Puplesis, general view (aedeagus removed), holotype, slide no. AG 405 (ZIN); 55, same, aedeagus and dorsal view of cornuti; 56, same, ventral view of cornuti; 57, S. amuriella Puplesis, general view (aedeagus removed), paratype, slide no. AG 409 (ZIN); 58, same, aedeagus, ventral view of cornuti and cathrema; 59, same, dorsal view. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 15EAST ASIA NEPTICULIDAE 1

  • 17. Stigmella palionisi Puplesis, 1984b: 596.Morphology update (Figs 65–67): Juxta developed, with narrow sclerotized apex (Fig. 65). Aedeagus with two

    lateral groups of larger, sclerotized cornuti and numerous tiny denticulate cornuti in between (Figs 65, 67).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 27.vii.1983, leg. R.

    Puplesis, genitalia slide no. AD 0362 (ZIN).

    18. Stigmella amuriella Puplesis, 1985: 62.Morphology update (Figs 57–59): Valva with rounded inner lobe (Fig. 57). Aedeagus with slender spine-like cornuti

    (Figs 58, 59).Material examined: 1 ♂, paratype, Primorskiy Kray, 20 km E Ussuriysk, 27.vii.1983, leg. R. Puplesis, genitalia slide

    no. AG 409 (ZIN).

    19. Stigmella alisa Puplesis 1985: 63.Morphology update (Figs 60–64): Juxta with narrow apex (Figs 60, 61). Gnathos with long anterior processes (Figs

    63, 64). Aedeagus with about 10 large cornuti and numerous small spine-like cornuti arranged in two lateral groups (Figs 60, 61).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 25.vii.1982, leg. R. Puplesis, genitalia slide no. AG 416 (ZIN).

    20. Stigmella auricularia Puplesis, Diškus & Juchnevič, 2003: 245, 246.Morphology update (Figs 68–75): Uncus with triangular lateral lobes (Figs 68, 70, 75). Ventral plate of vinculum

    with deep oval excavation (Figs 68, 71, 74, 75).Material examined: 1 ♂, holotype, Primorskiy Kray, 14 km S Slavyanka, Ryazanovka, 3.vii.1982, leg. R. Puplesis,

    genitalia slide no. AD 0366 (ZIN); 1 ♂, paratype, 15 km SW Slavyanka, 3.vii.1982, leg. S. Yu. Sinev, genitalia slide no. AD 0367 (ZIN).

    21. Stigmella ussurica Puplesis, 1987: 8–10; a junior syn. of Stigmella tranocrossa Kempermen & Wilkinson, 1985.Morphology update (Figs 76–79): Uncus with two large median and two smaller lateral lobes (76, 78). Aedeagus

    with three large apical cornuti and a compact group of 5 smaller basal cornuti (Figs 76, 79).Material examined: 1 ♂, paratype of Stigmella ussurica, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe,

    6.vii.1982, leg. R. Puplesis, genitalia slide no. AG 410 (ZIN).

    22. Stigmella uigurica Puplesis, 1985: 62, 63; a junior syn. of Stigmella continuella (Stainton, 1856).Morphology update (Figs 80–84): Lateral lobes of vinculum very long and narrow (Fig. 80). Valva with two short

    apical processes (Fig. 81). Aedeagus with numerous tiny denticulate cornuti and a few spine-like cornuti (Figs 82–84).Material examined: 1 ♂, holotype of Stigmella uigurica, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe,

    13.viii.1982, leg. R. Puplesis, genitalia slide no. AG 387 (ZIN).

    23. Stigmella magica Puplesis, 1985: 63–65; a junior syn. of Stigmella lediella (Schleich, 1867).Morphology update (Figs 85–88): Inner lobe of valvae with numerous papillae (Figs 85, 87). Apical process of valva

    very long and curved (Fig. 85). Aedeagus with about 7 apical and 7 basal cornuti (Fig. 86).Material examined: 1 ♂, holotype of Stigmella magica, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva

    on Rhododendron mucronulatum 6.x.1982, leg. R. Puplesis, genitalia slide no. AG 408 (ZIN).

    24. Stigmella rhododendri Puplesis, 1985: 65; a junior syn. of Stigmella lediella (Schleich, 1867).Note. This species was described from another host-plant (Rhododendron schlippenbachii Maxim.); male genialia

    with broad sublateral processes of transtilla and more numerous basal cornuti (Puplesis 1985). However, the holotype was not available for our study.

    25. Stigmella palmatae Puplesis, 1984a: 115.Morphology update (Figs 89, 90): Outer margin of valva sinuous, inner lobe papillated, apical process blunt (Fig.

    90). Aedeagus missing (lost).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Filipendula

    palmata 13.viii.1982, ex pupa 12.iii.1983, field card no. 2063, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2854 (ZIN).

    26. Stigmella monticulella Puplesis, 1984a: 114.Morphology update (Figs 91–93): Valva with slender apical process (Fig. 91). Aedeagus with manica (Figs 92, 93).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Lonicera sp.

    27.ix.1982, ex pupa 19.iii.1983, field card no. 2187, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2870 (ZIN).

    STONIS & ROCIENE16 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • FIGURES 60–64. Male genitalia of Stigmella alisa Puplesis, holotype, slide no. AG 416 (ZIN). 60, valvae, juxta and aedeagus; 61, same, other (more ventral) view; 62, uncus, gnathos and vinculum; 63, gnathos; 64, uncus. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 17EAST ASIA NEPTICULIDAE 1

  • FIGURES 65–70. Male genitalia of Stigmella species. 65, S. palionisi Puplesis, holotype, slide no. AD 0362 (ZIN); 66, same, inner lobe of valva; 67, same, cornuti and transtilla; 68, S. auricularia Puplesis, Diškus & Juchnevič, general view (aedeagus removed), holotype, slide no. AD 0366 (ZIN); 69, same, right valva; 70, same, left valva. Scale bar 100 µm.

    STONIS & ROCIENE18 · Zootaxa 3652 (1) © 2013 Magnolia Press

  • FIGURES 71–75. Male genitalia of Stigmella auricularia Puplesis, Diškus & Juchnevič, holotype, slide no. AD 0366 (ZIN). 71, general view (aedeagus removed), holotype, slide no. AD 0366 (ZIN); 72, same, aedeagus; 73, same, more ventral view of cornuti; 74, same, capsule, dorsal view; 75, paratype, slide no. AD 0367 (ZIN). Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 19EAST ASIA NEPTICULIDAE 1

  • FIGURES 76–79. Male genitalia of Stigmella ussurica Puplesis (a junior syn. of Stigmella tranocrossa Kempermen & Wilkinson), paratype, slide no. AG 410 (ZIN). 76, general view; 77, other view with inner lobe of valva; 78, view with vinculum and uncus exposed; 79, aedeagus. Scale bar 100 µm.

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  • FIGURES 80–84. Male genitalia of Stigmella uigurica Puplesis (a junior syn. of Stigmella continuella (Stainton)), holotype, slide no. AG 384 (ZIN). 80, capsule (aedeagus and valvae removed); 81, valva; 82–84, various views of aedeagus and cornuti. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 21EAST ASIA NEPTICULIDAE 1

  • FIGURES 85–88. Male genitalia of Stigmella magica Puplesis (a junior syn. of Stigmella lediella (Schleich)), holotype, slide no. AG 408 (ZIN). 85, general view (aedeagus removed); 86, aedeagus; 87, ventral view of capsule; 88, dorsal view of capsule. Scale bar 100 µm.

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  • FIGURES 89–93. Male genitalia of Stigmella species. 89, S. palmatae Puplesis, general view (valvae removed, aedeagus lost), holotype, slide no. E. J. van Nieukerken 2854 (ZIN); 90, valvae and transtilla; 91, S. monticulella Puplesis, general view (aedeagus removed), holotype, slide no. E. J. van Nieukerken 2870 (ZIN); 92, aedeagus; 93, same, other view. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 23EAST ASIA NEPTICULIDAE 1

  • 27. Stigmella dentatae Puplesis, 1984a: 114, 115.Morphology update (Figs 94, 95): Apical cornuti large, spine-like. Uncus may be concealed in slide-mounted

    preparations (Fig. 95).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 29.v.1983, leg. R.

    Puplesis, genitalia slide no. AG 399 (ZIN).

    FIGURES 94–97. Male genitalia of Stigmella species. 94, S. dentatae Puplesis, general view, holotype, slide no. AG 399 (ZIN); 95, same, valvae; 96, S. omelkoi Puplesis, general view, holotype, slide no. AG 398 (ZIN); 97, valva. Scale bar 100 µm.

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  • 28. Stigmella aladina Puplesis, 1984a: 115.Morphology update (Figs 98–101): Valva with rounded inner lobe (Figs 98, 100). Aedeagus with a large apical

    group of spine-like cornuti (Fig. 99).Material examined: 1 ♂, holotype, Primorskiy Kray, Spassk-Dalny, larva on Quercus mongolica 18.vii.1983, ex

    pupa 4.viii.1983, field card no. 3020, leg. R. Puplesis, genitalia slide no. AG 411 (ZIN).

    FIGURES 98–101. Male genitalia of Stigmella aladina Puplesis, holotype, slide no. AG 411 (ZIN). 98, general view (aedeagus removed); 99, aedeagus; 100, ventral view of capsule; 101, dorsal view of capsule. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 25EAST ASIA NEPTICULIDAE 1

  • 29. Stigmella omelkoi Puplesis, 1984b: 593.Morphology update (Figs 96, 97): Uncus appearing short if bent downward in preparations (Fig. 96). Uncus with

    broad lateral processes (Fig. 97). Valva with rounded inner lobe (Fig. 97). Aedeagus with very large manica (Fig. 96).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Quercus

    mongolica 8.ix.1982, ex pupa 4.iii.1983, field card no. 2135, leg. R. Puplesis, genitalia slide no. AG 398 (ZIN).

    30. Stigmella fervida Puplesis, 1984b: 593, 594.Morphology update (Figs 102–105): Lateral processes of gnathos broad basally (Figs 102, 104). Ventral plate of

    vinculum broad medially, shorter laterally (Fig. 104). Aedeagus with large, lobe-like apical cornutus and numerous spine-like cornuti (Figs 103, 105).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 3.vii.1982, leg. R. Puplesis, genitalia slide no. AG 391 (ZIN).

    31. Stigmella kurokoi Puplesis, 1984b: 594.Morphology update (Figs 106–109): Inner lobe of valvae heavily papillated (Fig. 107), apical process pointed (Fig.

    108). Aedeagus with manica (Fig. 109).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 10.viii.1982, leg. R.

    Puplesis, genitalia slide no. AG 390 (ZIN).

    32. Bohemannia manschurella Puplesis, 1984b: 587.Morphology update (Figs 110–113): Process of juxta bifurcate distally (Figs 110, 112). Aedeagus with one large,

    spine-like, apical cornutus (Fig. 113). Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 21.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 379 (ZIN).

    33. Bohemannia ussuriella Puplesis, 1984b: 588.Morphology update (Figs 114–117): Juxta with pointed lateral lobes (Figs 114, 116, 117). Aedeagus with large

    spine-like apical cornuti (Fig. 117).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 5.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 378 (ZIN).

    34. Bohemannia suiphunella Puplesis, 1984b: 588.Morphology update (Figs 118–121): Ventral plate of vinculum very short and rounded (Figs 118, 119). Aedeagus

    with apical group of spine-like processes (Figs 120, 121).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 5.vii.1982, leg. R.

    Puplesis, genitalia slide no. AD 0360 (ZIN); 1 ♂ (non-type), same locality, 19.vii.1983, leg. R. Puplesis, genitalia slide no. AD 0363 (ZIN).

    35. Bohemannia nubila Puplesis, 1985: 69–71.Morphology update (Figs 122–127): Valva almost triangular, with pointed apical process and papillated inner lobe

    (Figs 123, 124). Ventral plate of vinculum narrowed distally (Figs 122, 125).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 27.viii.1982, leg. R.

    Puplesis, genitalia slide no. AG 377 (ZIN).

    36. Bohemannia piotra Puplesis, 1984b: 586, 587.Morphology update (Figs 128–131): Valva broad (Fig. 129), with very slender apical process (Figs 128, 131).

    Aedeagus with spine-like cornuti (Figs 128, 130).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Malus

    manschurica 1.vii.1983, field card no. 3006, leg. R. Puplesis, genitalia slide no. AG 383 (ZIN).

    37. Ectoedemia emendata Puplesis, 1985: 69; a junior syn. of Ectoedemia amani Svensson, 1966: 200, 201, fig. 34, pl. 4, fig. 3.

    Remarks. The holotype of E. emendata was not available for our study.

    38. Ectoedemia admiranda Puplesis, 1984b: 588, 590.Morphology update (Figs 132–139): Valva abruptly narrowed in apical part (Figs 132–134). Aedeagus with three

    pairs of carinae (Figs 136, 137, 139), numerous tiny cornuti (Fig. 137), and long band of spine-like cornuti (Fig. 138).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 13.viii.1982, leg. R.

    Puplesis, genitalia slide no. AG 362 (ZIN).

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  • FIGURES 102–105. Male genitalia of Stigmella fervida Puplesis, holotype, slide no. AG 391 (ZIN). 102, general view (aedeagus removed); 103, aedeagus; 104, ventral view of capsule; 105, ventral view of cornuti. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 27EAST ASIA NEPTICULIDAE 1

  • FIGURES 106–109. Male genitalia of Stigmella kurokoi Puplesis, holotype, slide no. AG 390 (ZIN). 106, general view (aedeagus removed); 107, inner lobe of valva; 108, other view of capsule with pointed apical process of valva; 109, aedeagus. Scale bar 100 µm.

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  • FIGURES 110–113. Male genitalia of Bohemannia manschurella Puplesis, holotype, slide no. AG 379 (ZIN). 110, general view; 111–112, focused from ventral to more dorsal; 113, transtilla and dorsal view of aedeagus. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 29EAST ASIA NEPTICULIDAE 1

  • FIGURES 114–117. Male genitalia of Bohemannia ussuriella Puplesis, holotype, slide no. AG 378 (ZIN). 114, general view; 115, valva; 116, other view of capsule; 117, ventral view of aedeagus. Scale bar 100 µm.

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  • FIGURES 118–121. Male genitalia of Bohemannia suiphunella Puplesis, holotype, slide no. AD 0360 (ZIN). 118, capsule (valvae and aedeagus removed); 119, general view in glycerol; 120, valva and aedeagus, ventral view; 121, same, dorsal view. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 31EAST ASIA NEPTICULIDAE 1

  • FIGURES 122–127. Male genitalia of Bohemannia nubila Puplesis, holotype, slide no. AG 377 (ZIN). 122, general view; 123, 124, valva; 125, dorsal view; 126, distal part of genitalia; 127, transtilla. Scale bar 100 µm.

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  • FIGURES 128–131. Male genitalia of Bohemannia piotra Puplesis, holotype, slide no. AG 383 (ZIN). 128, general view; 129–131, focused from ventral to more dorsal. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 33EAST ASIA NEPTICULIDAE 1

  • FIGURES 132–135. Male genitalia of Ectoedemia admiranda Puplesis, holotype, slide no. AG 362 (ZIN). 132, general view; 133–134, apical part of genitalia, focused from ventral to more dorsal; 135, transtilla and dorsal carinae of aedeagus. Scale bar 100 µm.

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  • FIGURES 136–139. Apical part of aedeagus of Ectoedemia admiranda Puplesis, holotype, slide no. AG 362 (ZIN), focused from ventral to more dorsal. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 35EAST ASIA NEPTICULIDAE 1

  • 39. Ectoedemia sivickisi Puplesis, 1984b: 590.Morphology update (Figs 140–143): Sublateral processes of transtilla very long (Figs 140, 143). Aedeagus with

    three pairs of carinae (Figs 141–143) and numerous tiny cornuti (Figs 140, 142).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 08.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 363 (ZIN).

    FIGURES 140–143. Male genitalia of Ectoedemia sivickisi Puplesis, holotype, slide no. AG 363 (ZIN). 140, general view; 141–143, focused from ventral to more dorsal. Scale bar 100 µm.

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  • 40. Ectoedemia laura Puplesis, 1985: 68, 69.Morphology update (Figs 144–148): Sublateral processes of transtilla very long (Figs 145, 146). Aedeagus with

    three pairs of carinae (Figs 146–148) and numerous tiny cornuti on vesica (everted in the holotype) (Figs 145, 147).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 25.vii.1983, leg. R.

    Puplesis, genitalia slide no. AG 364 (ZIN).

    FIGURES 144–148. Male genitalia of Ectoedemia laura Puplesis, holotype, slide no. AG 364 (ZIN). 144, general view (valvae and aedeagus removed); 145, valvae and aedeagus; 146, dorsal view of transtilla and carinae; 147–148, aedeagus with everted vesica, focused from dorsal to more ventral. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 37EAST ASIA NEPTICULIDAE 1

  • 41. Ectoedemia arisi Puplesis, 1984a: 120–122.Morphology update (Figs 149–152): Valva papillated (Figs 149, 150). Aedeagus with single, very large dorsal carina

    (Figs 151, 152).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 23.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 365 (ZIN).

    FIGURES 149–152. Male genitalia of Ectoedemia arisi Puplesis, holotype, slide no. AG 365 (ZIN). 149–150, focused from dorsal to more ventral; 151–152, transtilla and aedeagus, focused from ventral to more dorsal. Scale bar 100 µm.

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  • 42. Ectoedemia christopheri Puplesis, 1985: 69 (originally described as Ectoedemia wilkinsoni Puplesis, 1984a: 122, a junior homonym of Ectoedemia wilkinsoni Scoble, 1983.

    Morphology update (Figs 153–160): Caudal process of gnathos strongly sclerotized and appearing truncate at apex (Fig. 155). Valvae with numerous papillae and long setae (Fig. 155). Transverse bar of transtilla thickened. Aedeagus with narrow band of spines between lateral carinae (Fig. 156); two pairs of carinae (lateral and ventral) pointed distally (Figs 157–160), except single sublateral lobe (sublateral carina) (Fig. 160).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 25.vii.1983, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2869 (ZIN); 1 ♂, paratype, same locality, 17.vii.1983, leg. R. Puplesis, genitalia slide no. AG 366 (ZIN).

    FIGURES 153–156. Male genitalia of Ectoedemia christopheri Puplesis 153, general view, holotype, slide no. E. J. van Nieukerken 2869 (ZIN); 154, same, ventral view; 155, same, apical part of genitalia; 156, dorsal view of transtilla, paratype, slide no. AG 366 (ZIN). Scale bar 100 µm.

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  • FIGURES 157–160. Apical part of aedeagus of Ectoedemia christopheri Puplesis. 157,159, holotype, slide no. E. J. van Nieukerken 2869 (ZIN); 158, 160, paratype, slide no. AG 366 (ZIN). Scale bar 100 µm.

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  • 43. Ectoedemia philipi Puplesis, 1984b: 590, 592.Morphology update (Figs 161, 162): Gnathos with elaborate and strongly slerotized lateral arms (Fig. 162).

    Aedeagus rounded apically, with two pairs of spine-like carinae (Figs 161, 162).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 19.vii.1982, leg. R.

    Puplesis, genitalia slide no. AG 367 (ZIN).

    FIGURES 161–164. Male genitalia of Ectoedemia species. 161, E. philipi Puplesis, holotype, general view, slide no. AG 367 (ZIN); 162, same, gnathos and apical part of aedeagus; 163, E. chasanella Puplesis, holotype, general view, slide no. AD 0361 (ZIN); 164, same, ventral view of capsule. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 41EAST ASIA NEPTICULIDAE 1

  • 44. Ectoedemia chasanella Puplesis, 1984a: 124.Morphology update (Figs 163–168): Caudal process of gnathos narrowly (Figs 163, 164) to broadly rounded distally

    (Fig. 166). Aedeagus with two triangular ventral carinae and some minute cornuti on vesica (everted in Figs 167, 168).Material examined: 1 ♂, holotype, Primorskiy Kray, 15 km SW Slavyanka, Ryazanovka, 06.viii.1983, leg. R.

    Puplesis, genitalia slide no. AD 0361 (ZIN); 1 ♂, paratype, same label data, genitalia slide no. AG 368 (ZIN).

    FIGURES 165–168. Male genitalia of Ectoedemia chasanella Puplesis 165, general view with focus on aedeagus, holotype, slide no. AD 0361 (ZIN); 166, capsule (valvae and aedeagus removed), paratype, slide no. AG 368 (ZIN); 167, 168, same, valvae and aedeagus, focused from ventral to more dorsal. Scale bar 100 µm.

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  • 45. Ectoedemia scoblei Puplesis, 1984a: 122.Morphology update (Figs 169–172): Valva heavily papillated on inner margin (Figs 169, 170). Lateral arms of

    gnathos very long and broadened distally (Figs 169, 171). Aedeagus with two ventral carinae (Figs 171, 172).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 6.vii.1983, leg. R.

    Puplesis, genitalia slide no. AG 375 (ZIN).

    FIGURES 169–172. Male genitalia of Ectoedemia scoblei Puplesis, holotype, slide no. AG 375 (ZIN). 169, general view; 170–172, focused from ventral to more dorsal. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 43EAST ASIA NEPTICULIDAE 1

  • 46. Ectoedemia aligera Puplesis, 1985: 67, 68.Morphology update (Figs 173–178): Inner margin of valva papillated (Figs 174, 178). Lateral arms of gnathos long

    and broadened distally (Fig. 175). Ventral carinae of aedeagus comprise two (Fig. 175) to five slender spine-like processes (Fig. 177).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 9.vii.1983, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2874 (ZIN); 1 ♂, paratype, same label data, genitalia slide no. AG 369 (ZIN).

    FIGURES 173–176. Male genitalia of Ectoedemia aligera Puplesis, holotype, slide no. E. J. van Nieukerken 2874 (ZIN). 173, general view; 174–176, focused from ventral to more dorsal. Scale bar 100 µm.

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  • 47. Ectoedemia ermolaevi Puplesis, 1985: 68.Morphology update (Figs 179, 180): Valva with strongly slerotized, pointed apical process (Fig. 179). Aedeagus

    with two ventral carinae (Fig. 180).Material examined: 1 ♂, holotype, Primorskiy Kray (Southern Primor‘e), Kedrovaya Pad‘ Nature Reserve,

    19.vii.1974, leg. Ermolaev, genitalia slide no. E. J. van Nieukerken 2873 (ZIN).

    FIGURES 177–180. Male genitalia of Ectoedemia species. 177, E. aligera Puplesis, general view, paratype, slide no. AG 369 (ZIN); 178, same, ventral view; 179, E. ermolaevi Puplesis, general view, holotype, slide no. E. J. van Nieukerken 2873 (ZIN); 180, same, ventral view. Scale bar 100 µm.

    Zootaxa 3652 (1) © 2013 Magnolia Press · 45EAST ASIA NEPTICULIDAE 1

  • 48. Ectoedemia maculata Puplesis, 1987: 11, 12.Morphology update (Figs 181, 182): Apical process of valva triangular (Fig. 181). Pseuduncus very large, broadly

    rounded (partially bent in Fig. 182).Material examined: 1 ♂, holotype, Primorskiy Kray, Khasan District, Ryazanovka, 3.vii.1982, leg. M. V. Kozlov,

    genitalia slide no. AG 374 (ZIN).

    FIGURES 181–185. Male genitalia of Ectoedemia species. 181, E. maculata Puplesis, general view, holotype, slide no. AG 374 (ZIN); 182, same, ventral view of capsule; 183, E. olvina Puplesis, general view, holotype, slide no. E. J. van Nieukerken 2871 (ZIN); 184, same, valva; 185, same, aedeagus. Scale bar 100 µm.

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  • 49. Ectoedemia olvina Puplesis, 1984a: 119, 120.Morphology update (Figs 183–185): Valva papillated on inner margin (Fig. 184). Aedeagus with two ventral carinae

    (Fig. 185).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, 26.vii.1983, leg. R. Puplesis, genitalia slide

    no. E. J. van Nieukerken 2871 (ZIN).

    50. Ectoedemia ornatella Puplesis, 1984a: 120.Morphology update (Figs 186–190): Valva papillated on inner margin and apical process (Fig. 187). Aedeagus with

    two ventral carinae (Figs 188, 189) and some minute cornuti on vesica (Fig. 190).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 27.vii.1983, leg. R.

    Puplesis, genitalia slide no. AG 371 (ZIN).

    51. Ectoedemia ivinskisi Puplesis, 1984a: 120.Morphology update (Figs 191, 192): Apical process of valva heavily papillated (Fig. 192). Gnathos without median

    excavation or processes (Figs 191, 192).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 10.vii.1983, leg. R.

    Puplesis, genitalia slide no. AG 373 (ZIN).

    52. Ectoedemia pilosae Puplesis, 1984a: 123, 124.Remarks. The holotype was not available for our study.

    53. Ectoedemia picturata Puplesis, 1985: 65, 67.Morphology update (Figs 193, 194): Pseuduncus very long and broadly rounded distally (Fig. 193). Valva broad, with

    very slender apical process. Aedeagus with two divided ventral carinae and some minute cornuti on vesica (Fig. 194).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, larva on Rosa rugosa,

    18.viii.1982, ex pupa 13.iv.1983, field card no. 2067, leg. R. Puplesis, genitalia slide no. AG 384 (ZIN).

    54. Ectoedemia sinevi Puplesis, 1985: 67.Morphology update (Figs 195–199): Valva gradually narrowed towards apex, with pointed apical process (Figs 195,

    198). Aedeagus with seven slender spine-like carinae (Figs 196, 197, 199).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 23.vi.1983. leg. S. Yu.

    Sinev, genitalia slide no. AG 376 (ZIN).

    55. Ectoedemia insularis Puplesis, 1985: 68.Morphology update (Figs 200–210): Valva with triangular apical process (Figs 201, 202, 207), heavily papillated

    (Figs 207, 208). Gnathos with large caudal process and broad lateral arms (Fig. 200). Aedeagus with two slender ventral and two lateral carinae (Figs 204, 205, 209, 210); minute cornuti on vesica (Fig. 210).

    Material examined: 1 ♂, holotype, Sakhalin Isle, Yuzhno-Sakhalinsk. env., 8.vii.1983, S. Yu. Sinev, genitalia slide no. E. J. van Nieukerken 2872 (ZIN); 1 ♂, paratype, same locality, 30.vi.1983, leg. A. Sharkov, genitalia slide no. AG 370 (ZIN).56. Fomoria permira Puplesis, 1984b: 592, 593.

    Morphology update (Figs 211–222): Aedeagus with four very long and slender cornuti (Figs 220, 222), two very long carinae (Figs 214–216, 220), and two short, distally pointed carinae (Figs 221, 222). Cathrema well developed, its pointed sublateral corners can be misstaken for cornuti (Figs 214, 218, 220).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 24.vii.1982, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2969 (ZIN); 1 ♂, paratype, same locality, 27.vii.1982, leg. R. Puplesis, genitalia slide no. AG 385 (ZIN).

    57. Etainia capesella (Puplesis, 1985) (Obrussa capesella Puplesis, 1985 in Puplesis & Ivinskis 1985: 39, 40).Morphology update (Figs 223–226): Inner margin of valva sinuous. Gnathos with two short triangular caudal

    processes (Fig. 225). Aedeagus with two very large apical cornuti and two apical lobe-like carinae; cathrema well developed, complex, and very large (Fig. 224).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 25.viii.1982, leg. R. Puplesis, genitalia slide no. AG 380 (ZIN).

    58. Etainia tigrinella (Puplesis, 1985) (Obrussa tigrinella Puplesis, 1985 in Puplesis & Ivinskis 1985: 40, 41).Morphology update (Figs 227, 228): Pseuduncus distinctly truncate (Fig. 227). Aedeagus with one very large, basally

    lobed, distally pointed cornutus and four apical carinae; cathrema well developed, complex, and very large (Fig. 224).Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 8.viii.1982, leg. R.

    Puplesis, genitalia slide no. AG 382 (ZIN).

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  • FIGURES 186–190. Male genitalia of Ectoedemia ornatella Puplesis, holotype, slide no. AG 371 (ZIN). 186, genitalia capsule (valvae and aedeagus removed); 187, valva; 188, general view of aedeagus; 189,190, same, focused from ventral to more dorsal. Scale bar 100 µm.

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  • FIGURES 191–194. Male genitalia of Ectoedemia species. 191, E. ivinskisi Puplesis, general view, holotype, slide no. AG 373 (ZIN); 192, same, ventral view; 193, E. picturata Puplesis, capsule (valvae and aedeagus removed), holotype, slide no. AG 384 (ZIN); 194, same, valvae and aedeagus. Scale bar 100 µm.

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  • FIGURES 195–199. Male genitalia of Ectoedemia sinevi Puplesis, holotype, slide no. AG 376 (ZIN). 195, general view; 196, 197, aedeagus, focused from ventral to more dorsal; 198, reconstruction; 199, same, apical part of aedeagus. Scale bar 100 µm.

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  • FIGURES 200–205. Male genitalia of Ectoedemia insularis Puplesis, holotype, slide no. E. J. van Nieukerken 2872 (ZIN). 200, general view (aedeagus removed); 201, valva, ventral view; 202, same, dorsal view; 203, dorsal view of capsule; 204, 205, aedeagus. Scale bar 100 µm.

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  • FIGURES 206–210. Male genitalia of Ectoedemia insularis Puplesis, paratype, slide no. AG 370 (ZIN). 206, general view; 207, 208, focused from ventral to more dorsal; 209, aedeagus, focus on ventral carinae; 210, same, focus on lateral carinae and minute cornuti on vesica. Scale bar 100 µm.

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  • FIGURES 211–216. Male genitalia of Fomoria permira Puplesis 211, capsule (valvae and aedeagus removed), holotype, slide no. E. J. van Nieukerken 2969 (ZIN); 212, same, ventral view of valva; 213, same, dorsal view of valva with dorsal spine exposed; 214, general view, paratype, slide no. AG 385 (ZIN); 215, aedeagus, holotype, slide no. E. J. van Nieukerken 2969 (ZIN); 216, same, other focus on cornuti and carinae. Scale bar 100 µm.

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  • FIGURES 217–222. Male genitalia of Fomoria permira Puplesis, paratype, slide no. AG 385 (ZIN). 217, 218, general view, focused from ventral to more dorsal; 219–222, aedeagus, focused from ventral to more dorsal. Scale bar 100 µm.

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  • FIGURES 223–226. Male genitalia of Etainia capesella (Puplesis) (Obrussa capesella Puplesis), holotype, slide no. AG 380 (ZIN). 223, general view (aedeagus removed); 224, aedeagus; 225, valvae, with inner margin exposed; 226, same, with apical processes exposed. Scale bar 100 µm.

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  • FIGURES 227–230. Male genitalia of Etainia species. 227, Etainia tigrinella (Puplesis) (Obrussa tigrinella Puplesis), general view, holotype, slide no. AG 382 (ZIN); 228, same, apical part of aedeagus; 229, Etainia sabina (Puplesis) (Obrussa sabina Puplesis), general view, holotype, slide no. AG 381 (ZIN); 230, same, aedeagus. Scale bar 100 µm.

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  • 59. Etainia peterseni (Puplesis, 1985) (Obrussa peterseni Puplesis, 1985 in Puplesis & Ivinskis 1985: 41–43).Morphology update (Figs 231–235): Valva with slender pointed inner process (Fig. 232). Aedeagus with three very

    large and one small cornuti, and two strongly sclerotized lateral and one lobe-like, median carinae; cathrema well developed, complex, and very large (Fig. 230).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 18.vii.1982, leg. R. Puplesis, genitalia slide no. E. J. van Nieukerken 2876 (ZIN); 1 ♂, paratype, same locality, 27.viii.1982, leg. R. Puplesis, genitalia slide no. AG 388 (ZIN).

    FIGURES 231–235. Male genitalia of Etainia peterseni (Puplesis) (Obrussa peterseni Puplesis. 231, general view (aedeagus removed), holotype, slide no. E. J. van Nieukerken 2876 (ZIN); 232, valva, paratype, slide no. AG 388 (ZIN); 233, same, general view (aedeagus removed); 234, aedeagus, holotype, slide no. E. J. van Nieukerken 2876 (ZIN); 235, aedeagus, paratype, slide no. AG 388 (ZIN). Scale bar 100 µm.

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  • 60. Etainia sabina (Puplesis, 1985) (Obrussa sabina Puplesis, 1985 in Puplesis & Ivinskis 1985: 43, 44).Morphology update (Figs 229, 230): Valva without apical process (Fig. 229). Aedeagus with two very large and two

    smaller apical cornuti and two lateral, distally sclerotized and pointed carinae; cathrema well developed, complex, and very large (Fig. 230).

    Material examined: 1 ♂, holotype, Primorskiy Kray, 20 km E Ussuriysk, Gornotayezhnoe, 10.viii.1982, leg. R. Puplesis, genitalia slide no. AG 381 (ZIN).

    Acknowledgements

    Dr. Sergey Yu. Sinev (Zoological Institute of the Russian Academy of Sciences) and Dr. Erik J. van Nieukerken (Naturalis, Leiden) provided the initial stimulus for this project. We thank Prof. Dr. Virginijus Sruoga (Lithuanian University of Educational Sciences, Vilnius) for his assistance during the project. We are very grateful to Dr. Erik J. van Nieukerken, Dr. Sergey Yu. Sinev and one anonymous referee for reviewing this paper and their useful suggestions.

    This study was conducted as part of the “New Faunas” project of the Division of Biosystematics Research of Lithuanian University of Educational Sciences with support from the Research Foundation of the Research Council of Lithuania (MIP-049/2011).

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    (* Stonis, J. R., formerly Puplesis, R.)

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    AbstractIntroductionMaterial and methodsPhotographic documentation of the types of Nepticulidae from East Asia described by R. PuplesisAcknowledgementsReferences