LITHUANIAN UNIVERSITY OF EDUCOLOGICAL SCIENCES FACULTY OF NATURAL SCIENCES, MATHEMATICS AND TECHNOLOGIES DEPARTMENT OF BIOLOGY

ANDRIUS REMEIKIS

NEW DATA ON AND GEOGRAPHICAL DISTRIBUTION OF NEPTICULOIDEA () OF THE NEOTROPICAL REGION, CRIMEA AND HIMALAYA MASTER‘S THESIS (Biomedical Sciences, Zoology, 05B)

Scientific supervisors:

Prof. Habil. Dr. Jonas R. Stonis, Biomedical Sciences, Zoology (05B), Biosystematics Research Group, Department of Biology, Lithuanian University of Educational Sciences (Vilnius, Lithuania); Dr. Donald R. Davis, Biomedical Sciences, Zoology (05B), Research Entomologist of USNM, Smithsonian Institution (Washington D.C., U.S.A.)

Vilnius, 2013

CONTENTS

1. INTRODUCTION ...... 4 1.1. RELEVANCE OF THE STUDY ...... 4 1.2. NOVELTY OF THE STUDY ...... 5 1.3. PRESENTATION AND APPROVAL OF THE RESULTS ...... 5 1.3.1. National and international conferences and workshops...... 5 1.3.2. Publications...... 7 1.4. ACKNOWLEDGEMENTS ...... 9 2. AND (NEPTICULOIDEA) (LITERATURE REVIEW) ...... 11 2.1. SOME PARTICULARITIES OF NEPTICULOIDS ...... 11 2.1.1. Family Opostegidae ...... 11 2.1.2. Family Nepticulidae ...... 14 2.2. STUDY HISTORY ...... 25 2.2.1. Studies of Opostegidae of Central and South America ...... 25 2.2.2. Studies of Nepticulidae of Europe and other regions of the world ...... 26 2.2.3. Studies of Nepticulidae of Central and South America ...... 27 3. GOAL AND OBJECTIVES ...... 30 4. MATERIAL AND METHODS ...... 31 4.1. SCIENTIFIC MATERIAL EXAMINED ...... 31 4.2. METHODS ...... 35 5. RESULTS ...... 39 5.1. NEW DATA TO THE NEOTROPICAL FAUNA OF NEPTICULOIDEA ...... 39 5.1.1. Contribution to the fauna of Opostegidae of the Neotropical Region ...... 39 5.1.1.1. New species of Opostegidae from Costa Rica and Mexico ...... 39 5.1.1.2. Geographical distribution of Central American Opostegidae ...... 45 5.1.1.3. New distribution data of the Neotropical Opostegidae ...... 51 5.1.2. Nepticulidae of Central and South America ...... 55 5.2. NEW DATA TO THE SOUTH EASTERN EUROPEAN FAUNA OF THE NEPTICULIDAE .... 62 5.2.1. The Easternmost record of a Sub-mediterranean species Acalyptris platani in Europe ...... 62 5.2.1.1. European Acalyptris as representatives of the Mediterranean and Sub-mediterranean fauna ...... 62 5.2.1.2. Documentation of Acalyptris platani specimens collected in Karadag Nature Reserve ..... 63 5.2.1.3. Discussion about Acalyptris platani in Karadag ...... 68 5.2.2. A re-assessment of the Crimean fauna of Nepticulidae, with new records for the peninsula ...... 68 5.3. CONTRIBUTION TO THE FAUNA OF NEPTICULOIDEA OF THE HIMALAYA ...... 70

2 5.3.1. A re-assessment of Opostegidae of the Himalaya, with description of new species ...... 70 5.3.2. New Nepticulidae discoveries in the Himalaya ...... 80 5.4. THE MOST RECENT DISCOVERIES. DESCRIPTIONS OF NEW SPECIES FROM YUCATAN (MEXICO ONTRIBUTION TO THE FAUNA OF NEPTICULOIDEA OF THE HIMALAYA) ...... 82 5.4.1. maya Remeikis & Stonis, 2013 ...... 82 5.4.2. Acalyptris yucatani Remeikis & Stonis, 2013 ...... 84 5.4.3. Taxonomic discussion about recorded Nepticulidae species in Yucatan and adjascent regions ..... 87 6. CONCLUSIONS ...... 90 REFERENCES ...... 93 LITHUANIAN SUMMARY (SANTRAUKA) ...... 101

3 1. INTRODUCTION

1.1. RELEVANCE OF THE STUDY

One of the most fundamental challenges for mankind in the 21st century is to document the extent and distribution of global , and to understand the ecological processes that generate and maintain it (Stonis et al., 2010, 2013). Such information will be essential to inform and guide efforts to safeguard the natural ecosystems that provide earth’s life support systems in the face of escalating threats from human habitat destruction and modification (Stonis, 2010; Stonis et al., 2011, 2013). Growing international concern over the biodiversity crisis together with the provisions of the Rio Convention on Biological Diversity have imparted an urgency to the need for taxonomic revisions of diverse groups and the provision of identification manuals, particularly for most unstudied, remote or tropical areas (Puplesis, Robinson, 2000; Stonis et al., 2011). Hence this research project. Opostegidae and Nepticulidae (Nepticuloidea) are very specialized superfamilies of Lepidoptera (Insecta) with a worldwide distribution. The minute size of the adults, the concealed mining life-style of the larvae, and the difficulty of rearing imagines goes some way towards explaining why these are still poorly studied in many regions (Remeikis et al., 2009, Stonis ir kt., 2012). The major comprehensive goals of nepticuloid studies are taxonomic revisions and reviews of these tiny and intriguing , with documentation and description of new taxa. It also includes fieldwork in remote areas (e. g. the Himalaya or even southern Europe, the Crimea), a documentation of new data on larval bionomics and species diversity; a study of sizable collection material currently deposited at LEUS (received from BMNH, London, USNM Washington, ZMUC, Copenhagen and other world‘s leading biodiversity research centres), and description of new taxa from the Himalaya and the Neotropics; a re-assessment of the European, Himalayan and Neotropical fauna of the studied organisms to support new phylogenetic and faunogenetic concepts of the boreal and tropical biotas. The proposed studies can provide the wherewithal for users to identify representatives of groups for which there was previously no ‘entry-point’. Without the baseline data of providing names and means of identification for the species in a region, no one can go further and properly plan their conservation in the case of endemic species, or their control in the case of introduced

4 species damaging endemic vegetation, or recognise if a species is newly introduced and a potential pest. Also -mining have already proved outstandingly well suited for inquiries into a number of fundamental problems in functional and evolutionary ecology. The conducted studies have broadened knowledge of biodiversity of leaf-mining moths. The study is noteworthy, since few have studied these tiny moths, yet many species remain to be discovered and described. We hope that the present thesis and the reviews of regional faunas will stimulate further studies of these intriguing groups of insects – Opostegidae and Nepticulidae.

1.2. NOVELTY OF THE STUDY

As a result of specified field collecting of 2009–2013 in the Crimea, Himalaya, Guatemala (Central America) and Colombia (South America), and study of a collection material from Central and South America 4 new species of Opostegidae were described and 2 new species of Nepticulidae were discovered and 11 prepared for description (in co-authorship with Prof. Dr. J. R. Stonis, Dr. A. Diškus or Prof. Dr. J. Schuster, Guatemala). Updated checklists of Opostegidae of Central America and the Himalaya, with new data on geographical distribution of species were presented. Five species of Nepticulidae were recognized as new to the fauna of Crimea (South Eastern Europe) for the first time. New taxonomic data provided in this thesis make up 2% for Opostegidae of the world and 3.4%, 7.5%, 7.7%, 27% and 14% for Opostegidae of the Neotropical Region, Central America, Costa Rica, Mexico and the Himalaya, respectively. In Nepticulidae, new unpublished taxonomic data provided in this thesis make up 1.4% of the world fauna and 9.8% of the Neotropical fauna.

1.3. PRESENTATION AND APPROVAL OF THE RESULTS

1.3.1. National and international conferences and workshops

The results of the research were presented at these 8 coferences or workshops (in Lithuania and abroad):

1. Stonis J. R., Paulavičiūtė B., Sruoga V., Diškus A., Remeikis A., Šimkevičiūtė A., Navickaitė A. Naujos filogenetinės hipotezės apie augalus minuojančius primityvius Lepidoptera remiantis

5 molekuliniais duomenimis (Insecta: Lepidoptera: Nepticuloidea, , ). Ekosistemų vystymosi kryptys ir populiacijų filogeografijos molekuliniai tyrimai Baltijos regione. Lithuanian Academy of Sciences, Vilnius, 27 th October, 2009.

2. Šimkevičiūtė A., Remeikis A. „Mažųjų drugių (Lepidoptera) kolekcinių egzempliorių parengimas moksliniam tyrimui“. Gyvūnijos ir žmogaus tyrimų aktualijos. Scientific conference of bachelor and master students. Vilnius Pedagogical University, 30th April, 2009.

3. Remeikis A. „Nauji duomenys apie Centrinės Amerikos baltųjų gaubtagalvių (Insecta, Lepidoptera: Opostegidae) fauną“. Scientific conference of bachelor and master students. Vilnius Pedagogical University, 30th May, 2010.

4. Stonis J. R., Remeikis A., Sruoga V. Šimkevičiūtė A., Diškus A., Navickaitė A. „Indijoje per didįjį musoną. VPU Biosistematikos tyrimų grupės lauko darbai Himalajuose (Utarakandas, 2010 m.)“. Lietuvos Entomologų draugijos visuotinis susirinkimas, Vilnius University, 13th November, 2010.

5. Remeikis A., Stonis J. R., Diškus A., Davis D. R. Taxonomic diversity and geographical distribution of the Neotropical Opostegidae (Insecta: Lepidoptera). V International Conference of Naturalists “From Biotechnology to Environment Protection”, Zielona Gora, 25–27th November, 2010.

6. Stonis, J. R., Remeikis, A., Diškus, A. LEU biosistematikai ieškojo naujų faunų. Plenarinis pranešimas Lietuvos Entomologų draugijos suvažiavimae. Vilnius, 10th December, 2011.

7. Stonis, J. R., Remeikis, A., Diškus, A. Mokslui nežinomų rūšių paieškos Centrinės Amerikos faunoje. Pranešimas šalies IX-ojoje konferencijoje „Biologinė įvairovė: būklė, struktūra, apsauga, Vilnius, 16th November, 2012.

8. Remeikis A. Remeikis A. New data on species taxonomy and geographical distribution of Nepticuloidea (Lepidoptera) of the Neotropical Region, Crimea and Himalaya. Presentation of Master‘s thesis. Department meeting at Biology Department of Biology, Lithuanian University of Educational Sciences, May 28, 2013.

6 1.3.2. Publications

1.3.2.1. Scientific papers in journals with Impact factor, reviewed in Web of Science 1. Stonis, J. R., Remeikis, A., Sruoga, V. 2013. An annotated list of the Opostegidae of the Himalaya, with a description of Pseudopostega brevicaudata sp. n. (Lepidoptera: Nepticuloidea). Zootaxa, 3609(2): 182–194. ISSN 1175-5326 (print), ISSN 1175-5334 (online) 2. Stonis, J. R., Remeikis, A., Diškus, A., Noreika, R. 2013. New Nepticulidae species (Insecta, Lepidoptera) from the Yucatán Peninsula (SE Mexico). Zootaxa, 3609(2): 223–230. ISSN 1175- 5326 (print), ISSN 1175-5334 (online)

1.3.2.2. Scientific papers in ISI journals Abroad 3. Stonis, J. R., Navickaitė, A., Rocienė, A., Remeikis, A., Diškus, A. 2013. A provisional checklist of the Nepticulidae (Insecta, Lepidoptera) of the Crimea. Zoology and Ecology, 1–9 [Great Britain]. ISSN 2165-8005 (print) ISSN 2165-8013 (online) In Lithuania 4. Remeikis A., Stonis J. R., Diškus A., Davis D. R. 2009. Contribution to the Opostegidae fauna of Central America, with an updated checklist and description of new species from Costa Rica and Mexico (Insecta: Lepidoptera). Acta Zoologica Lituanica, 19 (4): 278–286. ISSN 1392-1657 5. Stonis, J. R., Remeikis, A., Auksoriūtė, A., Baužys, D., Vilkas, A. 2010. Sunorminti ir nauji Lietuvoje aptinkamų straubliuočių būrio, blakių pobūrio (Insecta: Hemiptera: Heteroptera) vabzdžių vardai. Acta Zoologica Lituanica, 20 (3): 262–272. ISSN 1392-1657 6. Stonis, J. R., Remeikis A. 2011. Acalyptris platani (Müller-Rutz) in Karadag, – the Easternmost Record of the Submediterranean Species in Europe (Insecta: Lepidoptera: Nepticulidae). Acta Zoologica Lituanica, 21 (2): 7–13. ISSN 1392-1657

1.3.2.3. Chapters in monographs 7. Stonis, J. R., Diškus, A., Navickaitė, A., Remeikis, A., Rocienė, A. 2012. Morfologiniai Nepticulidae požymiai, naudojami identifikuojant taksonus ir paremiantys šiuolaikinę sistemą, 60–72 p. In: Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Monograph. Lututė Publishers, Kaunas. ISBN 978-9955-37- 148-9

7 1.3.2.4. Publications in international conference materials 8. Remeikis, A., Stonis, J. R., Diškus, A., Davis, D., R. 2010. Taxonomic diversity and geographical distribution of the Neotropical Opostegidae (Insecta: Lepidoptera). V International Conference of Naturalists “From Biotechnology to Environment Protection”. 41 p. University of Zielona Gora Publishers, Zielona Gora.

1.3.2.5. Science popular publications 9. Stonis J. R., Remeikis A. 2009. Krymo dienoraštis (1). Atsiminimai apie žygius Karadago gamtos rezervate. Mokslas ir gyvenimas, 11: 36–39. ISSN 0134-3084 10. Stonis J. R., Remeikis A. 2009. Krymo dienoraštis (2). Atsiminimai apie žygius Karadago gamtos rezervate. Mokslas ir gyvenimas, 12: 40–43. ISSN 0134-3084 11. Stonis J. R., Remeikis A. 2009. Gamtininkų žygiai Karadago rezervate (1). Šviesa 17 (22715):2–3. 12. Stonis J. R., Remeikis A. 2009. Gamtininkų žygiai Karadago rezervate (2). Šviesa 18 (22716):2–3. 13. Stonis J. R., Remeikis A. 2009. Patikėkit, Karadahas ir jums patiktų. 1. Ukrainos gamtos perlas. Žurnalas apie gamtą, 6 (36): 12–15. ISSN 1648-8938 14. Stonis J. R., Remeikis A. 2010. Patikėkit, Karadahas ir jums patiktų. 2. Naktį išlenda įvairūs gyviai. Žurnalas apie gamtą, 1 (37): 14–18. ISSN 1648-8938 15. Stonis J. R., Remeikis A. 2010. Patikėkit, Karadahas ir jums patiktų. 3. Šaitano vartai ir vabzdžių rojus. Žurnalas apie gamtą, 2 (38): 14–18. ISSN 1648-8938 16. Stonis J. R., Remeikis A. 2010. Patikėkit, Karadahas ir jums patiktų. 4. Šaitano vartai ir vabzdžių rojus. Žurnalas apie gamtą, 3 (39): 16–21. ISSN 1648-8938 17. Stonis J. R., Remeikis A. 2010. Patikėkit, Karadahas ir jums patiktų. 5. Nuostabūs augalai ir visokios gyvatės. Žurnalas apie gamtą, 4 (40): 16–20. ISSN 1648-8938 18. Stonis J. R., Remeikis A. 2010.Iš Himalajų dienoraščio. Biosistematikos tyrimų grupės lauko darbai Indijoje. Šviesa, 11: 1: 6. 19. Stonis J. R., Remeikis A. 2010. Rišikeše norisi džiaugtis, tyrinėti, bet nenumirti. VPU Biosistematikos tyrimų grupės lauko darbai Indijoje. Žurnalas apie gamtą, 5 (41): 18–23. ISSN 1648-8938. 20. Stonis J. R., Remeikis A. 2010. Himalajų gamta ne tik nustebino, bet ir išgąsdino. VPU Biosistematikos tyrimų grupės lauko darbai Indijoje (2). Žurnalas apie gamtą, 6 (42): 14–19. ISSN 1648-8938 21. Stonis J. R., Remeikis A. 2010. Pabaisų apsuptyje. Ar jaunosios doktorantės tikrai nepabūgo nei Indijos gyvačių, nei kraujasiurbių dėlių. Mokslas ir gyvenimas, 11/12: 6–11.

8 22. Stonis J. R., Remeikis A. 2011.Iš Himalajų dienoraščio. Biosistematikos tyrimų grupės lauko darbai Indijoje. Šviesa, 1: 6. 23. Stonis J. R., Remeikis A. 2011. Himalajų dienoraštis. VPU Biosistematikos tyrimų grupės lauko darbaiIndijoje. Mokslas ir gyvenimas, 1: 39–43. ISSN 0134-3084. 24. Stonis J. R., Remeikis A. 2011. Į Himalajus per musoną (3). VPU Biosistematikos tyrimų grupės lauko darbai Indijoje. Žurnalas apie gamtą, 1 (43): 16–21. ISSN 1648-8938. 25. Stonis J. R., Remeikis A. 2011. Į Himalajus per musoną (4). VPU Biosistematikos tyrimų grupės lauko darbai Indijoje. Žurnalas apie gamtą, 2 (44): 20–24. ISSN 1648-8938. 26. Stonis, J. R., Remeikis, A. 2011. Indijos dienoraštis. VPU biosistematikos tyrimų grupės lauko darbai Indijoje (5). Žurnalas apie gamtą, 3: 16–21. ISSN 1648-8938 27. Stonis, J. R., Remeikis, A. 2012. Atvykome į didžiosios gamtos katastrofos vietą. Žurnalas apie gamtą, 1: 18–20. ISSN 1648-8938 28. Stonis, J. R., Diškus, A., Remeikis, A. 2012. „Naujų faunų“ projektas nubloškė į pavojingą šalį. Žurnalas apie gamtą, 3: 17–19. ISSN 1648-8938 29. Stonis, J. R., Remeikis, A., Diškus, A. 2012. Skaitytojų gąsdinimai. Patarimai, jeigu keliausite arba tirsite Gvatemaloje. Žurnalas apie gamtą, 4: 19. ISSN 1648-8938 30. Stonis, J. R., Remeikis, A., Diškus, A. 2012. Gvatemalos dienoraštis. Gvatemalos gamtą nesunku įsimylėti, bet ar verta rizikuoti? Mokslas ir gyvenimas, 10: 22–25. ISSN 0134-3084 31. Stonis, J. R., Remeikis, A., Diškus, A. 2012. Gvatemalos dienoraštis. Šalies gamta nepakartojama, bet dar mažai ištirta Mokslas ir gyvenimas, 11/12: 42–48. ISSN 0134-3084

1.4. ACKNOWLEDGEMENTS

The work was started in 2009, but Master’s Thesis was carried out in 2011–2013 at Biosystematics Research Group of Biology Department of Lithuanian University of Educational Sciences. The scientific supervisors of the present thesis Prof. Habil. Dr. Jonas Rimantas Stonis (Head of Biosystematics Research Group, LEU, Vilnius) and Dr. Donald R. Davis (Leading researcher of Entomology Department, USNM, Washington D.C.) provided the initial stimulus for the study together with generous and enormous support during its course. We also thank Dr. Donald R. Davis (USNM, Washington), Prof. Dr. Niels P. Kristensen, Ole Karsholt (ZMUC, Copenhagen), Dr. David C. Lees (BMNH, London), Dr. S. Yu. Sinev, (ZIN, St. Petersburg), Dr. Yu. Budashkin (Karadag Nature Reserve, the Crimea), and Simon R. Hill (London) for the loan of material and (or) for providing valuable information.

9 A. Remeikis thanks Dr. Yu. I. Budashkin and Prof. Dr. J. Schuster for generous support during the course of fieldwork in the Karadag Nature Reserve (the Crimea) and Guatemala (Central America).

The author is indebted to the members from Biology Department of Lithuanian University of Educational Sciences particularly to the colleagues from Biosystemtatics Research Group: Doc. Dr. A. Diškus, Prof. Dr. V. Sruoga, dotoral students A. Navickaitė and A. Rocienė.

This study was conducted as part of the “New Faunas” Research Project of the Biosystematics Research Group of Lithuanian University of Educational Sciences with support from the Lithuanian State Science and Studies Foundation (2011, 2012).

10 2. OPOSTEGIDAE AND NEPTICULIDAE (NEPTICULOIDEA) (LITERATURE REVIEW)

2.1. SOME PARTICULARITIES OF NEPTICULOIDS

2.1.1. Family Opostegidae

Opostegidae comprise a morphologically distinct family of small, predominantly white moths whose females possess a primitive, monotrysian reproductive system. Together with their sister family, Nepticulidae, the Opostegidae contain some of the smallest Lepidoptera known, with a wing span ranging from 4 to 16 mm (Davis, 1989; Puplesis, Robinson, 1999; Davis, Stonis, 2007; Remeikis et al., 2009). Many factors combine to entitle the Opostegidae as arguably the most difficult family among all Lepidoptera to study (Davis, 1989). The small size and apparent rarity of most species of Opostegidae, coupled with the great difficulty in locating their usually well concealed, -mining larvae, undoubtedly have hindered previous attempts to collect and study this group (Davis, Stonis, 2007). Although generally global in distribution, evidence now indicates the greatest diversity for Opostegidae occurs in the continental tropical or subtropical regions (Davis, Stonis, 2007; Remeikis et al., 2009, 2010; Stonis et al., 2013). The morphology of opostegids has been extensively discussed by Davis (1989) and, to a lesser extent, by Buszko (1981), Johansson et al. (1990), Puplesis, Robinson (1999) and Davis, Stonis (2007). Head. Oval, slightly or strongly fIattened (depressed) dorso-ventrally. Frons and vertex with tufts of erect piliform scales (collectively termed a tuft, Fig. 1 A). Lower part of face practically naked, with tiny microtrichia. Neck covered by very large and distinctive group of lamellar scales behind eyes forming a collar (the term used below for this structure) (Fig. 1 B). Clypeus very small, triangular or rounded (Davis, 1989; Puplesis, Robinson, 1999; Puplesis, Diškus, 2003).

11 A B Fig. 1. Head structures of Opostegidae: A – lateral view, with lifted eye-caps (courtesy of Puplesis, Robinson, 1999); B – dorsal view (courtesy of Puplesis, Diškus, 2003)

Mandibles absent. Maxillary palpi five-segmented, spined at apex. Haustellum greatly reduced, but sometimes exceeding length of maxillary palpus. Labial palpi three-segmented, much shorter than maxillary palpi. Ocelli and chaetosemata absent. Compound eyes usually large (Fig. 1 A) (but small in Neotropical Notiopostega), occasionally only moderately large. Antennae long, but never as long as forewing. First segment of antenna (scape) greatly enlarged, even more so than in Nepticulidae, the sister group, forming eye-cap which can entirely cover the eye (Fig. 1) (but small in Neotropical Notiopostega) (Davis, 1989; Davis, Stonis, 2007; Remeikis et al., 2009). Antennal pedicel moderately short. Flagellum with 40-90 more or less cylindrical segments; most flagellomeres each with (among other types of sensilla) three palmately branched sensilla ascoidea, each with four to ten 'arms' (Davis, 1989). Thorax and appendages. Tegulae small. Thorax depressed dorso-ventrally. Tegulae and thorax scales usually very similar to ground colour of forewing. Forewing relatively broad, without sexual dimorphism in colour or scaling (in contrast to Nepticulidae, in which many have specialized sex scales) (Puplesis, Robinson, 1999). Forewing of both sexes with only scattered remnants of microtrichia, predominantly in subcostal region and along underside of hind margin of forewing base (Davis, 1989; Puplesis, Robinson, 1999; Puplesis, Diškus, 2003). Microtrichia forming usually well defined, distinctive colour patches (Puplesis, Robinson, 1999), especially in Opostegoides. Most of forewing underside surface covered with brown or cream scales. Upperside of forewing with varying patterns of a few simple types (Puplesis, Diškus, 2003). Forewings of most species white to silvery-white with brown terminaI strigulae and of ten with additional dorsal and/or costal spots; some species with broad, dark (usually brown) fascia on forewing (occasionally even two or three fasciae, including apical area); a few species with intense dark forewing coloration or with other pattern types (Fig. 2).

12

Fig. 2. Resting adult of Opostegidae (courtesy of Davis, Stonis, 2007)

Frenulum absent in both sexes; wing coupling mechanism comprising subcostal pseudofrenular bristIes on hindwing (Davis, 1989). Hindwing narrow, lanceolate, with long cilia; colour usually uniform, except in a few cases where the base is scaled with much paler scales. Wing venation greatly reduced but remaining veins well defined. Forewing characterized by a few straight unbranched veins (Sc, R, M, Cu and A in the most complete configuration, or just Sc, R and M in the most reduced). Hindwing with Sc, Rs, M, Cu and A, with just the first and the last represemed as separate veins. Wing venation is comprehensively figured by Davis (1989: figs 25– 30). Legs with pairs of unequal spurs: one pair on midtibia, two on hind-tibia; fore-tibia without epiphysis and with semi-erect, spinose setae. Abdomen. No specialized abdominal sex scaling is known among the Opostegidae, but males of ten possess a pair of tiny, normally confluent anal tufts. Second sternum (S II) rather variable in shape, and with distinct anterior fragment in a few genera, e.g. Pseudopostega (Davis, 1989). Additional information on abdominal morphology is given by Kristensen, Nielsen (1980), Kyrki (1983) and Davis (1989). However, with the present state of knowledge, the significance of such sporadic data for intrafamilial taxonomy and phylogeny is uncertain. Male genitalia. In contrast to many genera in the sister group Nepticulidae, the opostegid genital capsule generally exhibits the presumed nepticuloid ancestral condition, with the sclerites of segment IX (tegumen and vinculum) forming a complete fused ring (annulus). Tegumen usually well developed (Puplesis, Robinson, 1999; Remeikis et al., 2009), but anterior margin not always well sclerotized. Uncus usually a broad band, in some species with small broadly rounded or very extended lateral lobes with numerous setae, or with just three very long setae on each side;

13 Eosopostega with uncus elaborate and complex. Gnathos very strongly or completely reduced in Opostegoides, but well developed in all other genera, the sclerite usually with a huge central element, but sometimes forming a simple, caudalIy bent plate (Davis, 1989; Puplesis, Robinson, 1999; Puplesis, Diškus, 2003; Davis, Stonis, 2007). Valva in all Opostegidae with a very conspicuous pectinifer on the cucullar lobe with numerous long peg-like spines (usually 12-60). Pectinifer varying slightly or greatly in shape among genera and species, but (in contrast with Nepticulidae) allways on a discrete cucullar lobe which is attached to the main body of the valva via a slender and short to very long pedicellum. In more specialized cases, pedicellum strongly wrinkled transversely and exceeding length of body of valva (Puplesis, Robinson, 1999). Cucullar lobe in most genera very large, but relatively small in Opostegoides and Eosopostega (Davis, 1989; Puplesis, Robinson, 1999). Main body of valva relatively large (Puplesis, Diškus, 2003; Remeikis et al. 2009; Stonis et al., 2013).

2.1.2. Family Nepticulidae Nepticulidae are a family of minute monotrysian Microlepidoptera with a worldwide distribution and nearly 826 described species (Navickaitė et al., 2011; Stonis et al., 2013). Their morphology, biology and taxonomic composition of Nepticulidae have been extensively discussed by Scoble (1983), van Nieukerken (1986); Johansson et al. (1990); Puplesis (1994), Puplesis, Robinson (2000), Puplesis, Diškus (2003), and in our paper (Stonis, Remeikis, 2011) or our chapter of a monograph “Morfologiniai Nepticulidae požymiai, naudojami identifikuojant taksonus ir paremiantys šiuolaikinę sistemą“ (in Lithuanian) (Stonis et al., 2012). The head-capsule of nepticulids is hypognathous, approximately oval, and slightly flattened dorsoventrally. Only a few sulci are present on the head-capsule. The clypeus is a small triangular or rounded lobe; a clypeal sulcus is absent (Puplesis, Robinson, 2000). The mouthparts normally are only developed weakly; mandibles are absent; the maxillae are represented by galeae and maxillary palpi; laciniae are absent; the galeae together unite to form the proboscis (= haustellum) which may sometimes be functional. Occasionally the galeae are very long; the maxillary palpi are always five-segmented, with sensilla; the labial palpi are normally three-segmented and widely separated at their bases, occasionally two segmented; the first (basal) segment of the labial palpus is usually longer than the second and third segments (Scoble, 1983; van Nieukerken, 1986; Johansson et al., 1990; Puplesis, Robinson (2000). In slide preparations of the head-capsule, the sucking pump and membranous hypopharynx are usually visible (Scoble, 1983). Ocelli are absent; chaetosemata are present (Johansson et al., 1990). The compound eye is generally large, its size relative to that of head-capsule varying little. The eye-index is fairly high, in the range 0.9–1.2, and

14 is possibly related to time of flight (diurnal species have a lower index than night-flying ones). The antenna is always shorter, usually much shorter, than the forewing, its basal segment (scape) strongly enlarged and forming an eye-cap with a concave underside. The eye-caps cover the compound eyes when the is at rest. The flagellum comprises 12 to 70 segments (flagellomeres) that are more or less uniform except for the first (the pedicel) which is longer than the rest. Four types of sensilla have been found on the flagellomeres of Nepticulidae: sensilla trichodea, s. chaetica (two types), s. coeloconica and s. vesiculocladum (van Nieukerken, Dop, 1987). The sensillum vesiculocladum described by these authors occurs in all species examined but is unknown outside the family (Puplesis, Robinson, 2000). The tufted head is characteristic. Piliform scales arise from a large circular area on the front of the head, and also from two other patches on the vertex and are collectively named the frontal tuft; its colour has diagnostic value (Puplesis, Diškus, 2003; Stonis et al., 2012). Two groups of piliform or lamellar scales are attached to the back of the head, often overlapping the thorax, and collectively termed the collar. The frons is smoothly scaled below the frontal tuft (Fig. 3).

Fig. 3. Head of Nepticulidae, with lifted eye-caps (courtesy of Puplesis, Diškus, 2003)

The colour pattern of head, thorax and forewing is a very important diagnostic feature (Stonis, Remeikis, 2011); however it is of almost no phylogenetic use, since Neotropical nepticulids (like those from other regions) show great homoplasy (Puplesis, Robinson, 2000). A partial exception is Manoneura, with strikingly strong blue-purple lustre, and possibly a few Neotropical Enteucha with unique white-tipped forewings (Puplesis, Robinson, 2000). The forewing may be uniform or patterned (with one to three fasciae, with spot(s) or with both fasciae and spot(s)) (Puplesis, Diškus, 2003). Most nepticulids have uniformly grey or blackish forewings with a single fascia, the dark areas often with bright metallic lustre which may be bronze, purple, blue or green. But

15 forewing patterns are more diverse in the Neotropical species described in Puplesis, Robinson (2000): nearly 30% have a single white/light fascia on a dark background, 20% have irrorated forewings (irregularly dispersed darker scales on a paler background), 14% have a dark fascia on a paler background, 12% are unicolorous, 6% have three fasciae on a dark background; other species have two fasciae on a dark background, two opposing pale spots, and one spot. Very unusual pattern types among the Neotropical nepticulids include Enteucha cyanochlora with a triangular costal spot together with a white forewing tip, and two species (Stigmella andina and S. pruinosa) with striking sexual dimorphism: females of S. andina have distinctive black-tipped forewings whereas in males the forewing is uniformly pale; males of S. pruinosa have a broad band of whitish androconial scales over brown forewings, absent in females (Puplesis, Robinson, 2000; Stonis, Remeikis, 2011; Stonis et al., 2012). Generally fasciae or spot(s) vary from white to yellowish, with a silver or gold sheen or lustre, occasionally other colours. In females the scales usually appear more coarse, and pale markings (if present) are more extensive than in the male (Puplesis, Diškus, 2003; Stonis et al., 2012). The hindwing varies from creamy white to dark brown, and in males is sometimes covered by black or orange androconial scales (but these are not known in Neotropical species) (Puplesis, Robinson, 2000). A cilia-line, formed by dark-tipped, lamellar scales, may be present. Androconial scales may be present in males, and normally occur on the underside of the forewing or either surface of the hindwings. They may entirely cover the wing, but in other cases are distributed in patches. Sometimes, long specialized androconial scales may extend into the fringe (e.g. Neotropical Acalyptris trifidus, Stigmella pruinosa, partially S. albilamina) (Puplesis, Robinson, 2000). Additionally, van Nieukerken (l986) recorded patches of velvet-textured raised scales on the hindwing underside of European . Wing-coupling in males comprises a double mechanism: firstly, a composite frenulum arising on the hindwing base hooks into the costal retinaculum, which is comprised of hooked scales; secondly, a row of costal bristles (pseudofrenular bristles), arising on the humeral lobe, couples with a subdorsal retinaculum composed of hair-scales. In females there is no frenular bristle, and only the second mechanism is present. For details see van Nieukerken (1986). Some species possess pseudofrenular bristles modified into a hair-pencil (for example, a long blackish cluster in Ectoedemia species 29105 by Puplesis, Robinson (2000). Wing venation is reduced in Nepticulidae, although reduction is generally not as extreme as in Opostegidae (Puplesis, Robinson, 1999). In the largest nepticulid , Stigmella, forewing vein Sc is short and sometimes weakly visible. R+M coalesce, and there are three radial veins (R1,

R2+3, R4+5). Cu is present as a separate vein. Veins lA and 2A are fused along their entire length

16 and are well-developed. In the hindwing, Sc+R1 is short, and Rs and M either usually coalesce in the basal half of the hindwing or are semi-reduced (as in the Neotropical Stigmella albilamina). Cu and A are separate veins. However, in the Neotropical Stigmella ovata, uniquely for Stigmella, R4,

R5, M1 and M2 are preserved in the forewing as four separate veins, and Cu is very long and anastomoses with the medial branch; this striking configuration is probably a plesiomorphy (Puplesis, Robinson, 2000). The most complete venation pattern is characteristic of Etainia (most notably in Holarctic species) (Puplesis, 1994; Puplesis, Diškus, 2003). The forewing in this genus normally exhibits one subcostal, five radial, two medial, one cubital and one anal vein together with a closed cell between M and Cu. The most reduced type of venation within the Nepticulidae is characteristic of Neotropical Enteucha and Manoneura (Puplesis, Robinson, 2000). However, a few southern African Trifurcula and Etainia, and, probably, a few Neotropical Fomoria (Scoble, 1983; Puplesis, Robinson, 2000) show independent reduction of the venation, a feature which limits the value of that character for phylogenetic purposes. The legs are well-developed, frequently concolorous with the underside of the thorax, and varying from silvery-white to blackish. The foreleg is without spurs or epiphysis; the midleg has a single pair of spurs, the two pairs on the hindleg are situated at or near the middle and at the apex of the tibia. The position of the proximal pair of spurs is not constant (Puplesis, 1994; Puplesis, Diškus, 2003). The pregenital segments of the abdomen, including VII in females and VIII in males, are clothed uniformly with lamellar scales. The underside of the abdomen is often paler than the upperside. Tergite VIII of males and VII of females usually bears a pair of anal tufts. Large, paired tufts of extremely long piliform scales arise on tergites IV and VIII in members of the Asiatic Acalyptris repeteki species-group (Puplesis, 1994). They arise from T-shaped sclerotizations and completely cover the caudal part of the abdomen. Smaller but undoubtedly homologous tufts are present in some North American Acalyptris species (Wilkinson, 1979; Puplesis, 1984b), but have not been discovered in Neotropical Acalyptris (Puplesis, Robinson, 2000; Puplesis et al., 2002a). The phylogenetic significance of these tufts (more than half the length of the abdomen) is obscure, but they are a useful diagnostic feature at species level (Puplesis, Robinson, 2000). The colour of the scales on the pregenital and genital segments is useful for separating at least two Neotropical species (Stigmella albilamina and S. fuscilamina) in which this is practically the only external difference (Puplesis, Robinson, 2000). The male genitalia are symmetrical except for the slightly asymmetrical aedeagus and asymmetrically distributed cornuti. The tegumen is fused with the vinculum through the vinculum arms, completing a ring-shaped annulus (Stonis, Remeikis, 2011). The tegumen, a simple band, is

17 very variable in size. Occasionally the tegumen may be invaginated or extended posteriorly. A pseuduncus (posterior extension of the tegumen), if present, is situated dorsal to the uncus or coalesced with it (Etainia) (Puplesis, 1994); usually the pseuduncus is a broad or narrow lobe (Puplesis, Robinson, 2000), but it is sometimes divided into lateral lobes (Puplesis, Robinson, 2000; Stonis, Remeikis, 2011; Stonis et al., 2012). There is no boundary between the pseuduncus and tegumen. Additional rod-like apodemes arising from the tegumen may be present in the male genitalia of Acalyptris (Puplesis, Diškus, 2003; Šimkevičiūtė et al., 2010; Stonis, Remeikis, 2011). In Neotropical representatives these are still not fully separated from the tegumen or are very under- developed (Puplesis, Robinson, 2000), an apparently more plesiomorphic state than in the Palaearctic species of the genus (Puplesis, Robinson, 2000). The uncus is narrowly V-shaped, thickened or hoodlike; it is often bilobed or even with three narrow processes and rarely takes other forms (Puplesis, Robinson, 2000; Puplesis, Diškus, 2003; Stonis et al., 2012). The unusual uncus with a single anterior process in Manoneura has been misinterpreted as 'downfolded' (van Nieukerken, 1986b). Occasionally, an uncus is absent (Ectoedemia) (Johansson et al., 1990; Puplesis, 1994; Stonis et al., 2012). The gnathos is a small sclerite connected through its lateral arms to the tegumen. It consists of a transverse bar (occasionally not developed), posterior processes (or process) and lateral arms. Sometimes anterior processes are additionally developed, and in other cases posterior processes can be replaced by a central element (Puplesis, Diškus, 2003). An unusually broadened posterior process in Acalyptris platygnathos partially replaces the central plate (Puplesis, Robinson, 2000). A very unusual modification of the gnathos is observed in three Neotropical species united into the Stigmella eurydesma species group: the anterior plate-like part of the sclerite is strongly reduced, but the posterior part is developed into very strongly sclerotized and melanized lateral processes (Puplesis, Robinson, 2000). These processes partially coalesce with the more dorsal uncus, and it is sometimes difficult to recognise two separate sclerites (Puplesis, Robinson, 2000). The very unusual gnathos in Manoneura (see Puplesis, Robinson, 2000: figs 84, 86 or Puplesis et al., 2002a) has been misinterpreted as a complex structure comprising gnathos+uncus as a single fused sclerite (Davis, 1978). Together with the tegumen, the gnathos surrounds the tuba analis. Slightly sclerotized and sometimes spined, the dorsal part of the tuba (very rarely visible in Nepticulidae) is referred to as the scaphium, and its ventral region as the subscaphium. The valva is always strongly sclerotized and exhibits numerous diagnostic differences between species (Puplesis, Diškus, 2003; Stonis, Remeikis, 2011; Stonis et al., 2012). The apical region usually tapers to a variable distal process, in some species (predominantly in the Stigmella

18 salicis group) to two processes. In some taxa the apex of the valva remains broad. The inner margin of the valva may bear one or more well-developed lobes or processes, which may be numerous and elaborate in some Neotropical species (e.g., Puplesis, Robinson, 2000). In Fomoria species 29122 (by Puplesis, Robinson, 2000) the valva is actually divided into two lobes. Numerous setae are distributed on the valva, especially on the inner margin and at the apex; the Neotropical Stigmella barbata has very long, strong, hair-like selae from the apical process and in this respect resembles a few Nearctic species. In some cases (notably in the Australian Pectinivalva Scoble and some Asiatic species of Acalyptris) the valva bears a pectinifer, more or less resembling the pectinifer of opostegids. There is still no general agreement as to the homology of these pectinifers. However, the pectinifers of the Asiatic Acalyptris, Pectinivalva sp. and Opostega auritella (Hübner) have been well illustrated and compared by van Nieukerken (l986b: figs 60–64), and its believed they are homologous (Puplesis, Robinson, 2000). Only a single species in the Neotropical fauna (Acalyptris bovicorneus) has been discovered with a similar pectinifer (Puplesis, Robinson, 2000); other related Acalyptris species have none. After Puplesis, Robinson (2000) the presence of a pectinifer may be either a parallelism, or more likely a plesiomorphy preserved in Pectinivalva and some Acalyptris; the latter explanation gains some support from the widespread but sporadic occurrence of this structure elsewhere in the monotrysian , notably in Incurvarioidea. The pectinifer in Opostegidae, sister group to Nepticulidae, is unambiguously derived in comparison with the hypothetical ancestral state which appears to be expressed in those Nepticulidae with a pectinifer. In Opostegidae the pectinifer is borne on a cucullar lobe well separated from the main body of the valva, a striking autapomorphy of the family expressed in all the taxa known (Puplesis, Robinson, 2000). The transtilla forms a dorsal connection between the apodemes of the valvae (Stonis et al., 2012). The fully developed transtilla (with a transverse bar) is reduced or absent only in Pectinivalva, Enteucha, Glaucolepis and most world species of Acalyptris (but in less than a third of the Neotropical species). The Neotropical Ectoedemia fuscivittata also exhibits secondary loss (reduction) of the transverse bar of the transtilla, not previously recorded for the genus (Puplesis et al., 2002a). In other nepticulids the transtilla is well-developed, consisting of an uninterrupted or medially interrupted transverse bar (Stonis, Remeikis, 2011; Stonis et al., 2012). The term 'sublateral process(es) of transtilla' is widely used in the literature on Nepticulidae and for the most part refers to the valval apodeme (Johansson et al., 1990). Admittedly, it is difficult in some cases to determine where the transtilla bar ends and the apodeme begins, but the term is inappropriate; we use it here with considerable misgivings (Puplesis, Robinson, 2000). 'Sublateral processes' are invariably developed, sometimes strongly so but the fusion of the

19 apodeme and transtilla such that the former is indistinguishable and the whole is a continuous rounded band of sclerotization that could be interpreted entirely as transtilla; where the continuous structure is angular it could be inferred that the transverse element is the transtilla and the longitudinal elements are the apodemes, but there is no justification for this assumption. Where the apodeme runs anteriorly beyond the transtilla bar, Puplesis, Robinson (2000) have followed accepted practice and designated this as a 'sublateral process', i.e., 'sublateral process' is a synonym of 'apodeme'; where it does not, Puplesis, Robinson (2000) have recorded the sublateral process as absent, but the apodeme may well still be present. There is some intraspecific variability in the transtilla and apodemes; where slender sublateral processes are present, their curvature may vary; broad processes may vary in shape and quite often the left process may not be symmetrical with the right. Although the presence or absence of the transverse bar of the transtilla is a good diagnostic feature of a few genera, and may have some phylogenetic significance, the characteristics of size and shape (discounting intraspecific variability) are significant only for species diagnosis. We treat a well-developed transtilla with a broad and enlarged transverse bar or specialized sublateral processes as a derived feature (Stonis, Remeikis, 2011). The juxta is a small triangular, trapezoidal or irregularly shaped sclerotization of the membrane between the valvae ventral to the aedeagus (Puplesis, 1994; Stonis et al., 2012). It is variable in shape and in its degree of sclerotization and occurs irregularly in different genera of Nepticulidae with little phylogenetic predictability. However, the juxta in Stigmella is simpler and more plate-like, than in, for example, Acalyptris, where it tends to be more complicated or very complicated and probably contains elements derived from the posterior extension of the vinculum (Puplesis, Robinson, 2000; Stonis, Remeikis, 2011). In some Stigmella species the juxta is apparently not connected to the vinculum; sometimes the juxtal region is entirely membranous (i.e., a juxta is absent) (Puplesis, Robinson, 2000; Stonis et al., 2012). The vinculum is shaped like a large plate and is sometimes narrowed or bilobed anteriorly. Two types of vinculum are present in Nepticulidae. In the first, the vinculum is connected through wide and well-developed lateral arms with a large tegumen and these together form a ring surrounding the aedeagus ('ringshaped' vinculum). In the second type the vinculum has moderate lateral arms which connect with a strongly reduced and usually small tegumen ('U-shaped' vinculum). In the present treatment, the complex of vinculum plus tegumen is regarded as an annulus (Puplesis, 1994). The size of the ventral plate of the vinculum, lateral lobes and anterior invagination may vary strongly between species. The anterior region of the vinculum has been referred to as the saccus (Wilkinson, Scoble, 1979; Newton, Wilkinson, 1982).

20 The aedeagus, in contrast to many other Lepidoptera, is broad and strong1y sclerotized, cylindrical or retort-shaped, frequently broadened basally and occasionally apically; it is often 2–3 times longer than the valva. Carinae may be developed towards the apex of the aedeagus and vary considerably between species. The vesical cornuti vary from very small and spine-like or almost granular to large, horn-like, lamellar, or elongate and irregular. The number of cornuti varies interspecifically from a few to several hundred. They have great diagnostic value, in most cases at species level, very occasionally at higher taxonomic levels (e.g., in Trifurcula and Glaucolepis). The striate thickening surrounding the base of the ductus ejaculatorius (the arcuate plate through which the ductus passes into the vesica) was regarded as a pair of striate plates and was named the cathrema by Kemperman, Wilkinson (1985). In the Australian Pectinivalva, the cathrema is only weakly developed and this state is plesiomorphic. In some Stigmella species a distinct tubular membrane (the manica) surrounds the aedeagus. This membrane is usually spinose but such a modification has yet to be discovered among the Neotropical species (Puplesis, Robinson, 2000). Females are of the monotrysian type, with a common terminal anogenital opening. After Johansson et al. (1990) and Puplesis, Robinson (2000) the vagina is short, widening into a vestibulum which may sometimes be strongly folded, and almost always protrudes laterally to form an accessory sac in Stigmella. In Ectoedemia a distinctive spiculate pouch is developed. In Etainia, Acalyptris and Fomoria additonal sclerotization of the vagina-vestibulum forms structures that can be described as an antrum or sclerites of the vestibulum. The ductus spermathecae opens into the accessory sac or into the vestibulum if an accessory sac is absent. Usually the ductus is coiled, and occasionally it is spiculate. The bursa copulatrix is large, usually oval, with or without pectinations or signa. The signa are reticulate (in Trifurculinae sensu Puplesis, 1994; Stonis et al., 2012) or in a different pattern; sometimes they are large spines (in some species of the Palaearctic Stigmella paliurella species-group). In some species of the Palaearctic species-group, the accessory sac has functionally replaced the bursa copulatrix, which is minute or indiscernible (Johansson et al., 1990; Stonis et al., 2012). The apophyses are well-developed and vary interspecifically. The anal papillae usually form a broad, flat posterior margin to the abdomen. The ovipositor is weakly developed, occasionally protruding (Scoble, 1983; van Nieukerken, 1986a; Johansson et al., 1990; Stonis et al., 2012). Biology. The biologies of Neotropical nepticulids are known only from very scanty reared material (Puplesis, Robinson, 2000). Generally, nepticulid eggs are laid singly, glued to the surface of may be a leaf or another (often damaged) plant organ. The precise oviposition site is sometimes characteristic, as certain species lay their eggs either on the upper- or underside of a leaf. But other species may lay on either

21 side of the leaf. Usually just one egg per leaf is laid, but in some species (when population density is high) a few or numerous mines may be found in one leaf. For example, in the Western Palaearctic Stigmella paliurella (on Paliurus spina-christi) and Stigmella ficulnea (on Ficus carica), 10 to 30 mines have been noted in one leaf (in the Caucasus and Turkmenistan) (Puplesis, 1994; Puplesis, Diškus, 2003). Between 400 and 500 mines of the Eastern Palaearctic Ectoedemia picturata have been noted on one (composite) leaf of Rosa rugosa in Far Eastern Russia (Puplesis, 1985a). The duration of the egg stage varies greatly, from 8–11 days to 130–150 days. Although this has some correlation with temperature (Kino, 1981), it relates also to the voltinism of the species. Four or five larval instars are typical, but occasionally – as in the subgenus Ectoedemia (Zimmermannia) – 6–8 instars are known (Schönherr, 1958; van Nieukerken, 1985b; Johansson et al., 1990). After hatching, the young larvae bore into the leaf. The majority make upper-surface mines, but occasionally mines are under-surface ones (e.g., in the Asiatic Stigmella flavescens) (Puplesis, 1994). The majority of Nepticulidae are leaf-miners, but sometimes larvae mine buds and shoots (Etainia, and some Bohemannia species); the summer generation of Etainia species makes short mines in the keys () of Acer; Ectoedemia (Zimmermannia) mine in young bark (cambium). The Neotropical Ectoedemia species 29105 (Puplesis, Robinson, 2000) may be a bark-miner, because similar species of this group in the Holarctic region feed only in bark. Amongst the numerous leaf-miners, some species (Ectoedemia populella-group) start to mine in the petiole or midrib of a leaf and only later enter the tissues of the leafblade (Johansson et al., 1990). Old mines are sometimes completely transparent, or at least translucent and easily visible (e.g., the Neotropical Stigmella gossypii, Enteucha gilvafascia) (Puplesis, Robinson, 2000). In some species, various instars make different types of tunnels, each type with characteristic frass deposition (the Neotropical Acalyptris species 29140 on Lonchocarpus lineatus (Leguminosae) has uniform frass deposition throughout the whole mine (Puplesis, Robinson, 2000). The tunnels made by Nepticulidae have characteristic patterns (Fig. 4). Mines can be generally categorized as: linear (most species) blotch (very rare); a combination of gallery in the first half and blotch in the second half (Fig. 4 A, rare); spiral (= helical) (only in Enteucha). Bark mines are always linear (Johansson et al., 1990). The mines of the majority of species are characteristic and have diagnostic value at species level (Puplesis, Diškus, 2003). However, the shape of the mine and the colour and distribution of the frass may vary, depending (amongst many factors) on the thickness of the leaf, light conditions and hostplant species (Puplesis, Diškus, 2003; Stonis, Remeikis, 2011; Diškus, Stonis, 2012).

22

A B Fig. 4. Leaf-mines of Nepticulidae: A – , larva absent; B – S. hybnerella, with a feeding larva (orig.)

The larva never its mine until it is full-grown. However, in most investigated Glaucolepis, and some other species, the larvae use more than one leaf (Johansson et al., 1990). These species mine continuously from one leaf to another via petiole or stem. The duration of the larval stage is usually short, in the majority of cases a matter of a few days (in temperate regions, as well as in Belize) (Puplesis, Robinson, 2000), but sometimes this stage may take a few months. A prolonged larval stage is characteristic of most Ectoedemia, which mine from late summer to autumn, and for species overwintering as larvae (e.g., the Palaearctic Fomoria weaveri and Stigmella castanopsiella). The longest larval stages are known for species of Ectoedemia (Zimmermannia) (van Nieukerken, 1985b). The season of the larval stage varies, but larvae of most species mine in autumn. Larvae of certain Ectoedemia species mine in yellowed or even fallen leaves, with the conservation of characteristic 'green islands' (Ivinskis et al., 1985; Johansson et al., 1990). Except in a few species (the Palaearctic Fomoria weaveri group, Ectoedemia agrimoniae and Trifurcula eurema), the fully-grown larva emerges from the mine and descends on a silk thread, usually to the ground. Larvae may then hide in detritus where they spin a silk cocoon. In some species the cocoon is spun on a stem or trunk of the hostplant. The duration of the pupal stage is short in the summer generation(s) of bivoltine or multivoltine species, usually 10–20 days. In the overwintering generation, fully-grown larvae do not pupate but diapause through the long, cold period in a prepupal stage. After diapause, the pupal stage is as short as that of summer generations. The pupa is protruded halfway out from the cocoon by means of its abdominal spines, before the moth emerges (Johansson et al., 1990). The imago has a short lifespan. Indoors, when natural conditions are mimicked, the lifespan of the moths amounts to 2–3 days, or 5–6 days when fed with sugar syrup (Puplesis,

23 1994). The lifespan of adults collected in the wild and not fed is usually only one day, rarely three days. According to Johansson et al. (1990), individuals of several species aestivate in southern Europe. This may also be a characteristic of some central Asiatic species. Otherwise, only when the imago is kept at a low temperature (about 10°C), may the lifespan (with the moth generally in torpid condition) be lengthened to up to 5–6 days. The duration of the flight period of a species is about 20–30 days, sometimes longer. Emergence starts at different times in the case of different species. Usually, the first males emerge earlier than the first females. The moths usually fly around their hostplants in the afternoon and evening before dusk. Adults of most species are attracted to light, males being collected rather more frequently than females. Before copulation the male exhibits specific behaviour in the form of a 'dance' (Puplesis, 1994). The male runs close to or around the resting female, vibrates his raised wings and turns his abdomen slightly upward. If the female responds by raising her wings slightly and vibrating them a little, the male instantly turns his abdomen towards her, and mating takes place. The wings of both individuals are then lowered. Duration of copulation is at least 45 minutes in the eastern Palaearctic Stigmella kozlovi (Puplesis, 1984a) and about 20 minutes in the western Palaearctic Ectoedemia liebwerdella (Schönherr, 1958). Univoltine species are rare among the Nepticulidae. However, within the subfamily Trifurculinae, many species of Ectoedemia and Trifurcula and some Fomoria and Bohemannia have only one generation per year. In most univoltine species the larval stage is unusually long, especially in Ectoedemia (Zimmermannia). In this group this stage lasts from two to six or seven months, or even nearly two years. Bivoltine species are in the majority in the Holarctic region. Details of the biology of tropical species are largely unknown. A few species have been noted as multivoltine, but we would expect a great number of such species in the tropics. Even among the boreal fauna, the majority of the so-called bivoltine species might have three (or more) generations per year if climatic conditions permit (Puplesis, Diškus, 2003; Diškus, Stonis, 2012). Nepticulidae are trophically associated with dicotyledonous from the Magnoliophyta. According to van Nieukerken (1986a, 1986b), plants from 18 subclasses and numerous families are fed upon by nepticulid larvae. Exhaustive data on the hostplant relationships of western Palaearctic Nepticulidae have been presented by van Nieukerken (1986a) and by Johansson et al. (1990). The majority of nepticulid species in the Holarctic mine plants from a few families: Rosaceae, Betulaceae, Salicaceae and Leguminosae. Further south, more families are involved (Puplesis, 1988a,b, 1992; Johansson et al., 1990). But for many species the hostplant is still unknown (Puplesis et al., 1996). Two new hostplant families (Cunoniaceae and Eucryphiaceae) were recently recorded by Hoare (2000) for the Australian fauna; they were

24 considered as a part of the ancient angiosperm flora that covered large areas of Australia, Antarctica and South America in the late Cretaceous and early Tertiary (see Hoare, 2000). After Puplesis, Robinson (2000) currently only five nepticulid hostplant families have been recorded in the Neotropical Region: Leguminosae (Lonchocarpus lineatus), Compositae (Senecio bonariensis), Onagraceae (Ludwigia major), Malvaceae (Gossypium barbadense, occasionally G. hirsutum) and Polygonaceae (Coccoloba uvifera, the hostplant of two species). Monophagy is known to occur within the family, but by far the greatest proportion of nepticulid species are oligophagous (van Nieukerken, 1986a; Puplesis, 1994). Broad oligophagy is especially characteristic of species feeding on Rosaceae. Strict oligophagy is the most common trophic type within the family and applies to nearly 80% of all investigated Nepticulidae. A few cases of disjunct oligophagy are known in the Holarctic fauna but no actual polyphagy. There are numerous species among Nepticulidae which can be very abundant and damage the hostplant (the Neotropical Stigmella gossypii on Gossypium may fall in this category). However most species are no more than potential pests (Stonis, Remeikis, 2011), and significant host damage appears to be an exception brought about by particularly favourable ecological conditions. Nepticulid populations appear to be able to react with very rapid increases in density (Puplesis, Diškus, 2003; Diškus, Stonis, 2012).

2.2. STUDY HISTORY

2.2.1. Studies of Opostegidae of Central and South America

Within the last 20 years, notable efforts have appeared to raise this family from obscurity. The first of these was a generic review of the family and world catalogue (Davis, 1989), followed by a revision of the Oriental Opostegidae (Puplesis, Robinson, 1999), and most recently by a review and world catalogue of the Nepticuloidea and Tischerioidea by Puplesis, Diškus (2003). However, the biggest boost was given to number of species known from Central America by Davis, Stonis (2007) who described one new genus and 68 new species. Most recently a few papers dealing with diversity of the Opostegidae in Central America were published by us (Remeikis et al., 2009; Remeikis et al., 2010). One new species was described by us from the Himalaya (Stonis et al., 2013) Contribution of different authors to the species number in Opostegidae (and other related Lepidoptera) was recently reviewed by Navickaitė et al. (2011).

25 2.2.2. Studies of Nepticulidae of Europe and other regions of the world

These tiny moths are still poorly studied in many regions. Only the northern European nepticulid fauna can be considered to have been exhaustively studied (Johansson et al., 1990). Studies in other regions of the world fall some way short of reflecting the actual diversity of the group although coverage compares well with that of other groups of Microlepidoptera. The most exhaustive revisions or detailed taxonomic papers have been published for the western Palaearctic (Johansson, 1971; Nieukerken, 1983, 1985a, 1985b, 1986a; Johansson et al., 1990; Nieukerken, Puplesis, 1991; Laštuvka, Laštuvka, 1997, etc.), Central and Eastern Palaearctic, including Far Eastern Russia and Japan (Kemperman, Wilkinson, 1985; Puplesis, 1994; Puplesis, Diškus, 1995, 1996a, 1996b, 1996c; Puplesis et al, 1996, 1997), South Africa (Scoble, 1978a, 1978b, 1980a, 1980b, and, outstandingly significant, 1983), Nearctic (Davis, 1978; Wilkinson, 1979, 1981; Wilkinson, Scoble, 1979; Wilkinson, Newton, 1981; Newton, Wilkinson, 1982), New Zealand (Donner, Wilkinson, 1989), and recently for Australia (Hoare et al., 1997; Hoare, 2000). Long neglected, the diverse Australian fauna is being further studied by Robert Hoare; the Chinese and Japanese fauna is being revised by Erik van Nieukerken (NNM); a revision of the Central Asiatic nepticulids is currently in preparation by J. R. Stonis and A. Diškus (with co-authors, VPU). For genus Acalyptris, particularly valuable and exhaustive is a taxonomic revision of the Acalyptris platani and A. staticis groups in Europe and the Mediterranean by van Nieukerken (2007). Currently the checklist of the European fauna of Acalyptris includes nine species divided into two species groups (van Nieukerken, 2007) and listed below. The platani group: 1) Acalyptris minimella Rebel, 1926, known from Portugal, Spain (incl. Ibiza, Mallorca), France (incl. Corsica), Italy (incl. Sicily, Sardinia), Croatia, Slovenia, Greece and North Africa (Morocco, Algeria, Tunisia); larvae on Pistacia lentiscus, P. terebinthus; 2) A. platani Müller-Rutz, 1934, known Portugal, Spain, France (incl. Corsica), Italy (incl. Sicilia), Switzerland, Croatia, Greece, Cyprus, , and Iran (in the current paper for the first time also recorded in Ukraine (the Crimea); larvae on Platanus spp.; 3) A. loranthella Klimesch, 1937, known from Slovakia, Czech Republic, Austria, Hungary, Italy (incl. Sicily), Romania and Greece; larvae on Loranthus europaeus; 4) A. pistaciae van Nieukerken, 2007, known from Greece (Crete, Rodhos), Cyprus and Turkey; larvae on Pistacia terebinthus and P. lentiscus (van Nieukerken, 2007). The staticis group: 1) A. staticis Walsingham, 1907, known from Spain (incl. Canary Islands: Tenerife); larvae on Limonium pectinatum; 2) A. pyrenaica Laštuvka, Laštuvka, 1993, known from Spain, Germany (host-plant unknown) (Laštuvka, Laštuvka 1993); 3) A. maritima

26 Laštuvka, Laštuvka, 1997, known from Italy, Greece and Croatia; larvae on Limonium vulgare (Laštuvka, Laštuvka 1997); 4) A. limonii Laštuvka, Laštuvka, 1998, known from Greece and Croatia; larvae on Limonium vulgare (Laštuvka, Laštuvka 1998); 5) A. lesbia known from Greece; larvae on Limonium gmelinii (van Nieukerken, 2007). Most recently two our papers were published dealing with the Crimean Nepticulidae (South East Europe) (Stonis, Remeikis, 2011; Stonis et al., 2013)

2.2.3. Studies of Nepticulidae of Central and South America

The Neotropical Region (Central and South America), while vast and with a hugely diverse biota, is comparatively unexplored with respect to collection and study of Nepticulidae (Stonis et al., 2013). The first two species recorded from the region, Stigmella johannis and Fomoria molybditis, were described by Zeller in XIX century (as Nepticula) from Colombian material. Much later, five further species were reported from Peru and two from Guyana by Meyrick (1915); however, one of the Guyanese species (Enteucha cyanochlora) was placed in the and only recognized as nepticulid some 70 years later (Davis, 1984, 1985). During the last seven to eight decades there have been practically no investigations of Neotropical Nepticulidae, except for isolated descriptions of a few new species from Puerto Rico (Forbes, Leonard, 1930) and Argentina (Meyrick, 1931) and a few relatively recent discoveries in Florida (Davis, 1978; Wilkinson, 1981) of species with Neotropical affinities. Stigmella plumosetaeella was described by Newton, Wilkinson (1982) from Arizona, U.S.A., and here it is additionally recorded from south-western Mexico. However, until 2000 only 21 species have been recorded from Central and South America and the tropical states of the USA. Thirteen of these have been listed by Davis (1984) in the Neotropical checklist, but most have never been illustrated. The biggest boost was given to number of species known from the Neotropics by Puplesis, Robinson (2000) who presented in total 58 species of Nepticulidae from Central and South America. The authors also included 5 species currently known from southern Florida as potential members of the Neotropical fauna; the practicality of this was confirmed by recent discovery of 'Florida species' in Belize and Dominica (e.g., Manoneura basidactyla). Another 8 species with a distribution range from northern Florida into the northern states of the USA have not been incorporated into the revision by Puplesis, Robinson (2000) as their hostplant data and distribution suggests they are typical members of a temperate fauna; they were listed at the end of the checklist. The species revised by Puplesis, Robinson (2000) fall into seven genera: Enteucha Meyrick (6 species), Manoneura Davis (2 species), Stigmella Schrank (23 species), Ectoedemia

27 Busck (5 species), Fomoria Beirne (4 species), Acalyptris Meyrick (16 species) and Glaucolepis Braun (2 species). Only Manoneura is known exclusively from the Neotropics. The revision by Puplesis, Robinson (2000) was the first attempt to demonstrate to the scientific community an example of the diversity of Neotropical nepticulids and to prepare the ground for future investigations of this fauna. The authors studied a substantial unidentified material collected during the last thirty years and deposited mainly in the University of Copenhagen and in the United States National Museum of Natural History (USNM, Smithsonian Institution), together with material that R. Puplesis and S. R. Hill have collected in Belize. The Belize collecting trip, undertaken in April 1998 by R. Puplesis with the assistance of Simon Hill (University of Westminster, UK), has resulted in the description of a very interesting and rich rainforest fauna of nepticulids and other primitive Microlepidoptera. The nepticulid species found in Belize comprised exactly half of all species treated in the revision by Puplesis, Robinson (2000). Belize has given for scientists a baseline from which to estimate the possible extent of Neotropical nepticuloid diversity, allowed the authors to compare the overlap of the Belize collection with previous samples, and to supplement the existing BMNH holdings (otherwise almost entirely aged type material). Nearly one half of the country visited by R. Puplesis and S. R. Hill lies in dense protected forests, some still unexplored. Twenty-eight of species collected in Belize were new taxa (including four species left unnamed). However, none was known from the "heart" of the Neotropics – the Amazon basin – and it was assumed by Puplesis, Robinson (2000) that the absence of specimens from Amazonian rainforest reflected collecting effort, coupled with a lack of diversity. Later, the fieldwork in Ecuador of 1999 and 2001 resulted in the collecting new material. Therefore, in a following paper by Puplesis, Diškus, Robinson (2002a) 16 new species from the upper Amazon basin and the Andes (Ecuador) were recorded, increasing the number of species known from the Neotropics by more than one-fifth. The specified fieldwork in Ecuador also resulted in the acquisition of additional material of Manoneura basidactyla (Davis) and Ectoedemia fuscivittata Puplesis & Robinson (Puplesis et al., 2002b). These were the first records of these taxa from equatorial America. In this paper (Puplesis et al., 2002b) the authors amplified the descriptions and provided further illustrations of these species and updated the checklist of Neotropical Nepticulidae with a distribution chart and map (Puplesis et al., 2002b). Also the authors defined four species-groups and reviewed the known biologies of Neotropical Nepticulidae, adding observations on nepticulid leaf- mines in Ecuador, and review the diversity of Nepticulidae in tropical America (Puplesis et al., 2002b). This publication was followed by a summary of the Neotropical Nepticulidae by Puplesis and Diškus in a world review of the Nepticuloidea and Tischerioidea (Puplesis, Diškus, 2003).

28 Most recently, a paper dealing with two new species and a checklist of Central American Nepticulidae was published by us (Stonis et al., 2013). It describes Acalyptis maya Remeikis & Stonis and A. jucatani Remeikis & Stonis for the first time. Also 11 new species are pending for desription (Stonis, Remeikis, in prep.); these species were collected in Argentina, Colombia and Guatemala.

29 3. GOAL AND OBJECTIVES

The main goal of our study was a taxonomic re-assessment of Opostegidae and Nepticulidae (Nepticuloidea) on a basis of scientific examination of new material collected in the Neotropics, Crimea and Himalaya.

The following objectives were set for achieving the main goal:

Research Strategy 1 (Neotropical fauna) ■ An examination of collection sample of Opostegidae recently collected in Mexico, Costa Rica and Ecuador (currently deposited in LEU) and species identification and description of new taxa. ■ A re-assessment Central American fauna and a contribution to the most recent taxonomic revision of the Neotropical fauna of Opostegidae. ■ A new species designation, with a short re-assessment of the Neotropical fauna of the Nepticulidae on a basis of study of substantial unidentified material collected in Argentina and available to us from ZMUC and USNM.

Research Strategy 2 (South European fauna) ■ A review of nepticulid species diversity and distribution in the Crimea on a basis of field collecting (mainly in Karadag Nature Reserve) and investigaton of an unidentified material collected by other researchers.

Research Strategy 3 (Himalayan fauna) ■ An examination and taxonomic review of Opostegidae species occurring in the Himalaya, with description of new species. ■ An elucidation of the Himalayan Nepticulidae fauna on a basis of a fieldwork in the Indian Himalaya.

30 4. MATERIAL AND METHODS

4.1. SCIENTIFIC MATERIAL EXAMINED The studied material was collected during fieldworks in South Eastern Europe (the Crimea, 2009; see descriptions by Stonis, Remeikis, 2009a–e, 2010a–h), the Himalaya (2010; see descriptions by Stonis, Remeikis, 2010a–h, 2011b–e), Central America (Mexico: Yucatan, 2011 and Guatemala, 2012; see descriptions by Stonis, Remeikis, 2012; Stonis et al., 2012a,b,c, d), and South America (Colombia, 2013). Primary collecting-sites of author’s fieldwork in the Crimea: Karadag Nature Reserve (20 km W of Feodosia) 44°54'N, 35°13'E (Fig. 6), Sudak, 44°50'N, 34°57'E. In all these sites material was collected by rearing adults from mining larvae and collecting at light using standard method for light-collecting.

A

B C Fig. 6. Fieldwork in Karadag Reserve (2009): habitats: A–B – mountain slopes covered with Rosa sp. and Cotynus coggygria; C – Pistacia lentiscus on southern slopes of Karadag

31 Primary collecting-sites in the Himalaya: Rishikesh 30°07'N, 78°19'E (Fig. 7B,C), Mussoorie 30°27'N, 78°04'E. (Fig. 7A), Dhanolti 30°25'N, 78°15'E; Kadukhal 30°24'N, 78°17'E; Chamba 30°21'N, 78°23'E (see published descriptions by Stonis, Remeikis, 2010a–h, 2011b–e). In all these sites material was collected by rearing adults from mining larvae, but in Rishikesh also at light using standard method for light-collecting.

A

B C Fig. 7. Fieldwork in Guatemala (Central America) and the Himalaya: A – leaf-mine collecting on mountain slopes in SW Guatemala; B–C – leaf-mine collecting in Rishikesh, the Himalaya

Other substantial unidentified or identified material (incl. type material) was available to the author of the thesis from world‘s leading biodiversity research institutions: BMNH – The Natural History Museum, London, Great Britain; USNM – National Museum of Natural History, Smithsonian Institution, Washington D.C., U.S.A.; LEU – Lithuanian University of Educational Sciences (formerly Vilnius Pedagogical University), Vilnius, Lithuania; ZIN – Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia; ZMUC – Zoological Museum, University of Copenhagen, Denmark.

32 Description of the published new species (Remeikis et al., 2009; Stonis et al., 2013) were possible thanks to material collected by Simon R. Hill (London, U.K.) and Jonas R. Stonis (Vilnius, Lithuania) during fieldwork on the Pacific coast of Central America, in 2008, utilizing two primary collecting-sites: 1) Manuel Antonio (S of Quepos, Pacific coast, Costa Rica, tropical forest at 20 m, 84°10'W, 9°21'N) (Fig. 8 A); 2) Puerto Angel (Oaxaca Region, Pacific coast, Mexico, secondary forest at 45–65 m, 96°29'W, 15°39'N) (Fig. 9 B). Additional material was collected by Virginijus Sruoga, Jonas R. Stonis (Vilnius, Lithuania) and Simon R. Hill (London, U.K.) in Ecuador, in 2007. Primary sites in Ecuador were:

A B Fig 8. Primary collecting-sites: A – Manuel Antonio (S of Quepos, Pacific coast, Costa Rica, tropical forest at 20 m, 84°10'W, 9°21'N); B – Misahualli (Napo Region, SE of Tena, Ecuador, premontane tropical forest at 400–500 m, 77°36'W, 01°04'S)

A B Fig. 9. Primary collecting-sites: A – Bucay (=Cumanda, 80 km E Guayaquil, Ecuador, premontane tropical forest, 650–700 m, 79°5'W, 2°10'S); B – Puerto Angel (Oaxaca Region, Pacific coast, Mexico, secondary forest at 45–65 m, 96°29'W, 15°39'N)

33 3) Misahualli (Napo Region, SE of Tena, premontane tropical forest at 400–500 m, 77°36'W, 01°04'S) (Fig. 9 B); 4) Bucay (=Cumanda, 80 km E Guayaquil, premontane tropical forest, 650–700 m, 79°5'W, 2°10'S) (Fig. 9 A). In all these sites (in Central America and Ecuador) material was collected at light using standard method for light-collecting. The type series of the new species and other newly collected material currently are deposited in the collection of Biosystematics Research Group (LEU), with further re-deposition at ZMUC.

34 4.2. METHODS

Rearing adults from mining larvae and light-collecting Larvae were collected and reared, and adults were collected at light. Mined leaves (or other plant parts) were placed in Petri dishes which were then checked regularly for emerged adults. The rearing of adults is conducted under both natural and laboratory (indoors) conditions. For this purpose, leaves (or other organs of a plant) are carefully checked (Fig. 10), and infested parts with a larva (Fig. 11) are taken and placed in a Petri dish or special container. Detailed description of the method is given in Diškus, Stonis, 2012.

A B

C D Fig. 10. Leaf-mine collecting: A – checking leaves of Platanus for Acalyptris platani larvae in Karadag Nature Reserve (2009); B – checking leaves of Cotinus for Simplimorpha promissa larvae in Karadag Nature Reserve; C – leaf-mine collecting in the Himalaya (2010); D – a leaf-mine of nepticulid on Zizyphus

35

A B

C D Fig. 11. Leaf-mines and a cocoon of Nepticulidae: A, B – leaf-mines with feeding larvae; C – larva leaving the leaf-mine; a cocoon on host-plant

Adult moths were collected by attracting them to mercury-vapour light from a lamp suspended slightly above eye-level and 5–10 cm in front of a white screen. As many different habitats and sites as possible were sampled. Moths attracted to the screen (Fig. 12) were collected into small glass tubes and pinned after killing with ethyl acetate or by heath.

36

A B Fig. 12. Collecting at light: A – a nepticulid moth attracted by light; B – new “invention” widely used during fieldwork in Karadag Nature Reserve – a screen with a bent top

Dissection and genitalia slides Genitalia were prepared following the method described by Robinson (1976) and shortly summarized in Diškus and Stonis (2012) or in our publication in co-authorship with Stonis (Stonis et al., 2013). After maceration of the abdomen in 10% KOH and subsequent cleaning, male genital capsules were removed from the abdomen and mounted ventral side uppermost. Where the genital armature was particularly complicated, the genitalia were studied and sketched in glycerin before permanent mounting. The aedeagus was removed and mounted alongside the genital armature except in the case of some paratypes or where its removal would jeopardise study of fine structure and where it did not obscure other sclerites. Female genitalia were removed entirely from the abdomen, cleaned and mounted ventral side uppermost. Genitalia and abdominal pelts of both sexes were stained with Chlorazol Black (Direct Black 38/Azo Black) or, occasionally, mercurochrome, and mounted in Euparal (Remeikis et al., 2009; Stonis, Remeikis, 2011; Diškus, Stonis, 2012).

37

Fig. 13. Slide preparation using a Biolam AU-12

Adults and morphological structures of the genitalia were studied and photographed using a Leica DFC420 camera and a Leica DM2500 microscope from specimens in glycerol before permanent mounting in Euparal. Other details on methods and terminology are given in Puplesis, Diškus (2003), Davis, Stonis (2007) or in Stonis et al., 2013).

Fig. 14. Slide photography using a Leica DM2500 attached to a microscope

38 5. RESULTS

5.1. NEW DATA TO THE NEOTROPICAL FAUNA OF NEPTICULOIDEA

5.1.1. Contribution to the fauna of Opostegidae of the Neotropical Region

5.1.1.1. New species of Opostegidae from Costa Rica and Mexico

In this study, we report upon the results of recent field-collecting along the Pacific Coast of Costa Rica and Mexico (in 2008), and also in Ecuador (in 2007), i.e. collections unrepresented among material recently reviewed by Davis, Stonis, 2007. From these new collections, we describe below 3 new species (Pseudopostega robusta from Costa Rica and P. mexicana and P. latiplana from Mexico) (Remeikis et al., 2009, 2010).

5.1.1.1.1. Pseudopostega robusta Remeikis & Stonis (Fig. 15)

MALE. Forewing 2.6–2.8 mm; wingspan 5.9–6.2 mm. Head: vestiture entirely white; scape white; flagellum pale beige; palpi cream. Thorax and tegulae white. Forewing white, marked with a small, pale brown, dorsal spot just below the middle of dorsal margin, two dark brown, subapical costal strigulae (possibly other strigulae present, but appear rubbed in type series), and a fuscous apical spot; cilia mostly light beige or pale brown. Hindwing and cilia pale brown or beige. Legs cream. FEMALE. Unknown. MALE GENITALIA. Uncus a pair of small, narrow and truncate lobes, widely separated at a distance of length of cucullar lobe. Vinculum broad; anterior margin slightly concave. Gnathos well-sclerotized, basally divided into two lateral parts (each bearing a lateral fold), with stout caudal process; basal fold indistinct or absent. Valva with large cucullar lobe, bearing a pectinifer consisting of a single row of 18–20 blunt spines; distal apex of cucullar lobe slightly extended, slender; pedicel broad; valva 0.6 the length of genital capsule. Juxta long rod-like median extention from vinculum. BIOLOGY. Adults fly in March. Otherwise unknown. DIAGNOSIS. Pseudopostega robusta belongs to the lateriplicata group and most resembles P. lateriplicata Davis & Stonis and P. floridensis Davis & Stonis. The male of P. robusta can be

39 distinguished by the divided basal part of the gnathos with the thickened lateral folds and a stout caudal process (Remeikis et al., 2009). DISTRIBUTION. Costa Rica (Pacific coast). MATERIAL EXAMINED. Holotype ♂, Costa Rica, Pacific Coast, S of Quepos, Manuel Antonio, 25.iii.2008, at light, Stonis and Hill, genitalia slide no. RA216 (VPU). Paratypes, 6 ♂♂, data as holotype, genitalia slides no. RA213, RA214, RA215, RA217, RA218, RA219 (VPU). ETYMOLOGY. The species name is derived from the Latin robusta (stout) in reference to the large and strongly thickened gnathos in the male genitalia.

40 A B

C D Fig. 15. Pseudopostega robusta Remeikis & Stonis, Costa Rica: A – adult; B – male genitalia, holotype, slide No. RA216; C – gnathos, ventral view, holotype, slide No. RA216; D – gnathos, lateral view, paratype, in glycerol

41 5.1.1.1.2. Pseudopostega mexicana Remeikis & Stonis (Fig. 16)

MALE. Forewing 1.7–1.8 mm; wingspan 4.0–4.3 mm. Head: vestiture entirely white; scape white, flagellum cream to pale beige; palpi white. Thorax and tegulae white. Forewing white, with two to three fuscous and almost straight subapical strigulae, one curved or straight apical strigula and black apical dot; underside of forewing densely irrorated with fuscous scales; pale brown. Hindwing and cilia greyish brown. Forelegs darkened with fuscous scales; midlegs cream, with pale brown banding on terminal tarsal segments; hindlegs cream. FEMALE. Unknown. MALE GENITALIA. Uncus a pair of relatively small, rounded, setose lobes, separated at a distance of 0.4 length of cucullar lobe. Vinculum narrowed, distally truncate. Gnathos with triangular basal part and large, distally broadened caudal process; basal fold absent; gnathos thickened laterally (but not anteriorly). Valva with large cucullar lobe, bearing a pectinifer consisting of 36–38 blunt spines; distal apex of cucullar lobe swollen; pedicel short and slender; saccular lobe short, triangular; valva short. Juxta absent or poorly developed. BIOLOGY. Adults fly in November. Otherwise unknown. DIAGNOSIS. Pseudopostega mexicana belongs to the spatulata group and most resembles P. spatulata Davis & Stonis and P. truncata Davis & Stonis but differs in the combination of a large cucular lobe of the valva, a narrow vinculum and a non thickened gnathos anteriorly (Remeikis et al., 2009). DISTRIBUTION. Mexico (Pacific coast). MATERIAL EXAMINED. Holotype ♂, Mexico, Oaxaca Region, Pacific Coast, Puerto Angel, secondary forest, 29.xi.2008, at light, Stonis and Hill, genitalia slide no. RA201 (VPU). Paratype, 1 ♂, data as holotype, genitalia slide no. RA202 (VPU). ETYMOLOGY. The species name refers to the country from which the type series originated (Mexico).

42 A B

C D

Fig. 16. Pseudopostega mexicana, holotype, Mexico: A – capsule,slide No. RA201; B – uncus, slide No. RA201; C – part of cucullar lobe of valva, slide No. RA201; D – forewing

43 5.1.1.1.3. Pseudopostega latiplana Remeikis & Stonis (Figs 17, 18) MALE. Forewing 1.8–2.0 mm; wingspan 4.0–4.4 mm. Head: vestiture entirely white; scape white, flagellum cream; palpi cream to white. Thorax and tegulae white. Forewing white, with dark brown oblique fascia, three fuscous subapical strigulae, black elongated apical dot and three short apical strigulae; underside of forewing greyish brown; cilia grey. Hindwing and cilia greyish brown. Forelegs fuscous; midlegs with brown darkenings distally; hindlegs cream. FEMALE. Unknown. MALE GENITALIA. Uncus a pair of large, rounded, setose lobes, separated by narrow excavation. Vinculum rounded. Gnathos consisting of a very large, anteriorly deeply divided lobe, with a stout caudal process and a large, broadly rounded basal fold. Valva with large cucullar lobe, bearing a pectinifer consisting of 44–50 blunt spines. Juxta absent. BIOLOGY. Adults fly in November. Otherwise unknown. DIAGNOSIS. Pseudopostega latiplana belongs to the saltatrix group, but differs in the large lobes of the uncus and gnathos, which comprises a very large, anteriorly deeply divided lobe, with a stout caudal process and large, broadly rounded basal fold (Remeikis et al., 2009). DISTRIBUTION. Mexico (Pacific Coast). MATERIAL EXAMINED. Holotype ♂, Mexico, Oaxaca Region, Pacific Coast, Puerto Angel, secondary forest, 29.xi.2008, at light, Stonis and Hill, genitalia slide RA204 (VPU). Paratype, 1 ♂, data as holotype, genitalia slide no. RA205 (VPU). ETYMOLOGY. The species name is derived from the Latin lata (broad, wide) and plana (flattened) in reference to the prominent, lateral lobes of the uncus and gnathos.

17 pav. Adult of Pseudopostega latiplana, holotype, Mexico

44 A B

C D

Fig. 18. Male genitalia of Pseudopostega latiplana, Mexico: A – capsule, holotype, slide No. RA204; B – gnathos, lateral view, paratype, in glycerol; C–D – uncus and gnathos, holotype, slide No. RA204

5.1.1.2. Geographical distribution of Central American Opostegidae

Prior to the study by Davis and Stonis (2007), the family Opostegidae had never been revised nor reviewed for Central or South America. As a result of their taxonomic revision of the New World fauna, 4 genera and 91 species of Opostegidae were reported for North, Central and South America (including 1 new genus and 68 new species, 2 new subspecies). Recently one additional species was described from Guatemala (Heppner, Davis, 2009). The greatest diversity of opostegid species was

45 reported for the Neotropics (a total of 68), including an unusually large number of species for Costa Rica, which were also briefly discussed in Stonis et al. 2008. Based on new collections in this thesis (after Remeikis et al., 2009, 2010) we provide a revised checklist of the Central American Opostegidae, with distributional data for the species now known to occur outside of Central America (Table 1). A total of 40 species of Opostegidae are reported in the checklist, with the greatest number of species occurring in Costa Rica (26 spp) (Fig. 19).

Fig. 19. Number of currently known opostegid species in the countries of Central America and the Caribbean (Abbreviations in ISO 2-alpha): MX – Mexico, BZ – Belize, GU – Guatemala, NI – Nicaragua, CR – Costa Rica, PA – Panama, US – Florida, CU – Cuba, JM – Jamaica, DM – Dominica, PR – Puerto Rico, VI – United States Virgin Islands, VG – British Virgin Islands, GD – Grenada, TT – Trinidad and Tobago) (Remeikis et al., 2009)

Fig. 20. Distribution map of recently described new species (Remeikis et al., 2009) (a dot – Pseudopostega robusta, a star – P. mexicana and P. latiplana)

46 Table 1. Checklist of the Central American Opostegidae, with distributional data for the taxa now known to occur outside of the region

Caribbean Central America South America No. Genera and species US CU JM DM PR VI VG GD TT MX BZ GT NI CR PA GF GY VE CO EC PE BR PY AR . Neopostega Davis & Stonis, 2007    1 falcata Davis & Stonis, 2007  2 petila Davis & Stonis, 2007  3 distola Davis & Stonis, 2007   4 nigrita Heppner & Davis, 2009  Pseudopostega Kozlov, 1985                        5 rotunda Davis & Stonis, 2007   6 serrata Davis & Stonis, 2007    7 lateriplicata Davis & Stonis, 2007  8 robusta Remeikis & Stonis, 2009  9 microacris Davis & Stonis, 2007  10 fumida Davis & Stonis, 2007  11 diskusi Davis & Stonis, 2007  12 spatulata Davis & Stonis, 2007  13 mexicana Remeikis & Stonis, 2009  14 attenuata Davis & Stonis, 2007    15 conicula Davis & Stonis, 2007  16 breviapicula Davis & Stonis, 2007    17 tanygnatha Davis & Stonis, 2007  18 saltatrix (Walsingham, 1897)             19 dorsalis Davis & Stonis, 2007  20 parakempella Davis & Stonis, 2007   21 latiplana Remeikis & Stonis, 2009  22 adusta (Walsingham, 1897)       23 longipedicella Davis & Stonis, 2007   24 lobata Davis & Stonis, 2007      25 sublobata Davis & Stonis, 2007   26 duplicata Davis & Stonis, 2007  

US CU JM DM PR VI VG GD TT MX BZ GT NI CR PA GF GY VE CO EC PE BR PY AR

27 tenuifurcata Davis & Stonis, 2007  28 concava Davis & Stonis, 2007  29 brevifurcata Davis & Stonis, 2007  30 brevivalva Davis & Stonis, 2007  31 bidorsalis Davis & Stonis, 2007  32 latifurcata Davis & Stonis, 2007      33 latiapicula Davis & Stonis, 2007   34 pumila (Walsingham, 1914)  35 bicornuta Davis & Stonis, 2007  36 constricta Davis & Stonis, 2007  37 brachybasis Davis & Stonis, 2007  38 venticola (Walsingham, 1897)           39 elachista (Walsingham, 1914)  40 perdigna (Walsingham, 1914) 

* – The species comprises two subspecies: P. latifurcata latifurcata Davis & Stonis, 2007 (currently known from Puerto Rico, Virgin Islands and Dominica) and P. latifurcata apoclina Davis & Stonis, 2007 (currently known from Costa Rica).

Abbreviations: MX – Mexico, BZ – Belize, GT – Guatemala, NI – Nicaragua, CR – Costa Rica, PA – Panama, US – Florida, CU – Cuba, JM – Jamaica, DM – Dominica, PR – Puerto Rico, VI – United States Virgin Islands, VG – British Virgin Islands, GD – Grenada, TT – Trinidad and Tobago, GF – French Guiana, GY – Guyana, VE – Venezuela, CO – Colombia, EC – Ecuador, PE – Peru, BR – Brazil, PY – Paraguay, AR – Argentina

48 Pseudopostega adusta (Walsingham, 1897) New to Costa Rica (this species previously was known from Cuba, Dominica, US Virgin Islands, Belize and Ecuador). Material examined: 11 ♂♂, Costa Rica: Pacific Coast, S of Quepos, Manuel Antonio, 25.iii.2008, at light, Stonis and Hill, genitalia slides RA206, RA207, RA208, RA209, RA210, RA211, RA212, RA232, RA233, RA234, RA235 (VPU) (Fig. 21).

A

B C

Fig. 21. Pseudopostega adusta, male genital structures: A – caudal part of capsule; B – capsule; C – gnathos, ventral view

Pseudopostega parakempella Davis & Stonis, 2007 New to Mexico (this species previously was known only from the type locality – Florida, U.S.A.). Material examined: 1 ♂, Mexico, Oaxaca Region, Pacific Coast, Puerto Angel, secondary forest, 29.xi.2008, at light, Stonis and Hill, genitalia slide RA203 (VPU) (Fig. 22).

Fig. 22. Pseudopostega parakempella, male genitalia, ventral view

50 5.1.1.3. New distribution data of the Neotropical Opostegidae

Pseudopostega lobata Davis & Stonis, 2007 (Fig. 23) New to Ecuador (this species previously was known from Belize, Nicaragua, Costa Rica and Argentina). Material examined: 1 ♂, Ecuador: Oriente, Napo Region, East of Tena, Misahualli, at light, 06-09.ii.2007, Sruoga, Stonis and Hill, genitalia slide RA222 (VPU).

Fig. 23. Pseudopostega lobata. Male genital structures (caudal part of capsule), ventral view

Pseudopostega didyma Davis & Stonis, 2007 (Fig. 24) New to the Amazonian Oriente Region of Ecuador (this species previously was known only from the type locality – a moist tropical site located on the western slopes of the Andes of Ecuador). Material examined: 1 ♂, Ecuador: Oriente, Napo Region, East of Tena, Misahualli, at light, 06- 09.ii.2007, Sruoga, Stonis and Hill, genitalia slide RA225 (VPU); 1 ♂, Ecuador: 80 km E of Guayaquil, Bucay (= Cumanda), 700 m, at light, 18.ii.2007, Sruoga, Stonis & Hill, genitalia slide RA226 (VPU).

51

Fig. 24. Pseudopostega didyma. Male genital structures (caudal part of capsule), ventral view

New data provided in this thesis (after published paper by Remeikis et al., 2009) increases the number of the world Opostegidae fauna to 202. Of the currently known 202 species, approximately 88% occur in subtropical to tropical regions (Fig. 25). Of these, 87 species (or 43% of the world fauna) are known to be restricted to the Neotropical Region. Currently, the Central American fauna comprises 2 genera and 40 species (i.e., 19.8 % of the opostegid species for the World). More than half of these (26 or 65%) are recorded from Central America alone and only a few of the Central American Opostegidae species now are known to possess a broader distribution (Table 1). An unusually large number of species was reported for Costa Rica (26 species). According to the current distributional data Costa Rican species fall into 4 groups. Half of the species (13 or 50%) are recorded from Costa Rica alone and only a few of these are known to possess broader distributions. One species known from Costa Rica also occurs in neighboring Panama, two species (7.7%) from Central America (Costa Rica) and the Caribbean, seven species (27%) are known from Central America and South America (but not the Caribbean), three species (11.5%) are known to have with a transneotropical distribution. These data most likely reflect the lack of adequate field surveys in other regions of the Neotropics (Davis, Stonis, 2007; Remeikis et al., 2009). No other Neotropical area has been subjected to such intensive collecting efforts for Microlepidoptera as has been conducted in certain areas of Costa Rica. The of La Selva survey, for example, resulted in 17 species of Opostegidae (of which only Pseudopostega saltatrix was previously known) from a very restricted area of Heredia Province, Costa Rica (Davis, Stonis, 2007).

52 New taxonomic data provided in this paper make up 1.5% for Opostegidae of the world and 3.4%, 7.5%, 7.7% and 27% for Opostegidae of the Neotropical Region, Central America, Costa Rica and Mexico, respectively (Table 2).

Fig. 25. Number of currently known opostegid species in the countries of the Neotropics (Abbreviations in ISO 2-alpha): MX – Mexico, BZ – Belize, GU – Guatemala, NI – Nicaragua, CR – Costa Rica, PA – Panama, US – Florida, CU – Cuba, JM – Jamaica, DM – Dominica, PR – Puerto Rico, VI – United States Virgin Islands, VG – British Virgin Islands, GD – Grenada, TT – Trinidad and Tobago, GF – French Guyana, GY – Gayana, VE – Venesuela, CO – Colombia, EC – Ecuador, PE – Peru, BR – Brasil, PY – Paraguay, AR – Argentina, CL – Chile

53

Table 2. New contribution to the Opostegidae fauna

Fauna Number of currently New discoveries known species (in total)

World fauna 203 (100%) 3 (1.5%)

Fauna of Costa Rica 26 (100%) 2 (7.7%)

Mexican fauna 11 (100%) 3 (27%)

Central American fauna 40 (100%) 3 (7.5%)

54 5.1.2. Nepticulidae of Central and South America

The checklist of the Nepticulidae of Central and South America included seven genera and 58 species (Puplesis, Robinson, 2000). However, it was followed by two later papers (including a review of the Neotropical fauna) (Puplesis et al., 2002a, b), where 16 new species were added: Enteucha acuta, E. guajavae, Stigmella austroamericana, S. montanotropica, S. nubimontana, S. rubeta, Fomoria repanda, F. tabulosa, Acalyptris ecuadoriana, A. onorei, A. basihastatus, A. pseudohastatus, A. articulosus, A. rotundus, A. amazonius, A. insolentis) (Puplesis et al., 2002a). Thus the most recent chekclist includes 74 species (Puplesis et al., 2002 b). Examination of material collected from South America (mostly from Patagonia: Argentina) yielded 8 species of Nepticulidae (including seven new to science) (Fig. 29). Therefore, in this thesis 82 species of Nepticulidae can be recognized for the Neotropical fauna (Table 3). All newly discovered American nepticulid species are left unnamed and unillustrated until they will be published in a scietific paper (Remeikis, in prep.). All newly recognized species belong to same genus (Stigmella) and two species groups (S. salicis group and an undesignated group).

Table 3. Taxonomic checklist of currently known Nepticulidae taxa in the Neotropical Region with species distribution data to Central and South American countries: US – Florida and Arizona only (U.S.A.), MX – tropical regions of Mexico, BZ – Belize, DM – Dominica, GY – Guyana, VE – Venezuela, CO – Colombia, EC – Ecuador, PE – Peru, AR – Argentina, CL – Chile (Country Codes in ISO 2-alpha) Genera and species U M B D G V C E P A C S X Z M Y E O C E R L Enteucha Meyrick      1. cyanochlora Meyrick, 1915  2. gilvafascia (Davis, 1978)  3. hilli Puplesis & Robinson, 2000  4. contracolorea Puplesis &  Robinson, 2000 5. terricula Puplesis & Robinson,  2000 6. snaddoni Puplesis & Robinson,  2000

55 7. acuta Puplesis & Diškus, 2002  8. guajavae Puplesis & Diškus, 2002  Manoneura Davis      9. basidactyla (Davis, 1978)     10. trinaria Puplesis & Robinson,  2000 Stigmella Schrank          11. plumosetaeella Newton &   Wilkinson, 1982 12. barbata Puplesis & Robinson,  2000 13. austroamericana Puplesis &  Diškus, 2002 14. kimae Puplesis & Robinson, 2000  15. eurydesma (Meyrick, 1915)  16. albilamina Puplesis & Robinson,  2000 17. fuscilamina Puplesis & Robinson,  2000 18. johannis (Zeller, 1877)  19. epicosma (Meyrick, 1915)  20. cuprata (Meyrick, 1915)  21. andina (Meyrick, 1915)  22. olyritis (Meyrick, 1915)  23. rudis Puplesis & Robinson, 2000   24. marmorea Puplesis & Robinson,  2000 25. peruanica Puplesis & Robinson,  2000 26. schoorli Puplesis & Robinson,  2000 27. hamata Puplesis & Robinson,  2000

56 28. imperatoria Puplesis & Robinson,  2000 29. montanotropica Puplesis &  Diškus, 2002 30. nubimontana Puplesis & Diškus,  2002 31. rubeta Puplesis & Diškus, 2002  32. maya Remeikis & Stonis, 2013  33. Species nova by Remeikis &  Stonis 34. Species nova by Remeikis &  Stonis 35. Species nova by Remeikis &  Stonis 36. Species nova by Remeikis &  Stonis 37. Species nova by Remeikis &  Stonis 38. Species nova by Remeikis &  Stonis 39. gossypii (Forbes & Leonard,  1930) 40. hylomaga (Meyrick, 1931)  41. costalimai (Bourquin, 1962)  42. guittonae (Bourquin, 1962)  43. pruinosa Puplesis & Robinson,  2000 44. ovata Puplesis & Robinson, 2000  Ectoedemia Busck    45. mesoloba Davis, 1978  46. reneella Wilkinson, 1981  47. helenella Wilkinson, 1981  48. species 291058 

57 49. fuscivittata Puplesis & Robinson,   2000 Fomoria Beirne    50. molybditis (Zeller, 1877)  51. diskusi Puplesis & Robinson, 2000  52. species 29122  53. repanda Puplesis & Diškus, 2002  54. tabulosa Puplesis & Diškus, 2002  Acalyptris Meyrick    55. latipennata (Puplesis & Robinson,  2000) 56. dividua Puplesis & Robinson,  2000 57. ecuadoriana Puplesis & Diškus,  2002 58. onorei Puplesis & Diškus, 2002  59. bicornutus (Davis, 1978)  60. tenuijuxtus (Davis, 1978)  61. bovicorneus Puplesis & Robinson,  2000 62. martinheringi Puplesis & Robinson,  2000 63. fortis Puplesis & Robinson, 2000  64. hispidus Puplesis & Robinson,  2000 65. novenarius Puplesis & Robinson,  2000 66. lascuevella Puplesis & Robinson,  2000 67. bifidus Puplesis & Robinson, 2000  68. trifidus Puplesis & Robinson,  2000 69. unicornis Puplesis & Robinson, 

58 2000 70. laxibasis Puplesis & Robinson,  2000 71. species 29135  72. platygnathos Puplesis &  Robinson, 2000 73. species 29140  74. basihastatus Puplesis & Diškus,  2002 75. pseudohastatus Puplesis &  Diškus, 2002 76. articulosus Puplesis & Diškus,  2002 77. rotundus Puplesis & Diškus, 2002  78. amazonius Puplesis & Diškus,  2002 79. insolentis Puplesis & Diškus, 2002  80. jucatani Remeikis & Stonis, 2013  Glaucolepis Braun   81. aerifica (Meyrick, 1915)  82. argentosa Puplesis & Robinson,  2000

Previous study of Neotropical material (Puplesis et al., 2002 b; Puplesis, Diškus, 2003) has shown a high level of endemism of the Nepticulidae fauna in general, and also demonstrated the phenomenon of Acalyptris predomination (Fig. 26). For example, among the recognized species in Belize (Puplesis, Robinson, 2000), 48% belong to the genus Acalyptris, in the western part of the Amazon basin this genus comprises about 50% of the fauna (Puplesis et al., 2002 b). Earlier, Acalyptris was known with a highly restricted distribution that excluded the Neotropics (or, indeed, any tropical area) and also was not known as very diverse in species. Our new addition to the fauna from the Andean part of Argentina dramatically increased species number known for fauna of the country (Figs 28, 29), slightly increased predomination of Stigmella in the Neotropics but slightly decreased the phenomenon of Acalyptris predomination (Fig. 27).

59

Glaucolepis Enteucha

11% 3% Manoneura

2%

Acalyptris 34%

Stigmella 36%

Fomoria 7% Ectoedemia 7% Fig. 26. Taxonomic composition of previously known fauna of the Neotropical Nepticulidae (after Puplesis et al., 2002 b)

Glaucolepis Enteucha 2% 10% Manoneura 3%

Acalyptris 31%

Stigmella 42% Fomoria 6%

Ectoedemia 6%

Fig. 27. Taxonomic composition of currently recognized fauna of the Neotropical Nepticulidae

60

35

30

25

20

15 29

10 18 5

11 5 9 7 2 2 0 1 1 1 1 US MX BZ DM GY VE CO EC PE AR CL

Fig. 28. Number of currently known nepticulid species in the countries of the Neotropics (with seven new species to the fauna of Argentina). Abbreviations in ISO 2-alpha: US – Florida and Arizona (U.S.A.), MX – Mexico, BZ – Belize, DM – Dominica, GY – Gayana, VE – Venesuela, CO – Colombia, EC – Ecuador, PE – Peru, AR – Argentina, CL – Chile

50

45

40

35

30

25

20 38

15 Number of species described

10 16 5 8 7 5 2 2 2 2 0 1 1

1877 1915 1930 1931 1962 1978 1981 1982 2000 2002 2011

Fig. 29. Chronology of nepticulid species discovery in the Neotropics (with seven undescribed new species to the fauna of Argentina in 2013)

61 5.2. NEW DATA TO THE SOUTH EASTERN EUROPEAN FAUNA OF THE NEPTICULIDAE

5.2.1. The Easternmost record of the Sub-mediterranean species Acalyptris platani (Müller-Rutz) in Europe

5.2.1.1. European Acalyptris as representatives of the Mediterranean and Sub-mediterranean fauna Althought genus Acalyptris Meyrick is diverse in tropical regions, including the Neotropics, (Šimkevičiūtė et al., 2009; Stonis, Remeikis, 2011a) and also abundant in desert and steppe regions of Central Asia (Puplesis 1990, 1994; Puplesis, Diškus, 1995, 2003), in Europe, Acalyptris forms a small fraction (in total 9 species) of the nepticulid fauna. Most of Acalyptris species occur in Greece (7), also Spain (4) and Croatia (4) (Fig. 30).

6

4

Number of species 2

0

Italy

Spain

Czech

France

Austria Greece

Croatia Cyprus

Ukraine

Bulgaria

Portugal Slovakia

Hungary Slovenia

Romania Germany

Switzerland

Fig. 30. Number of currently known Acalyptris species in the countries of Europe (after Stonis, Remeikis, 2011a)

All European Acalyptris species mainly occur within Mediterranean or Sub-mediterranean regions. The latter region represents a large ecoregion in the temperate mixed forest biome (with hot dry summer and mild, rainy winter).

62 5.2.1.2. Documentation of the Acalyptris platani specimens collected in the Karadag Reserve

Males (Fig. 31 B). Forewing length 1.9–2.3 mm, wingspan 4.2–5.0 mm. Head: palpi cream; frontal tuft brown-cream, on vertex brown to fuscous (differently coloured tufts distinctly separated); collar inconspicuous, comprises of dark cream piliform scales; scape and pedicel yellowish cream (not white as usually in the species); flagellum pale grey-brown in proximal half, dark cream in distal half; antenna with 28–29 segments (not 34, as usually in the species). Thorax yellowish cream (not brown, as common in the species). Forewing basal 1/4 yellowish cream densely irrorated with pale brown scales, followed by broad yellowish cream fascia to 1/2; distal part of wing fuscous to brown apically, with yellowish cream tornal and costal spots united in a second fascia; cilia shiny cream; cilia line, in contrast to the general description of the species (van Nieukerken, 2007), indistinct or absent. Underside of forewing cream. Hindwing very broad at basal 2/3, distinctly and abruptly cuspidate towards tip; costal bristles present (brown); upper surface of the whole broadened basal

A B Fig. 31. Adult specimens of Acalyptris platani recorded from the Crimea: A – female; B – male (right side, with broadened hindwing covered with white androconial scales) part (2/3 of hindwing) except anal margin, covered with a thick indumentum of white raised androconial scales, irrupted along midline by a distinct, straight, longitudinal furrow, without scales; costal margin of hindwing with a row of short pale brown broadened (not piliform) scales instead of cilia. Underside cream. Abdomen brown, with yellow inconspicuous anal tufts. Legs yellow-cream. Females (Fig. 31 A). Forewing length 2.2–2.3 mm, wingspan 4.7–4.9 mm. Antenna with ca. 26 segments. Hindwing lancelolate, without white androconial scales. Male genitalia (Fig. 32). Vinculum anteriorly and posteriorly concave, but in the specimens from Karadag anterior incision tends to be shallower than usual. After Stonis, Remeikis (2011a) otherwise as in the species descriptions given by Johansson et al. (1990), Laštuva, Laštuvka (1997)

63 and van Nieukerken (2007). Tegumen narrowly rounded, forming specific pseuduncus; uncus band- shaped, with short central process (indistinct in ventral view). Gnathos with long and almost pointed central element (caudal tip usually weakly sclerotized in the specimens from Karadag) and with a prominent inner lobe at base. The shape of the inner lobe may vary, usually right and left lobes slightly asymmetrical. Transtilla without transverse bar but, in contrast to the descriptions by Johansson et al. (1990), Laštuva, Laštuvka (1997) and van Nieukerken (2007), sublateral process of valva less deve loped, almost invisible and right valva joined with left one via sclerotized juxta.

A B Fig. 32. Male genitalia of Acalyptris platani, Karadag Reserve, 44o54’N 35o13’E, the Crimea: A – capsule; B – aedeagus (genitalia slide no. RA238, RA239, RA240, RA241)

Carinae of aedeagus ending in tiny forked lobes, tightly fused to ventral process; pair of lateral carinae straight and pointed. Vesica with numerous very small triangular cornuti and one large cornutus. Female genitalia (Fig. 33). Anal papillae separated by a shallow square-like excavation. Vestibulum with vaginal sclerotization comprising three plates. The main, “nose”-shaped, broad and

64 tends to be dentate on proximal margin. Otherwise as in the species descriptions given by van Nieukerken (2007). Total bursa length ca. 740μm. Corpus bursae elongate, without pectinations, with very narrow and long reticulate signa (length ca. 380–400 μm), margins crenate. Ductus spermathecae with 2 convolutions and long and conspicuous vesicle (as in the description by van Nieukerken (2007) (broken in slides no. RA243 and RA243, therefore, not shown in Fig. 33).

A B Fig. 33. Female genitalia of Acalyptris platani, Karadag Nature Reserve, 44o54’N 35o13’E, the Crimea: A – general view; B – details of caudal part (genitalia slide no. RA242, RA243)

Bionomics. Host-plants: Platanus spp. (icluding P. orientalis, P. hybrida and also P. x acerifolia Willd., P. digitifolia Palib., P. occidentalis L. planted in Karadag Reserve) (Stonis, Remeikis, 2011). Eggs on leaf underside, usually against a vein. Leaf-mine (Fig. 34 A–C) as a long gallery with contorted green or brown (when dried) frass. Larva (while feeding) yellowish with green or dark green central line and brown head; before pupation larva turns to brownish yellow.

65 Cocoon yellowish brown (Fig. 34 D). Mortality rate of the larvae collected in the Karadag Reserve in 2009 was 26%; of the cocoons reared indoors – 29%. In total, mortality rate for the whole sample collected in Karadag Reserve was 55 %. The species is most likely bivoltine; in Karadag, fresh mines were found in August together with old mines from the previous season.

A B

C D

Fig. 34. Leaf-mines and cocoon of Acalyptris platani, Karadag Nature Reserve, 44o54’N 35o13’E, the Crimea: A–C – leaf-mines; D – a cocoon (sample/field card no. 4965)

Distribution (Fig. 35). Acalyptris platani exhibits Mediterranean or typical Sub- mediterranean distribution (Stonis, Remeikis, 2011a): until now it has been known from Portugal, Spain (incl. Menorca), France (incl. Corsica), Italy (incl. Sicilia), Switzerland, Croatia, Greece, Bulgaria and Cyprus (Laštuvka, Laštuvka, 1997; van Nieukerken, 2007). Following van Nieukerken (2007), a supposed record from the Netherlands was based on misidentification, and the northernmost locality for A. platani has for more than 70 years been the region of Paris. In the current paper, the record from the Crimea (Ukraine) represents the easternmost locality within Europe (Stonis, Remeikis, 2011a). However, A. platani is also reported in western Asia (van

66 Nieukerken, 2007): Turkey, Iran and Georgia. We suposse that the later record of A. platani in George still needs verification.

Fig. 35. Distribution of Acalyptris platani, with newly recorded locality in the Crimea (South Eastern Europe) (after Stonis, Remeikis, 2011)

Fig. 36. Habitat of Acalyptris platani in Karadag Nature Reserve, 44o54’N 35o13’E, Ukraine

Material examined. 4♂ 3♀, UKRAINE: Crimea, 20 km SW Feodosiya, Karadag Reserve (Botanical Park of the Reserve, i.e. “Biostanciya”), on Platanus spp. (incl. P. x acerifolia Willd., P. digitifolia Palib., P. occidentalis L.), 12–26 August 2009, sample/field card no. 4965, genitalia slide

67 no. RA238, RA239, RA240, RA241 (males), RA242, RA243 (females), leg. A. Remeikis, J. R. Stonis. Other 2 ♀ specimens are preserved in 70% ethanol and kept in a freezer for molecular studies.

5.2.1.3. Discussion about Acalyptris platani in Karadag

Acalyptris platani collected in the Karadag Nature Reserve represents an isolated population of the species with generally Sub-mediterranean distribution (Stonis, Remeikis, 2011a), and slightly differs from the general description of A. platani given in the taxonomic revision by Erik van Nieukerken (van Nieukerken 2007). In contrast to the description, the males from Karadag are smaller, antennae have less flagellomeres, the cilia-line of forewing is absent, the costal margin of the hindwing is with a row of short broadened (not piliform) scales, the forewing pattern and colouration of antennae are paler. However, differences in colouration could possibly be caused by the climate of Karadag: hot dry summer, cool rainy winter. The discovery of Acalyptris platani in the Crimea supports the concept of the South Crimean Nepticulidae fauna as mostly Sub- mediterranean (or Mediterranean) (Stonis, Remeikis, 2011a).

5.2.2. A re-assessment of the Crimean fauna of Nepticulidae, with new records for the peninsula

The recently published updated checklists of the Nepticulidae of Europe and Asia (Van Nieukerken, Puplesis, 1991; Puplesis, 1994; Puplesis, Diškus, 2003, etc.) or Karadag Nature Reserve (Budaskin, 2004) record 35 species for fauna of the Crimea. Most of them (19 species) belong to Stigmella, the remaining to Simplimorpha (1), Ectoedemia (9), Trifurcula (2), Glaucolepis (2) and Etainia (2) (Table 4). Our fieldwork in Karadag Nature Reserve and Sudak (2009) and study of material collected from the Crimea by other researchers (Yu. Budashkin, J. Sinev, A. Zagulaev, J. R. Stonis, formerly R. Puplesis) confirmed the occurrence only of 17 Nepticulidae species (Fig. 37). However, four of these species are new to the Crimean fauna: Stigmella ulmiphaga, S. plagicolella, Acalyptris plantani and Ectoedemia albifasciella. Therefore, the Crimean fauna of Nepticulidae currently comprises 39 species (not 35); it was published in our publication in co-authorship with other researchers (Stonis et al., 2013).

68

25

20 New addition to the fauna

15 Species confirmed by our material 10 Previously known species 5

Number Number of nepticulid species 0

Etainia

Stigmella

Trifurcula

Acalyptris Glaucolepis

Ectoedemia Simplimorpha

Fig. 37. Taxonomic composition of the Crimean fauna of Nepticulidae confirmed by our material (collected during our fieldwork and by other researchers)

A B Fig. 38. Male genitalia of Stigmella dorsigutella, Karadag Nature Reserve, the Crimea, 44°54'N 35°13'E: A – capsule; B – aedeagus (genitalia slide no. RA 251)

69 Table 4. Particularities of geographical distribution of the Nepticulidae fauna discovered in the Crimea Geographical distribution (% of species Simplimorpha Stigmella Acalyptris Ectoedemia Trifurcula Glaucolepis Etainia within the fauna) Species with a wide European - 90,4% - 80% - - 50% distribution Species with a limited (mostly 100% 9,6% 100% 20% 100% 100% 50% Mediterranean) distribution

The Crimea (particularly the southern slopes of the penninsula) is recognized as the area in South Eastern Europe most diverse in nepticulid species. However, 71,8% of the Crimean fauna is represented by species with a wide European or Euro-Asian distribution, in contrast to the Central Asian nepticulid fauna which characterized in average by from 71% of endemic species. However, a restricted geographical distribution (mainly Mediterranean) prevails among the Crimean species of Simplimorpha, Acalyptris, Trifurcula and Glaucolepis (Table 4).

5.3. CONTRIBUTION TO THE FAUNA OF NEPTICULOIDEA OF THE HIMALAYA

5.3.1. A re-assessment of Opostegidae of the Himalaya, with description of new species

India, while interesting with a hugely diverse biota, is comparatively unexplored with respect to collection and study of Opostegidae. The first nine species recorded from the region were described in 1907–1922. Much later, eight further species were reported by Puplesis, Robinson (1999) (Fig. 46). However, only nine species of 18 occur in the Himalaya (Fig. 47). The first species recorded from Himalaya was described in 1910 by E. Meyrick (In total Meyrick is responsible for description of 6 Himalayan opostegids). The biggest boost to species number of the Himalaya was given by Puplesis, Robinson (1999) who mostly recorded “Meyrick’s Indian” species for the Himalaya and described two new species. Currently (including our first discovery of Pseudopostega frigida in the Indian Himalaya in 2011 and description of one new species), Himalayan Opostegidae fall into three genera: Opostegoides (4 species), Pseudopostega (4 species) and Opostega (1 species).

70 This review of the Himalayan Opostegidae is the first attempt to demonstrate to the scientific community an example of the diversity of Himalayan opostegids and to prepare the ground for future investigations of this fauna. Opostegoides epistolaris (Meyrick, 1911) Diagnosis. This species is a little difficult to separate from many other Opostegoides species using only external features, but the shape of vaIva (notably the bulged inner lobe) in the male provides a good character for its distinction.

Material examined. INDIA: Lectotype ♂, [Nilgiri Hills], N. Coorg: Dibidi, 13.v.1907 (Newcome), genitalia slide no. 28625 (BMNH); paralectotype: 1♀ [recorded incorrectly as a male in original description], data as lectotype, genitalia slide no. 28639 (BMNH). Opostegoides pelorrhoa (Meyrick, 1915) Diagnosis. Differs from other Oriental species known by the irregularly marked forewing and relatively large cucullar lobes of the valva.In the female genitalia the configuration of the corpus bursae is unique within the Opostegidae. Material examined. INDIA: Holotype ♂, Assam, Khasi Hills, vii. 1906, genitalia slide no. 28653 (BMNH). NEPAL: Nepal: 2♂, 6♀ , Kathmandu District, Phulchoki, 27-31.v.1983 (Allen. Brendell, Robinson & Tuck), genitalia slide nos. 28683♂, 28684♀ (BMNH).

A B

71 C Fig. 39. Male genitalia of Opostegoides pelorrhoa: A – capsule; B – cucular lobe; C – aedeagus

Opostegoides index (Meyrick, 1922) Diagnosis. This species may be differentiated from all currently known Opostegoides by the presence of a brown costal spot on the forewing.

Material examined. INDIA. Ho1otype ♀, Assam: Shillong, 1922 (T.B.Fletcher), genita1ia slide no. 28652 (BMNH). Opostegoides species 1005 Diagnosis. Most similar to Opostegoides malaysiensis, but differing in the darker dorsal forewing spot and in the yellowish area between the first and second terminal strigulae; the costal spot is also darker than that of species 28641 but the single specimen known of the latter is damaged and comparison of the costal pattern is impossible (Puplesis, Robinson, 1999). Material examined. NEPAL: 1♀; 70 km W of Kathmandu, Baikuntapuri, 19.iv.1995 (Puplesis), genitalia slide no. RP1005 (VPU).

72 A B Fig. 40. Female genitalia of Opostegoides species 1005: A – caudal part; B – general view

Pseudopostega frigida (Meyrick, 1906) Diagnosis. After Puplesis, Robinson (1999) currently there are just three other species with a dark costal spot on the forewing known from the Oriental region: P. myxodes, P. nigrimaculella and Opostegoides index. Pseudopostega frigida differs from all of them in genital features. The male genital structure is most similar to that of similantis, but differs in the undivided gnathos plate and broader cucullar lobe of the valva (Fig. 41). Material examined. SRI LANKA: Lectotype ♂, Peradeniya, ii.1905 (Green), genitalia slide no. 28626 (BMNH). Paralectotypes: 10♂♀, data as lectotype (BMNH). SRI LANKA: 1 ex. (♂?), Maskeliya, ii.1906 (Pole) (BMNH). NEPAL: 2 ex., Chitwan National Park, Sauraha, 3–6.vi.1983 (Allen, Brendell, Robinson & Tuck) (BMNH); 1♂, 3♀, 70 km W of Kathmandu, Baikuntapuri, 19- 21.iv.1995 (Puplesis), genitalia slide nos RP1006, RP007, RP1008, RP1009 (VPU). THAILAND: 1♂, Chiang Mai, 325 m, 15–30.x.1984 (Karsholt et al.), genitalia slide no. Pupl.402 (ZMUC). INDONESIA: 1♂, eastern Sumba, Bajog, 25 m, vi.1949 (Sutter & Wegner), genitalia slide no. Pupl.012 (NNM); 1♀, westem Sumba, Wainangura, 450 m, viii.1949 (Sutter & Wegner) (NNM); 1♀, central Sumba, Lindi Watju, 27.ix–15.x.1949 (Sutter & Wegner) (NNM).

73 A

B C Fig. 41. Male genitalia of Pseudopostega frigida: A – capsule; B – gnathos; C – uncus

Pseudopostega brevicaudata Remeikis and Stonis, 2013 Male. Forewing length: 2.9–3.0 mm. Wingspan: 6.6 mm. Head: palpi cream; piliform scales on vertex, collar and scape white; flagellum grey-brown.Thorax and tegulae white. Forewing white with broad dark brown median fascia characterized by small triangular indentation in proximal margin which is marked by darker scales; with black apical dot and two parallel cilia-lines.

74 Underside of forewing dark 'brown, except basal third and apical margin; cilia shades of brown. Hindwing grey-brown, cilia pale ochreous grey. Legs pale yellowish ochre or ochreous cream; forelegs almost fuscous. Colour of abdomen unknown. Female. Unknown.

A B Fig. 42. Male genitalia of two mostly related Pseudopostega species: A – P. brevicaudata Remeikis and Stonis, 2013; B – P. frigida

Male genitalia (Fig. 42 A). Uncus with two tiny, slender and very widely separated lobes directed subcaudally; emargination between lobes rather shallow. Gnathos with caudally-directed, broad and distally truncated process covered by minute spines and bearing a large sclerotized plate with a shallow distaI emargination. Valva with large cucullar lobe bearing pectinifer and supported by ashort pedicel; pectinifer of about 24 spines arranged in a single row; valvallobe distally triangular; basal process of valva straight, pointed apically and relatively short,' moderately sclerotized; costaI process strongly sclerotized, shorter than basal process. Juxta damaged, probably weakly wrinkled and with slender elongate median process. Vinculum broadly rounded anteriorly Biology. The single specimen known was collected at light in June. Larval biology unknown.

75 Diagnosis. Externally very similar to euryntis, from which it differs by the small triangular proximal indentation on the fascia, darker (grey-brown) antenna, and greyish to grey hindwing. The male genital structure is most similar to that of P. frigida, but differs in the short lobes of uncus, stout process of gnathos; juxta absent. Distribution. Nepal (Kathmandu valley, 1400 m). Material examined. NEPAL: 1♂ holotype, Kathmandu, British Embassy, 1400 m, 12– 13.vi.1984 (Allen) genitalia slide no. 28623 (BMNH). Pseudopostega nepalensis Puplesis & Robinson, 1999 Diagnosis. The brown forewing and tuft of piliform scales on the head distinguish nepalensis from all other Oriental opostegids except the other members of the P. velifera group, P. velifera and P. sumbae. From these it differs by the very long spermathecal duct in the female genitalia (Fig. 43)

Fig. 43. Female genitalia of Pseudopostega nepalensis (genitalia slide no. RP1010 BMNH)

and, additionally, from P. velifera by the absence of irregular yellow-cream areas along the forewing costa.

Material examined. Holotype 1♀, NEPAL: 70 km W of Kathmandu, Baikuntapuri, 19–

76 20.iv.1995 (Puplesis) genitalia slide no. RPl0l1 ♀ (VPU). Paratypes: 1♂, 2♀, same data as holotype, genitalia slide no. RPl0l0 (VPU). Pseudopostega zelopa (Meyrick, 1905) Diagnosis. Despite some variation of forewing pattem, this species is extemally distinctive,

A

B Fig. 44. Genitalia of Pseudopostega zelopa: A – male; B – female characterized by the broad brown antemedian fascia and brown apical area of the forewing. In the male genitalia zelopa is distinguished by the inverted V- shaped gnathos in combination with a very long, distally slightly concave uncus and apically broadened juxta (Fig. 44). Material examined. Holotype ♀, SRI LANKA: Pundalu-oya (Green), v.1903 (BMNH). INDIA: 1♂, Assam: Khasi Hills, v.1907, genitalia slide no. 28621 (BMNH); NEPAL: 1♂, Terai, Dharan, secondary forest, 18.viii.1984 (Allen) (BMNH); 51♂♀, 70 km W of Kathmandu,

77 Baikuntapuri, 19.iv.1995 (Puplesis), genitalia slide nos. RPl001♂, RPl002♂, RPl012♀ (VPU). THAILAND: 1♀, Uthai Thani Distr., Khao Nang Rum, l.iii.1986 (Allen), genitalia slide no. 28622 (BMNH). INDONESIA: 1♀, SW Timor, xi–xii.1891 (Doherty) genitalia. slide no. 28806 (BMNH); Sumba: 2♂, 2♀, Lindi Watju, 27.ix–15 .x.l949 (Sutter & Wegner), genitalia slide nos. Pupl.004, Pupl. 020 (NNM); 2 ♂, 7 ♀, Melolo, v–vi.1949 (Sutter & Wegner) (NNM). Opostega chalcophylla Meyrick, 1910 Diagnosis. Differs from all known Oriental species in the uniformly brown forewings. In the male genitalia the species is recognizable by the distinctive shape of the juxta (abruptly broadened in distal half), very broad vaIva and the gnathos in the shape of an inverted 'V'. Distribution. Central and eastern Himalayas and Assam (eastern India and Nepal).

Fig. 45. Male genitalia of Opostega chalcophylla (genitalia slide no. 28662, BMNH)

Material examined. Lectotype ♂: INDIA: eastern Himalaya, Kurseong, 7.ix.1909 (N.A.), genitalia slide no. 28662 (BMNH). Paralectotype: 1♂, data as lectotype, genitalia slide no. 28811 (BMNH). 1♂ [?– abdomen in glue], Assam: Khasi Hills, 1906 (Doncaster) (BMNH); 1♂, Sikkim: Gangtok, 6000 ft, 21.v.28 (Bailey) (BMNH). NEPAL: 1 ex. [without abdomen], Phulchoki, 2000– 2500 m, oak-laurel forest, 22.v.1984 (Allen) (BMNH).

78 8

2 2 Number 1of species recorded 1 1 1 1 1

1907 1910 1911 1915 1916 1920 1922 1999 2011

Fig. 46. Pattern of species records of the Indian Opostegidae through time (Note compression of periods 1923–1998)

18

9 Number of species recorded

India Himalaya

Fig. 47. India (and the Himalaya), while interesting with hugely diverse biotas, are comparatively unexplored with respect to collection and study of Opostegidae: 18 opostegid species are known for India, and only nine recorded in the Himalaya

79 5.3.2. New Nepticulidae discoveries in the Himalaya

Our fieldwork in the Himalaya confirmed the occurrence of several Nepticulidae species (15 new to the fauna). Eight nepticulid species belonging to Stigmella, one to Enteucha, four species to Acalyptris and one species to Ectoedemia were recognized as new for science but they are omitted from the thesis until publication in a scietific journal (Remeikis et al., in prep.). The Himalayan trip, undertaken in August 2010 by the author of the thesis with companionship of five other members of Biosystematics Research Group (LEU), has also resulted in the discovery of a very interesting species – Fomoria festivitatis (van Nieukerken, 2008) – a new species for the Indian Himalaya (Figs 48–50).

Fig. 48. Leaf-mine of Fomoria festivitatis van Nieukerken, 2008 (the first record for the Indian Himalaya)

Fig. 49. Adult of Fomoria festivitatis (India: Uttarakand, Mussoorie)

80

A B

Fig. 50. Genitalia of Fomoria festivitatis (India: Uttarakand, Mussoorie): A – male (slide no. RA250); B – female (slide no. RA268)

81 5.4. THE MOST RECENT DISCOVERIES. DESCRIPTIONS OF NEW SPECIES FROM YUCATAN (MEXICO)

5.4.1. Stigmella maya Remeikis & Stonis, 2013

Type material. Holotype: ♂, MEXICO, Yucatán, Quintana Roo, Tulum, 20°12'35"N, 87°25'55"W, elevation 15 m, mining larvae 29.xi.2011, ex pupa 11.xii. 2011, leg. A. Remeikis & J.R. Stonis, gen. slide no. RA 473. Paratypes: 2♂, 3♀, same label data, ex pupae 8–16.xii.2011, gen. slide nos RA 472♂, RA 469♀, RA 470♀, RA 471♀ (all specimens of the type-series are currently deposited in LEUS, with further re-deposition at ZMUC). Diagnosis. A very distinctive, very small dark-winged species with a silver-white fascia in the forewing. Externally, males of the new species easily distinguished from all other currently known Stigmella species, including Neotropical ones, by black hindwings densely covered with androconial scales. In male genitalia, S. maya differs from all Neotropical Stigmella species by a combination of a specialized uncus, a broad U-shaped gnathos and a simple apically pointed valva. Male (Fig. 51 A, C). Forewing length 1.4–1.5 mm; wingspan 3.04–3.27 mm (n=3). Head: palpi cream; frontal tuft cream, on vertex brown to orange; collar and scape whitish to cream; antenna with 23–24 segments, slightly longer than half forewing; flagellum brown-grey to cream- glossy (particularly first 8 segments) on upper side, cream on underside. Thorax fuscous grey; tegulae black. Forewings fuscous grey, speckled in apical part, with silver-white slightly shiny fascia; terminal cilia white, comprised of broadened (lamellar) scales; tornal cilia blackish grey, comprising piliform scales; underside of forewing black. Hindwings fuscous grey, densely covered with black androconia, except at apex (Fig. 51 C); cilia of hindwings blackish grey. Legs dark cream to black on upper side, dark cream on underside. Abdomen black on upper side, dark grey, with purple and some green iridescence on underside; anal tufts white-cream. Female (Fig. 51 B). Forewing length 1.3–1.4 mm; wingspan 2.8–3.0 mm (n=3). Hindwings grey to blackish grey; no androconial scales. Abdomen shiny black on upper side, grey on underside, without purple and green iridescence. Otherwise as male. Male genitalia (Figs 51 E, F, 52 A,B). Capsule longer (215 mm) than wide (160 mm) (n=2). Vinculum with two large lateral (anterior) lobes. Uncus with small heavier sclerotised lateral (posterior) lobes. Gnathos narrow, broadly U-shaped, posterior projections far apart (Fig. 52 A). Valva (Fig. 52 B) 130 mm (n=2) long, basally broad, narrowed apically, with thickened and pointed apical process; transtilla with short triangular pointed sublateral processes. Aedeagus (Fig. 51 F) 230 mm (n=2); vesica with numerous minute cornuti.

82

A B

C D

E F

Fig. 51. Morphology of the new species: A – adult of Stigmella maya; B – the same; C – details of hindwing with adroconial scales; D – Acalyptris jucatani; - male genitalia of Stigmella maya, capsule, gen. slide No. RA 473; F – the same, aedeagus

83 Female genitalia (Fig. 52 C). Total length 575–585 mm (n=3). Vestibulum relatively broad, without sclerites. Corpus bursae broadly oval, very large, covered with numerous pectinations, without signa. Ductus spermathecae with 2.5 large convolutions. Bionomics. Mines in leaves. Host-plant: Karwinskia humboldtiana (Schult.) Zucc. () (Fig. 53 A). Egg on upper side of the leaf. Larvae mine in late November and early December. Contorted or sinuous gallery of mine filled with dark brown to blackish frass (Fig. 53 A). Larva pale green to green, with dark green intestine. Larval exit slit on upper side of the leaf. Cocoon very pale brownish cream to grey-beige; length 1.4 – 1.75 mm, maximal width 0.7–1 mm (n=6). Adults emerged in December. Distribution (Fig. 54). Lowland coastal forest of Yucatán (SE Mexico: Quintana Roo) (Fig. 53 C). Etymology. This species is named after the Maya people, a Mesoamerican civilization.

5.4.2. Acalyptris yucatani Remeikis & Stonis, 2013

Type material. Holotype: ♀, MEXICO, Yucatán, Quintana Roo, Tulum, 20°12'35"N, 87°25'55"W, elevation 15 m, mining larva 29.xi.2011, ex pupa 14.xii.2011, leg. A. Remeikis & J. R. Stonis, gen. slide no. RA 468♀ (currently deposited in LEUS, with further re-deposition at ZMUC). Diagnosis. Externally, the new species can be distinguished from most Acalyptris species, including Neotropical ones, by the presence of two white spots (costal and tornal) on the speckled forewing. In female genitalia it differs from all known Neotropical Acalyptris species by a combination of specialized large asymmetrical signae and a vestibulum without distinct sclerites. Male. Unknown. Female (Fig. 51 B). Forewing length 1.65 mm, wingspan 3.63 mm (n=1). Head: palpi brownish cream; frontal tuft cream, on vertex dark brown; collar whitish cream, comprising piliform scales; scape white; antenna with ca. 25 segments, slightly shorter than half forewing; flagellum dark grey on upper side, pale grey on underside. Thorax, tegulae and forewings cream- white, speckled with blackish brown tipped scales. Forewing with two cream-white oblique spots:

84

A B

C D

Fig. 52. Morphology of the new species: A – Stigmella maya, male genialia, slide No. RA 470, gnathos and uncus; B – the same, valva; C – the same, female genitalia; D – Acalyptris jucatani, female genitalia

85 costal and tornal; cilia grey; underside of forewing brownish cream, grey-brown distally and along costa. Hindwings grey (at certain angle of view may look pale grey) on upper side and underside; cilia of hindwings grey. Legs cream, with blackish darkening on upper side; forelegs darkened on both sides. Female genitalia (Fig. 52 D). Total length 565 mm (n=1). Vestibulum narrow, folded, without shaped sclerites. Corpus bursae oval, with very large, distinctly asymmetrical signae (245 and 350 mm) (n=1). Bionomics. Mines in leaves (Fig. 53 B). Host-plant: Schinus sp. (Anacardiaceae). Egg on upper side of the leaf. Larvae mine in late November and early December. Sinuous gallery of mine filled with dark green to blackish or black frass (Fig. 53 B). Larva yellow, with dark green intestine. Larval exit slit on upper side of the leaf. Cocoon very pale ochre-brown; length 1.65 mm, maximal width 1.1 mm (n=1). The single adult emerged in December. Distribution (Fig. 54). It occurs in the lowland coastal forest of Yucatán (SE Mexico: Quintana Roo) (Fig. 53 C). Etymology. This species is named after the Yucatán Peninsula.

A B

C Fig. 53. Bionomics of the new species: A – leaf-mine of Stigmella maya; B – the same, Acalyptris jucatani; C – habitat of the new species

86 5.4.3. Taxonomic discussion about recorded Nepticulidae species in Yucatan and adjacent regions

Currently 31 species of Nepticulidae are recorded from Yucatán (Puplesis and Robinson 2000) and seven others from the adjacent areas (Puplesis & Robinson 2000; Šimkevičiūtė et al. 2009): Enteucha hilli Puplesis & Robinson, 2000; E. contracolorea Puplesis & Robinson, 2000; E. snaddoni Puplesis & Robinson, 2000; Manoneura basidactyla (Davis, 1978); Stigmella plumosetaeella Newton & Wilkinson, 1982; S. barbata Puplesis & Robinson, 2000; S. kimae Puplesis & Robinson, 2000; S. albilamina Puplesis & Robinson, 2000; S. fuscilaminaPuplesis & Robinson, 2000; S. pruinosa Puplesis & Robinson, 2000; S. racemifera Šimkevičiūtė & Stonis, 2009; S. species 018 (described but unnamed in Šimkevičiūtė & Stonis 2009); S. maya Remeikis & Stonis, 2013.; Ectoedemia (Zimmermannia) species 29105 (described but unnamed in Puplesis & Robinson 2000); E. (Ectoedemia) fuscivittata Puplesis & Robinson, 2000; Fomoria diskusi Puplesis

Fig. 54. Nepticulidae of Yucatán and adjacent areas. 1, currently known distribution: 1. Stigmella maya, Acalyptris yucatani. 2. Enteucha hilli, E. contracolorea, Stigmella barbata, S. kimae, S. albilamina, S. fuscilamina, S. pruinosa, Ectoedemia sp. 29105, E. fuscivittata, Fomoria diskusi, F. sp. 29122, Acalyptris latipennata, A. dividua, A. bovicorneus, A. martinheringi, A. hispidus, A. novenarius, A. bifidus, A. trifidus, A. unicornis, A. laxibasis, A. sp. 29135, A. platygnathos, A. sp. 29140, Glaucolepis argentosa. 3. Enteucha snaddoni, Acalyptris fortis. 4. Manoneura basidactyla. 5. Acalyptris lascuevella. 6. Stigmella racemifera, S. sp. 018, Acalyptris paradividua, A. terrificus, A. sp. 015, A. sp. 016. 7. Stigmella plumosetaeella

87

A B

C D

E F

Fig. 55. The taxonomic composition of Nepticulidae: A – fauna of Yucatan; B – fauna of Florida (USA); C – fauna of the Crimea (SE Europe); D – fauna of Central Asia; E – fauna of East Asia; F – fauna of South Africa

88 & Robinson, 2000; F. species 29122 (described but unnamed in Puplesis & Robinson 2000); Acalyptris latipennata (Puplesis & Robinson, 2000); A. dividua Puplesis & Robinson, 2000; A. bovicorneus Puplesis & Robinson, 2000; A. martinheringi Puplesis & Robinson, 2000; A. fortis Puplesis & Robinson, 2000; A. hispidus Puplesis & Robinson, 2000; A. novenarius Puplesis & Robinson, 2000; A. lascuevella Puplesis & Robinson, 2000; A. bifidus Puplesis & Robinson, 2000; A. trifidus Puplesis & Robinson, 2000; A. unicornis Puplesis & Robinson, 2000; A. laxibasis Puplesis & Robinson, 2000; A. platygnathos Puplesis & Robinson, 2000; A. species 29135; A. species 29140 (described but unnamed in Puplesis & Robinson 2000); A. paradividua Šimkevičiūtė & Stonis, 2009; A. terrificus Šimkevičiūtė & Stonis, 2009; A. species 015; A. species 016 (described but unnamed in Šimkevičiūtė & Stonis 2009); A. yucatani Remeikis & Stonis, 2013; Glaucolepis argentosa Puplesis & Robinson, 2000. Nepticulids are rather abundant in the tropical forests of the Maya Mountains (Belize) (Puplesis & Robinson 2000). However, none has been described from the “heart” of Yucatán – the lowland forests. The results of the most recent trip to the northeastern areas of the Yucatán Peninsula (2011) suggest the following peculiarities of seasonal activities and distribution of Nepticulidae. There were no adults observed in November, i.e. shortly after the rainy season (May to mid-October). Larvae started mining in very late November or early December. No larval activity of Nepticulidae or Tischeriidae leaf-miners was observed – only old or very old leaf-mines could be found during October and most of November. The diversity of leaf-miners seemed lower than in the pre-montane forests of Belize. The taxonomic composition of Nepticulidae occurring in Yucatán is dominated by the genus Acalyptris Meyrick. Further fieldwork in the peninsula is now needed to estimate the true nepticulid diversity there. Stigmella maya represents a species that is among the smallest nepticulids and the smallest Lepidoptera known in the world. The new species exhibits remarkable morphology and provides new data for further taxonomic studies: in its male genitalia, S. maya mostly resembles a group of undescribed species occurring in the Andes (Patagonia: Argentina) (Stonis & Remeikis, unpublished). The taxonomic composition of Nepticulidae occurring in Yucatan (Fig. 55 A) exhibits unusually strong predomination of genus Acalyptris Meyrick. It differs from that in other regions (Fig. 55), both much smaller (like the Crimea, fig. 55 C) and much larger (like Central Asia, fig. 55 D) or both at close distance (like Florida, fig. 55 B) and very remote (like East Asia or South Africa, fig. 55 E, F).

89 6. CONCLUSIONS

Research Strategy 1 (Neotropical fauna) 1. The examination of collection sample recently collected in areas of Central America (unrepresented in the material used in the most recent review of Opostegidae by other researchers), has resulted in the discovery and description of three new species: Pseudopostega robusta from Costa Rica and P. mexicana and P. latiplana from Mexico. New data increased the number of species of the world‘s Opostegidae fauna to 202.

2. Pseudopostega robusta belongs to the lateriplicata group and it can be recognized by the divided basal part of the gnathos with the thickened lateral folds and stout caudal process; P. mexicana belongs to the spatulata group, but differs in the combination of a large cucular lobe of the valva, narrow vinculum and non thickened gnathos anteriorly; P. latiplana belongs to the saltatrix group, but differs in the large lobes of the uncus and gnathos, which comprises a very large, anteriorly deeply divided lobe, with a stout caudal process and large, broadly rounded basal fold.

3. The Central American fauna of Opostegidae comprises 2 genera and 40 species (i.e., 19.8% of described opostegid species of the world). More than half of these (26 or 65%) are recorded from Central America alone (with the greatest number of species occurring in Costa Rica) and only a few Central American species now are known to possess a broader distribution.

4. Pseudopostega adusta from Costa Rica, P. parakempella from Mexico, P. lobata from Ecuador and P. didyma from the Amazonian Oriente Region of Ecuador represent new geographical records of the Neotropical Opostegidae.

5. The study of substantial unidentified material from South America (Argentina) has has resulted in the acquisition of additional material of Stigmella ovata and discovery of seven new species of Nepticulidae increasing the number of species known from the Neotropics by 1/10. Thus, in total 81 species of Nepticulidae can be recognized now for the Neotropical fauna. However, all newly recognized nepticulid species belong to group, not confirming the phenomenon of Acalyptris predomination in the Patagonian fauna (earlier described in the Neotropical fauna).

6. Nepticulids are rather abundant in the tropical forests of Yucatan. Currently 31 species of Nepticulidae are recorded from Yucatán and adjacent areas, including two new species described in

90 2013: Stigmella maya Remeikis & Stonis and Acalyptris jucatani Remeikis & Stonis. Stigmella maya represents a species that is among the smallest nepticulids and the smallest Lepidoptera known in the world, and exhibits remarkable morphology (mostly resembles a group of undescribed species occurring in the Andes.

7. The results of the most recent trips to the northeastern areas of the Yucatán Peninsula (Mexico) and Guatemala suggest the following peculiarities of seasonal activities and distribution of Nepticulidae: there were no adults observed in November, i.e. shortly after the rainy season (May to mid-October), larvae started mining in very late November or early December.

8. The taxonomic composition of Nepticulidae occurring in Yucatan exhibits unusually strong predomination of genus Acalyptris Meyrick; it differs from that in other regions.

Research Strategy 2 (South European fauna) 9. Results of fieldwork in Karadag Nature Reserve together with investigaton of unidentified material collected by other researchers resulted in the acquisition of additional material and documented a total seven genera and 39 species of Nepticulidae from the Crimea (incl. Stigmella ulmiphaga, S. plagicolella and Acalyptris plantani which are new for the Crimean fauna).

10. The isolated population of Sub-mediterranean Acalyptris platani in Karadag Nature Reserve represents the easternmost record of the species in Europe and supports the concept of the South Crimean Nepticulidae fauna as mostly Sub-mediterranean or Mediterranean. The restricted geographical distribution (mainly Mediterranean) also prevails among the Crimean species of Simplimorpha, Trifurcula and Glaucolepis.

Research Strategy 3 (Himalayan fauna) 11. In total nine species of Opostegidae are currently known as occurring in the Himalaya: Opostegoides epistolaris, O. pelorrhoa, O. index, O. species 1005, P. frigida, Pseudopostega nepalensis, P. zelopa, Opostega chalcophylla, including one new species (Pseudopostega brevicaudata Remeikis & Stonis), occurring in Nepal (Kathmandu valley).

12. Among congeneric species (icluding similar P. frigida), the new Pseudopostega apparently distinguishable by the shape of the gnathos and uncus. Pseudopostega frigida, collected in Himalaya (Rishikesh) represents a new record for the Indian fauna.

91

13. Recent Nepticulidae discoveries in the Himalaya comprise 14 new species belonging to Stigmella, Enteucha, Acalyptris and Ectoedemia, and Fomoria festivitatis (a species new to the Indian Himalaya fauna).

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99 Stonis, J. R., Remeikis, A., Diškus, A., Noreika, R. 2013. New Nepticulidae species (Insecta, Lepidoptera) from the Yucatán Peninsula (SE Mexico). Zootaxa, 3609(2): 223–230. Stonis, J. R., Remeikis, A., Sruoga, V. 2013. An annotated list of the Opostegidae of the Himalaya, with a description of Pseudopostega brevicaudata sp. n. (Lepidoptera: Nepticuloidea). Zootaxa, 3609(2): 182–194. Šimkevičiūtė, A., Stonis, J. R., Sruoga, V. 2010. Male genitalia characters and their diagnostic value in taxonomy of Nepticulidae (Insecta, Lepidoptera). From Biotechnology to Environment Protection. Zielona Gora. Šimkevičiūtė, A., Stonis, J. R., Diškus, A. 2009. Taxonomic checklist of Nepticulidae of Mexico, with the description of three new species from the Pacific Coast (Insecta, Lepidoptera). Acta Zoologica Lituanica, 19 (4) 268–277. Walsingham, Lord. 1907. Microlepidoptera of Tenerife. Proceedings of the General Meetings for Scientific Business of the Zoological Society of London, 1907: 911–1034, pls 51–53. Wilkinson, C. & Newton, P. J. 1981. The micro-lepidopteran genus Ectoedemia Busck (Nepticulidae) in North America. Tijdschrift voor Entomologie, 124 (2): 27–92. Wilkinson, C. & Scoble, M. J. 1979. The Nepticulidae (Lepidoptera) of Canada. Memoirs of the Entomological Society of Canada, 107: 1–129, pls 10. Wilkinson, C. 1979. A taxonomic study of the micro-lepidopteran genera Microcalyptris Braun and Fomoria Beirne occurring in the United States of America (Lepidoptera, Nepticulidae). Tijdschrift voor Entomologie, 122 (4): 59–90. Wilkinson, C. 1981. A supplement to the genus Ectoedemia Busck (Nepticulidae: Lepidoptera) in North America, dealing with some difficult species and also some new ones. Tijdschrift voor Entomologie, 124 (3): 93–110.

100 LITHUANIAN SUMMARY (SANTRAUKA)

Pastaruoju laiku apie biologinės įvairovės tyrimų svarbą labai daug kalbama. Entobiontiniai vabzdžiai yra paplitę įvairiuose Žemės ekosistemose, tačiau duomenų apie šių praktinę ir teorinę reikšmę turinčių vabzdžių taksonominę įvairovę ir geografinį paplitimą vis dar labai maža. Tai parodė ir mūsų atlikta literatūros apžvalga. Iš viso parengtame darbe cituojame apie 60 šaltinių. Visi jie buvo darbo metu prieinami Biosistematikos tyrimų grupės bibliotekoje. Biosistematikos tyrimų grupė disponavo didele moksline kolekcine neidentifikuota medžiaga, gauta iš užsienio mokslo centrų: ZMUC (Danija), USMN (JAV), BMNH (D. Britanija). Medžiagą tyrimams taip pat rinkome ekspedicinių lauko darbų metu Ukrainoje, Karadago gamtiniame rezervate, Indijoje (Himalajuose, Utarakando valstijoje), Jukatane (Meksikoje), Guatemaloje (Centrinėje Amerikoje) ir Kolumbijoje (Pietų Amerikoje). Tyrimų metu buvo panaudotos įprastos taksonominio darbo metodikos, kurios aprašytos bakalauro darbe. Esminis darbo metodologinis elementas buvo taksonų identifikacija, remiantis genitalinių struktūrų morfologija. Tam buvo paruošta apie 270 pastoviųjų mikropreparatų. Pagrindinis mūsų darbo tikslas buvo naujas taksonominis Opostegidae ir Nepticulidae (Nepticuloidea) įvertinimas, remiantis medžiagos, surinktos Neotropiniame regione, Kryme ir Himalajuose moksliniais tyrimais. Darbo tikslui pasiekti buvo numatyti šie uždaviniai: 1 tyrimų kryptis – Neotropinė fauna Meksikoje, Kosta Rikoje ir Ekvadore surinktos Opostegidae medžiagos ištyrimas ir rūšių identifikavimas bei naujų taksonų aprašymas. Naujas Centrinės Amerikos Opostegidae faunos įvertinimas, remiantis padarytu indėliu į baltųjų gaubtagalvių taksonomiją.  Naujų Nepticulidae rūšių paieška ir naujas Neotropinės faunos įvertinimas, remiantis neidentifikuotos kolekcinės medžiagos, surinktos Argentinoje ir asmeninės medžiagos, surinktos Meksikoje bei aplinkiniuose kraštuose, tyrimais. 2 tyrimų kryptis – Pietų Europos fauna Krymo Nepticulidae rūšių įvairovės ir paplitimo apžvalga, remiantis ekspedicinių lauko darbų rezultatais ir neidentifikuotos medžiagos tyrimais. 3 tyrimų kryptis – Himalajų fauna Himalajuose aptinkamų Opostegidae rūšių ištyrimas ir taksonominė apžvalga, su naujų rūšių aprašymu.

101 Himalajų Nepticulidae faunos išaiškinimas, remiantis ekspedicinių lauko darbų Indijos Himalajuose rezultatais. 1 Tyrimų kryptis – Neotropinė fauna Ištyrus kolekcinę medžiagą, kitų tyrėjų surinktą Kosta Rikoje ir Meksikoje, identifikuotos 3 naujos baltųjų gaubtagalvių (Opostegidae) rūšys. Remiantis patino genitalinių struktūrų tyrimais Pseodopostega robusta, aptiktai Kosta Rikos medžiagoje išaiškinta itin stipri analinės plokštelės (gnathos) šoninė sklerotizacija ir labai išvystyta kaudalinė išauga. Remiantis šiais požymiais, kuriuos vertiname kaip naujadarus, t. y. apomorfijus, teigiame, kad ši rūšis yra iki šiol nežinoma mokslui. Remiantis patino genitalinių struktūrų tyrimais Pseodopostega mexicana, aptiktai Meksikos medžiagoje, nustatyta didelė porinės plokštelės kaudalinė skiautė, siaura pamatinė plokštelė ir stipri priekinės analinės plokštelės (gnathos) sklerotizacija. Tiriant naują mokslui rūšį Pseudopostaga latiplana, aptikta neįprasta analinės ir galinės plokštelių forma. Tai pateikė naujų duomenų apie Opostegidae genitalinių struktūrų įvairovę ir evoliuciją ir sudarė prielaidas aprašymui kaip naujos ir išskirtinės rūšies. Ši rūšis, kaip ir ankščiau paminėta P. mexicana, aptikta Meksikos medžiagoje. Taigi, šiuo metu Centrinėje Amerikoje jau išaiškinta 40 Opostegidae rūšių, iš kurių daugiausia taksonų paplitę Kosta Rikoje (26 rūšys). Centrinės Amerikos Opostegidae geografinio paplitimo analizė (kurios rezultatus pateikiame pirmoje bakalauro lentelėje) parodė, kad daugeliui šių rūšių yra būdingi riboti arealai ir tik nedaugelis jų yra paplitę už Centrinės Amerikos ribų. Ištyrus papildomą Opostegidae medžiagą, surinktą Ekvadore, patikslintas kai kurių rūšių geografinis paplitimas. Pseudopostega adusta: ši rūšis ankščiau buvo žinoma tik iš Kubos, Dominikos, Belizo ir Ekvadoro, tačiau atlikti medžiagos tyrimai pateikė neginčijamų įrodymų apie šios rūšies paplitimą Kosta Rikoje. Pseudopostega didyma: ši rūšis iki šiol buvo žinoma tik iš vienintelės Andų kalnų vakarinių šlaitų vietovės Ekvadore. Kaip žinoma, Andų kalnų grandinė yra svarbus gamtinis barjeras rūšių paplitimui, todėl ištyrus medžiagą ir išaiškinus naują P. didyma paplitimą Amazonės baseine, kilo daug diskusijų apie rūšių paplitimo ypatybes Neotropiniame regione. Pseudopostega lobata: ši rūšis ankščiau buvo žinoma tik iš trijų Centrinės Amerikos šalių ir Argentinos. Tačiau atlikti medžiagos tyrimai pateikė neginčijamų įrodymų apie šios rūšies paplitimą Ekvadore. Nauji duomenys padėjo tiksliau įvertinti Ekvadoro fauną, o taip pat keičia viso Neotropinio regiono Opostegidae faunos sampratą. Tyrimų metu buvo identifikuota didelė Nepticulidae mokslinė medžiaga iš Argentinos, kurią tyrimam perdavė užsienio institucijos. Tarp išaiškintų rūšių nustatyta Stigmella ovata (iki šiol

102 žinoma tik iš dviejų egzempliorių) ir dar 7 kitos mokslui naujos rūšys. Reikšmingiausia ne tik tai, kad visos šios naujos rūšys pasižymi išskirtiniais apomorfiniais požymiais, leidžiančiais teigti apie intensyvius autochtoninės evoliucijos procesus Patogonijos kalnuose, bet ir tai, kad visos rūšys priklauso tik dviems rūšių grupėms – Stigmella salicis (ir nepublikuotai rūšių grupei) Tai keičia iki šiol vyravusią sampratą apie Acalyptris genties dominavimą Neotropinėje Nepticulidae faunoje. Remiantis naujais duomenimis Acalyptris tyrinėtoje faunoje sudaro 31 proc. (o ne 34 proc. kaip buvo žinoma ankščiau); didžiausią dalį (42 proc.) sudaro Stigmella genties rūšys. Taip pat nauji atradimai leidžia teigti, kad šiuo metu Argentinoje aptinkama 12 Nepticulidae rūšių, nors didžiausias rūšių skaičius žinomas Belize ir Ekvadore. Nustatyta didelė Nepticulidae įvairovė Jukatano ir kaimyninių kraštų atogrąžų miškuose. Iš viso šiuo metu žinoma 31 mažųjų gaubtagalvių rūšis, tarp kurių dvi (Stigmella maya Remeikis & Stonis ir Acalyptris jucatani Remeikis & Stonis) buvo aprašytos 2013 m. Stigmella maya yra mažiausia šiuo metu žinoma rūšis tarp visų pasaulio Lepidoptera būrio atstovų ir pasižymi išskirtine morfologija. Lauko darbų rezultatai pirmą kartą leidžia teigti apie Nepticulidae gyvybinius ciklus Centrinės Amerikose atogrąžų miškų buveinėse; vikšrų minavimas prasideda lapkričio pabaigoje – gruodžio mėn., tačiau neregistruotas per lietingąjį sezoną. Jukatano Nepticulidae faunos sudėčiai būdingas neįprastai stiprus Acalyptris Meyrick genties dominavimas, kurio neaptinkame kitose, nedidėlėse ir kaimyninėse ar didelėse ir tolimose faunose. 2 tyrimų kryptis – Pietų Europos fauna Krymo Nepticulidae faunos tyrimai buvo atliekami dviem būdais: vykdėme lauko darbus Kryme, Karadago gamtos rezervate bei ištyrėme kitų tyrėjų surinktą kolekcinę medžiagą. Atlikus šiuos darbus, pakartotinai nustatyta 13 jau ankščiau Krymo faunoje žinomų Nepticulidae rūšių. Papildomai išaiškintos 4 naujos gaubtagalvių rūšys, iki šiol nežinomos Krymo faunoje. Remiantis gautais rezultatais išaiškintos 39 gaubtagalvių rūšys, kurių daugumai (apie 72 proc.) yra būdingas platus europinis paplitimas. Ribotu paplitimu Mediteraniniame regione pasižymi Simplimorpha, Acalyptris, Trifurcula ir Glaucolepis rūšys. Ypatingą reikšmę turi plataninio gaubtagalvio (Acalyptris platani) aptikimas Kryme. Ištyrę surinktą kolekcinę medžiagą ir palyginę mūsų duomenis su kitų autorių skelbtais duomenimis (van Nieukerken, 2007 ir kt.) apie A. platani Viduržemio jūros kraštų faunoje, išaiškinome šiuos morfologinius skirtumus: 1) spalviniai Karadage aptiktų plataninio gaubtagalvio priekinių sparnų raštai yra šviesesni; 2) užpakalinių sparnų priekinio krašto androkoniniai žvyneliai yra pakitę (išplatėję); 3) tiek patinų, tiek patelių genitalinėse struktūrose pastebėjome nežymių skleritų formos

103 skirtumų (lyginant šias struktūras su individais, aptiktais kituose kraštuose). Taigi A. platani aptikimas Kryme leidžia teigti ne tik apie toliausiai į rytus nutolusią plataninio gaubtagalvio geografiškai izoliuotą populiaciją, bet ir apie šios populiacijos individų morfologinius skirtumus. Šie skirtumai nėra pakankamai dideli, kad būtų galima teigti apie naują A. platani porūšį Krymo faunoje; vis dėlto, A. platani išaiškinimas Kryme gali būti įvertintas kaip gana reikšmingas mokslo faktas, nes Acalyptris gentis ik šiol nebuvo žinoma Rytų Europoje. 3 tyrimų kryptis – Himalajų fauna Remiantis atliktų ekspedicinių lauko darbų rezultatais, BMNH tipinės medžiagos tyrimais ir sudaryta Opostegidae taksonominė apžvalga, nustatyta, kad vienos iš 9 analizuojamų rūšių kaudalinės plokštelės (uncus) trumpos ataugos ir išvystyta šoninė analinės plokštelės (gnathos) sklerotizacija yra pakankamas pagrindas, kad traktuoti šią Pseudopostega brevicaudata rūšį kaip naują mokslui. Kita Himalajuose aptikta Opostegidae rūšis (P. frigida) yra nauja Indijos faunai. Tarp Himalajuose aptiktų 15 Nepticulidae rūšių tik viena (Fomoria festivitatis) buvo žinoma mokslui (visos kitos aptiktos rūšys yra naujos mokslui), tačiau surinktos medžiagos tyrimai pirmą kartą leidžia teigti apie F. festivitatis paplitimą Indijos Himalajuose.

Išvados 1 tyrimų kryptis – Neotropinė fauna 1. Tyrimų metu identifikuotos 3 naujos mokslui baltųjų gaubtagalvių (Opostegidae) rūšys, paplitusios Kosta Rikoje arba Meksikoje; šioms naujoms rūšims būdinga: itin stipri analinės plokštelės (gnathos) šoninė sklerotizacija ir labai išvystyta kaudalinė išauga (Pseodopostega robusta); itin didelė porinės plokštelės kaudalinė skiautė ir siaura pamatinė plokštelė bei stipri priekinė analinės plokštelės (gnathos) sklerotizacija (Pseodopostega mexicana); neįprasta analinės ir galinės plokštelių forma (Pseudopostaga latiplana).

2. Remiantis tyrimų duomenimis Centrinėje Amerikoje išaiškinta 40 Opostegidae rūšių, iš kurių 26 aptinkamos Kosta Rikoje. Geografinio paplitimo analizė parodė, kad daugeliui šių rūšių yra būdingi riboti arealai ir tik nedaugelis yra paplitę už Centrinės Amerikos ribų.

3. Pseudopostega adusta ankščiau buvo žinoma tik iš Kubos, Dominykos, Belizo ir Ekvadoro, tačiau atlikti tyrimai pateikė įrodymų apie šios rūšies paplitimą Kosta Rikoje; P. didyma iki šiol buvo žinoma tik iš Andų vakarinių šlaitų, tačiau tyrimai parodė, kad ši rūšis (nežiūrint Andų kalnų barjero) paplitusi ir Amazonės baseine; P. lobata ankščiau buvo žinoma tik iš trijų Centrinės

104 Amerikos šalių ir Argentinos, tačiau atlikti medžiagos tyrimai pateikė neginčijamų įrodymų apie šios rūšies paplitimą Ekvadore.

4. Nepticulidae medžiagos identifikacija padėjo išaiškinti 7 mokslui naujos rūšys. Tačiau visos šios naujos rūšys ne tik pasižymi išskirtiniais apomorfiniais požymiais, leidžiančiais teigti apie intensyvius autochtoninės evoliucijos procesus Patogonijos kalnuose, bet ir tai, kad jos visos priklauso tik dviems rūšių grupėms – Stigmella salicis ir vienai iki šiol neaprašytai grupei.

5. Argentinos Nepticulidae tyrimų rezultatai keičia iki šiol vyravusią sampratą apie Acalyptris genties dominavimą Neotropinėje Nepticulidae faunoje. Remiantis naujais duomenimis Acalyptris tyrinėtoje faunoje sudaro 31 proc. (o ne 34 proc. kaip buvo žinoma ankščiau); didžiausią dalį, 42 proc., sudaro Stigmella genties rūšys.

6. Nustatyta didelė Nepticulidae įvairovė Jukatano ir kaimyninių kraštų atogrąžų miškuose. Iš viso šiuo metu žinoma 31 mažųjų gaubtagalvių rūšis, tarp kurių dvi (Stigmella maya Remeikis & Stonis ir Acalyptris jucatani Remeikis & Stonis) buvo aprašytos 2013 m. Stigmella maya yra mažiausia šiuo metu žinoma rūšis tarp visų pasaulio Lepidoptera būrio atstovų ir pasižymi išskirtine morfologija.

7. Lauko darbų rezultatai pirmą kartą leidžia teigti apie Nepticulidae gyvybinius ciklus Centrinės Amerikose atogrąžų miškų buveinėse; vikšrų minavimas prasideda lapkričio pabaigoje – gruodžio mėn., tačiau neregistruotas per lietingąjį sezoną.

8. Jukatano Nepticulidae faunos sudėčiai būdingas neįprastai stiprus Acalyptris Meyrick genties dominavimas, kurio neaptinkame kitose, nedidėlėse ir kaimyninėse ar didelėse ir tolimose faunose.

2 tyrimų kryptis – Pietų Europos fauna 9. Atlikus lauko darbus bei identifikavus iki šiol netyrinėtą kolekcinę medžiagą, nustatytos 39 rūšys.

10 . Daugumai Kryme aptiktų rūšių (apie 72 proc.) yra būdingas platus europinis paplitimas. Ribotu paplitimu Mediteraniniame regione pasižymi tik Simplimorpha, Acalyptris, Trifurcula ir Glaucolepis rūšys.

105 11. Acalyptris platani aptikimas Kryme leidžia teigti ne tik apie toliausiai į rytus nutolusią šios sub- mediteraninės rūšies geografiškai izoliuotą populiaciją, bet ir apie šios populiacijos individų morfologinius skirtumus. Tačiau šie skirtumai nėra pakankamai dideli, kad būtų galima teigti apie naują A. platani porūšį.

3 tyrimų kryptis – Himalajų fauna 12. Nustatyta, kad vienos iš Himalajų faunoje analizuojamų Opostegidae rūšių trumpos kaudalinės plokštelės (uncus) ataugos ir išvystyta šoninė analinės plokštelės (gnathos) sklerotizacija yra pakankamas pagrindas, kad traktuoti šią Pseudopostega brevicaudata rūšį kaip naują mokslui; kita, Himalajuose aptikta Opostegidae rūšis (P. frigida) yra nauja Indijos faunai.

13. Tarp Himalajuose aptiktų 15 Nepticulidae rūšių, tik viena (Fomoria festivitatis) buvo žinoma (visos kitos aptiktos rūšys yra naujos mokslui), tačiau surinktos medžiagos tyrimai pirmą kartą leidžia teigti apie F. festivitatis paplitimą Indijos Himalajuose.

Šių tyrimų medžiaga buvo pristatyta aštuoniuose moksliniuose forumuose (įskaitant tarptautinę konferenciją užsienyje). Taip pat darbo metu paskelbiau (kartu su bendraautoriais) 31 straipsnį: 8 mokslinius ir 23 mokslo sklaidos.

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