Asta Navickaitė TAXONOMY, CHOROLOGICAL

LITHUANIAN UNIVERSITY OF EDUCATIONAL SCIENCES

Asta Navickaitė

TAXONOMY, CHOROLOGICAL COMPOSITION AND TROPHIC RELATIONSHIPS OF THE EURONEMORAL FAUNA OF NEPTICULIDAE (INSECTA, LEPIDOPTERA)

Summary of Doctoral Dissertation Biomedical Sciences, Biology (01 B): Entomology and Plant Parasitology, B–250

Vilnius, 2014

1 The work was carried out in 2009–2014 at Lithuanian University of Educational Sciences

Research supervisor: Assoc. Prof. Dr. Arūnas Diškus (Lithuanian University of Educational Sciences, Zool- ogy, 05B and Biology, 01B: Entomology and Plant Parasitology, B–250)

Research advisor: Prof. Habil. Dr. Jonas Rimantas Stonis (Lithuanian University of Educational Scienc- es, Zoology, 05B and Biology, 01B: Entomology and Plant Parasitology, B–250)

Chairman: Prof. Habil. Dr. Rimantas Rakauskas, Vilnius University (Biology, 01B: Entomology and Plant Parasitology, B–250)

Members: Dr. David C. Lees, Cambridge University, UK (Biology, 01B: Entomology and Plant Parasitology, B–250) Dr. M. Alma Solis, the Smithsonian Institution, USA (Biology, 01B: Entomology and Plant Parasitology, B–250) Assoc. Prof. Dr. Jurga Turčinavičienė, Vilnius University (Biology, 01B: Entomology and Plant Parasitology, B–250) Assoc. Prof. Dr. Jana Radzijevskaja, Vytautas Magnus University (Biology, 01B)

Opponents: Prof. Dr. Sigitas Podėnas, Vilnius University (Biology, 01B: Entomology and Plant Parasitology, B–250) Dr. Jolanta Rimšaitė, the Nature Research Centre (Biology, 01B: Entomology and Plant Parasitology, B–250)

The defence of the dissertation will be held at a public meeting of the Biology (En- tomology and Plant Parasitology) Coucil at Lithuanian University of Educational

Sciences on September 26, 2014, at 11 a.m. (Auditorium A11). Address: 39 Studentų St., Vilnius, LT–08106, Lithuania.

The summary of the doctoral dissertation was send on August ___, 2014. The dissertation is available at the Library of the Lithuanian University of Educational Sciences.

Asta Navickaitė

EURONEMORALINĖS FAUNOS MAŽŲJŲ GAUBTAGALVIŲ (INSECTA, LEPIDOPTERA, NEPTICULIDAE) TAKSONOMINĖ IR CHOROLOGINĖ ANALIZĖ BEI TROFINIAI RYŠIAI

Daktaro disertacijos santrauka Biomedicinos mokslai, Biologija (01 B): Entomologija ir augalų parazitologija, B–250

Vilnius, 2014 Disertacija parengta 2009–2014 metais Lietuvos Edukologijos universitete.

Mokslinis vadovas: Doc. dr. Arūnas Diškus (Lietuvos edukologijos universitetas, Zoologija, 05B, ir Biolo- gija, 01B: entomologija ir augalų parazitologija, B–250)

Mokslinis konsultantas: Prof. habil. dr. Jonas Rimantas Stonis (Lietuvos edukologijos universitetas, Zoologija, 05B, ir Biologija, 01B: entomologija ir augalų parazitologija, B–250)

Pirmininkas: Prof. habil. dr. Rimantas Rakauskas, Vilniaus Universitetas (Biologija, 01B: entomologija ir augalų parazitologija, B–250)

Nariai: Dr. David C. Lees, Kembridžo universitetas, Jungtinė Karalystė (Biologija, 01B: entomologija ir augalų parazitologija, B–250) Dr. M. Alma Solis, Smitsono centras/ Smithsonian Institution, JAV (Biologija, 01B: entomologija ir augalų parazitologija, B–250) Doc. dr. Jurga Turčinavičienė, Vilniaus Universitetas (Biologija, 01B: entomologija ir augalų parazitologija, B–250) Doc. dr. Jana Radzijevskaja, Vytauto Didžiojo Universitetas (Biologija, 01B)

Oponentai: Prof. dr. Sigitas Podėnas, Vilniaus Universitetas (Biologija, 01B: entomologija ir augalų parazitologija, B–250) Dr. Jolanta Rimšaitė, Gamtos tyrimų centras (Biologija, 01B: entomologija ir augalų parazitologija, B–250)

Disertacija bus ginama viešame Biologijos (entomologijos ir augalų parazitologijos) tarybos posėdyje, kuris vyks 2014 m. rugsėjo 26 d., 11 val., Lietuvos Edukologijos Uni- versitete, A11 auditorijoje. Adresas: Studentų g. 39, Vilnius, LT–08106, Lietuva Disertacijos santrauka išsiuntinėta 2014 m. rugpjūčio ___d. Su disertacija galima susipažinti Lietuvos edukologijos universiteto bibliotekoje.

THE PROBLEM AND RELEVANCE OF THE RESEARCH

Owing to active destruction of landscapes as well as frequent frag- mentation of habitats, the global biodiversity is undergoing rapid decline. The increasing concern by the world‘s scientific community about the biodiversity crisis stimulated researchers to more actively inventorize the Earth‘s main biomes (Stonis, 2010; Navickaitė et al., 2011b). The object of our research is one of the most primitive (and from this point of view, one of the most interesting) taxonomic groups of the Lepi- doptera order (Nepticuloidea: Nepticulidae) comprising 850 species and 16 genera: Pectinivalva, Roscidotoga, Simplimorpha, Enteucha, Mononeura, Areticulata, Stigmella, Bohemannia, Ectoedemia, Fomoria, Acalyptris, Parafomoria, Trifurcula, Glaucolepis, Etainia and Varius. The majority of these genera are widely distributed throughout many continents; they include the world‘s smallest Microlepidoptera, which are characterized not only by their archaic morphology but also, as plant-miners, greatly specialized (Stonis, 2010; Diškus, Stonis, 2012). Because of stenophagy, sedentary lifestyle and a high rate of ende- mism, these small insects occur in almost all land biomes (from tundra or deserts to tropical rainforests) and could serve as a perfect tool for characterizing the richness, origin and faunal relationships of biomes under investigation (Navickaitė et al., 2011b). However, research data concerning these insects, which are both of practical and theoretical importance, are insufficient. Prior to the investigation carried out by the author of the dissertation or her colleagues, the fauna of many regions had been unknown and undescribed and the data about the species diversity of Nepticulidae of the Crimea and Lithuania had been far from exhaustive. The deficiency of investigation data, economical (practical) and theoretical (scientific) significance of the family Nepticulidae and increasing concern by the scientific community about the biodiversity crisis stimulated the preparation of this revision of Nepticulidae of the Euro-Nemoral zone; the importance of such studies was also emphasized by the Convention on Biological Diversity of Rio de Janeiro.

5 GOAL AND OBJECTIVES OF THE RESEARCH

Goal of the dissertation: to present the first taxonomic, chorological and trophic review of the Nepticulidae of the Crimea and Lithuania within the context of the global and European Nepticulidae fauna.

The objectives of the research:

General assessment of the Nepticulidae fauna 1. To perform general assessment of taxonomic composition and distribution of the currently known Nepticulidae fauna within the Earth’s biogeographical regions. 2. To estimate data on trophic relationships within the world’s Nep- ticulidae fauna with re-assessment of trophic relationships among the Stigmella hemargyrella species. 3. To evaluate taxonomic and diagnostic characters of species groups of Stigmella associated with Quercus as a host-plant by presenting dichotomous identification systems for the species groups. 4. To perform cladistic analysis of species groups of Stigmella trophically associated with Quercus.

The Crimean Nepticulidae within the context of the European Fauna 5. To estimate the taxonomic composition and trophic relationships of the European Nepticulidae. 6. To perform chorological analysis of the European fauna. 7. To identify Nepticulidae species of the Crimean fauna on the basis of the performed fieldwork and examination of scientific material collected by other authors. 8. To identify geographical distribution and chorological groups of the Crimean fauna on the basis of the performed chorological analysis of Nepticulidae of the Crimea. 9. To evaluate occurrence of species registered within the Crimean fauna and identify the common and rare species. 10. To analyse features of mining of common species of the Crimean Nepticulidae and their belonging to chorological groups.

6 11. To analyse features of mining of rare species of the Crimean Nep- ticulidae and their belonging to chorological groups. 12. To perform chorological analysis of the Nepticulidae fauna of the Crimea and define periods of larval activity.

Taxonomic, chorological and trophic analysis of the Lithuanian Nepticulidae Fauna 13. To evaluate the taxonomic composition of the Lithuanian Neptic- ulidae and present a checklist. 14. To perform chorological analysis of the Lithuanian Nepticulidae and identify chorological groups. 15. To compare chorological data on the Nepticulidae fauna of various regions of Lithuania. 16. To define trophic features of the Lithuanian Nepticulidae and identify the predominant life cycles. 17. To perform analysis of trophic relationships and identify families of host-plants and genera of the Lithuanian Nepticulidae. 18. To estimate abundance of mining of the Lithuanian Nepticulidae in line with the methods for evaluating the distribution of species and mining. 19. To estimate occurrence of species registered within the Lithuani- an fauna by identifying common and rare species and linking the occurrence data with the results of the chorological analysis.

NOVELTY OF THE RESEARCH

The first taxonomic catalogue of the Western Palaearctic Neptic- ulidae fauna has been prepared, the taxonomic position of all the 850 species of the world’s Nepticulidae fauna has been checked and 15 new taxonomic combinations have been proposed. We (together with co-authors) have prepared an original taxonomic analysis of the Quercus-feeding Nepticulidae based on newly discovered data of morphology (incl. characters of genitalia). We have done comprehensive diagnostic and trophic analysis of the Stigmella species groups associated with Quercus. We did the first chorological assessment of the Lithuanian Nepticuli- dae fauna, supplemented the checklist of Lithuania’s fauna with new data

7 and performed the first comprehensive analysis of trophic relationships among species known from Lithuania. By carrying out frequent research fieldwork on Nepticulidae in Lithuania, we identified the Nepticulidae fauna of the Curonian Spit for the first time; the distinctive feature of this region’s fauna is a greater number of species of Euro-Mediterranen distribution than in the continental Lithuania. We for the first time dissected and studied hitherto unidentified Nepticulidae material collected in the Crimea and deposited at the Zoo- logical Institute of the Russian Academy of Sciences. We also performed the first taxonomic and chorological analysis of the Crimean fauna. During the fieldwork in the Karadag Reserve and other localities in the Crimea, we registered 64 Nepticulidae species, among which 26 (41%) were recorded for the first time for the Crimean fauna. We did the first analysis of the Crimean species, prepared new descriptions matching the requirements and the first original catalogues of leaf-mines and genitalia. When investigating Nepticulidae from regions related to the Eu- ro-Nemoral fauna, we discovered 8 new species of Nepticulidae (Navick- aitė et al., in prep.) of the Stigmella, Enteucha, Ectoedemia and Acalyptris. We (together with co-authors) described and published a new species of the Stigmella hemargyrella species group – Stigmella acuta.

THEORETICAL AND PRACTICAL SIGNIFICANCE

The International Rio de Janeiro Convention, ratified by the Sei- mas of the Republic of Lithuania, encourages pursuing and develop- ing various biodiversity research. Because of the recent technology developments and the overall intention to describe the extent of the biodiversity research, considerable effort was undertaken to investi- gate endobiontic insects in recent years. One of the most prominent characteristics of Nepticulidae, their mining within the green tissue of plants, presents an issue which is interesting and important for ecology (Stonis, 2010, 2011). The Nepticulidae family belonging to the Nepticuloidea superfam- ily analysed in the dissertation is widely distributed throughout various terrestrial ecosystems; it has been, however, poorly investigated from the theoretical point of view. The climate change, intensive migration

8 of the population and commercial projects pursuing economic benefits (in particular involving exotic territories) provide conditions for an incidental introduction of pest fauna (Diškus, Stonis, 2012). Because of the above-mentioned lifestyle within assimilating tissues of leaves, buds, stems, young bark or fruits of plants, a number of organisms that we investigated could be considered as pests or potential pests; for that reason they are important from the economic point of view (Puplesis, 1994, Kuznetzov, Puplesis, 1994). It is considered that oligophagous endobiontic insects are capable to rather easily adapt in other territories that are new to them and establish trophic relationships with alien host-plants (Diškus, Stonis, 2012). The investigation of life-cycles, trophic relationships or other biological characteristics of potential pests, as well as continuous cooperation with professionals of nature protection, could enable the preparation of effective measures preventing alien pests from infiltrating our fauna and application of a quarantine policy.

STATEMENTS TO BE DEFENDED

General assessment of the Nepticulidae fauna 1. There has been a considerable change in taxonomic composition of the Nepticulidae fauna because of numerous descriptions of new species and synonymization of some of the names; however, the species are not equally distributed among various biogeographical regions of the Earth. 2. Though host-plants of the majority of the Nepticulidae species have already been identified, the description of a new species Stigmella( acuta) has altered the conception of the Stigmella hemargyrella species group. 3. Within the world’s fauna there are six species groups of the Stigmella trophically associated with Quercus: S. caesurifasciella, S. saginella, S. quercipulchella, S. ruficapitella, S. castanopsiella and S. hemargyrella. 4. TheStigmella quercipulchella, S. hemargyrella, S. ruficapitella and S. castanopsiella groups are closely related, whereas S. saginella, S. caesurifasciella and S. cornuta are three phylogenetically independent clades; it could be stated that the species groups of Stigmella trophi-

9 cally associated with Quercus and occurring in the Eastern and West- ern Hemisphere (e.g. in America) do not share the same origin and represent different evolutionary tendencies.

The Crimean Nepticulidae within the context of the European fauna 5. There has been a considerable change in taxonomic composition of the European Nepticulidae fauna and conception of trophic relationships because of numerous descriptions of new species and synonymization of some of the names. At present the prevailing species within the European fauna are those which are trophically related to woody plants, mostly belonging to Rosaceae and Fagaceae and in a lesser degree, to Betulaceae, Salicaceae and other families. 6. The European Nepticulidae fauna is not homogeneous: the species from this region belong to 10 chorological groups, the largest of which are the Mediterranean and Euro-Submediteranean. 7. At present the Crimean fauna includes 64 species, 40% of which were identified for the first time during the dissertation investigation. 8. The Crimean Nepticulidae fauna is not homogeneous: the species from this region belong to 8 chorological groups; the majority of them (72%) belong to those chorological groups which are related to the Mediterranean ecoregion, whereas extremely vast Euro-Siberian or Trans-Palaearctic ranges are characteristic only to 1–5% of species occurring in the Crimea. 9. The majority of the Nepticulidae species of the Crimean fauna are characterized by wide distribution and abundant or very abundant mining. 10. The majority of the common Crimean species are leaf-miners of Quercus, Sorbus, Crataegus and Prunus plants and they mainly belong to the Euro-Submediterranean chorological group. 11. The majority of the rare species also are leaf-miners of Quercus, as well as the Lotus, Alnus and Acer, plants and mainly belong to the Euro-Mediterranean, Euro-Submediterranean, Euro-Nemoral or Sub-Mediterranean chorological groups. 12. The major part of the Crimean Nepticulidae fauna is associated with Rosaceae and Fagaceae plants; about one tenth of the species identified in the Crimea are leaf-miners of cultivated plants (including

10 Stigmella irregularis, the host-plant of which (Pyrus elaeagnifolia, was identified for the first time). 13. At present there are 76 Nepticulidae species within Lithuania’s fauna, which belong to 6 genera. 14. The Lithuanian Nepticulidae fauna is not homogeneous: it belongs to 8 chorological groups, the most abundant of which are those of moderately limited European distribution – the Euro-Submediterranean and Euro-Nemoral. 15. Throughout all the regions of Lithuania, the species of the Eu- ro-Submediterranean distribution constitute the major part of the Nepticulidae fauna except the Sea-Coast Region, where they are less numerous. Among Lithuania’s regions, the Curonian Spit is the most distinctive as it is characterized by species of the Sub-Baltic group which do not occur in other regions of Lithuania. 16. There are many oligophagous species among the Lithuanian Nep- ticulidae (37%), though most of them are monophagous (63%). The majority of Lithuania’s Nepticulidae are bivoltine species and the remaining are univoltine; only one species of Lithuania‘s fauna is polivoltine. 17. Lithuania’s Nepticulidae mine the plants belonging to 15 families and 35 genera (most of Lithuania’s Nepticulidae are trophically associated with Rosaceae, Salicaceae, Fagaceae and Betulaceae plants; the major part of them mine leaves of woody plants). 18. The predominant type of mining of Nepticulidae within the Lithua- nian fauna is abundant. Most species are characterized by wide distribution and abundant mining. 19. About 40% of the Lithuanian Nepticulidae species belong to the category of rare (rare and very rare) and 23%, to the common species. Nepticulidae of both common and rare species are mostly characterized by the Euro-Nemoral and Euro-Submediterranean ranges (there are no significant chorological differences between the Lithuania’s common and rare species).

11 APPROBATION

The key results that are the basis of the dissertation were presented for discussion and were approbated at the following conferences, semi- nars and meetings of research teams: • The National Scientific Conference “Biological Diversity of Lithuania: Status, Structure, Protection”: the co-authored report “The Fauna and Trophic Relationships of Nepticulidae (Lepidoptera) of Vilkaviškis District” (Vilnius, 2008). • The Scientific Seminar “Developments of Ecosystems and Molecular Investigation of the Phylogeography of Populations in the Baltic Re- gion”: the co-authored report “New Phylogenetic Hypotheses about Plant-Mining Primitive Lepidoptera Based on Molecular Data (Lep- idoptera: Nepticuloidea, Tischeriodea, Gelechiodea)” (Vilnius, Lithu- anian Academy of Sciences, 2009). • The International Seminar “Urban Ecology – Introduction: How to Work with Guide No. 8” (Sønderborg, Denmark, 2010). • The General Meeting of the Lithuanian Entomological Society: co-authored report “In India During the SW Monsoon: the Fieldwork of the Biosystematics Research Group in Uttarakhand, the Himala- yas, 2010” (Vilnius University, 2010). • The 5th International Conference of Naturalists “From Biotechnolo- gy to Environment Protection – Interdisciplinary Meeting of Natu- ralists”: the co-authored “First Survey of Nepticulidae (Insecta: Lep- idoptera) of the Curonian Spit (Baltic Coast of Lithuania)”. (Zielona Góra, Poland, 2010). • The National Conference “The Art of Living 2011”: the report “The Himalayas: Secrets of Nature and Dangers of Research” (Panevėžys, 2011). • The Scientific Conference of the Faculty of Natural Sciences of the Lithuanian University of Educational Sciences: the co-authored report “Will the Record of 500 New Species be Achieved Soon?” (Vil- nius, 2011). • Meetings of the Biology Department of the Lithuanian University of Educational Sciences (Vilnius, 2009, 2010, 2011, 2012, 2013, 2014).

12 • The First Baltic International Conference on Field Entomology and Faunistics: the co-authored report “What is New and Most Interest- ing About the Nepticulidae of Lithuania and the Crimea” (Vilnius, 2014). • The First Baltic International Conference on Field Entomology and Faunistics: the co-authored report “Field Methods of Nepticulidae Research: the Studies Leaf-Mining Lepidoptera Could Be Exciting” (Vilnius, 2014).

The approbation of the results was also done during the project “New Faunas: Taxonomic Analysis and Endobiontic Lepidoptera Species New to Science (MIP-049/2011)” financed from the Research Foundation of the Research Council of Lithuania (2010–2011) and the project “Taxo- nomic Identification of the Lithuanian Endobiontic Fauna” financed by the Research Council of Lithuania and carried out within the activities of the Recommended Scientific Research and Experimental Development (MTEP) (2012).

PUBLICATION OF THE RESEARCH RESULTS

The results of the dissertation research have been published in 19 papers that appeared in journals and other publications. Among them, 5 articles were published in the following periodical scientific journals: 2 in Acta Zoologica Lituanica, 1 in Ecology and Zoology and 2 in Zootaxa (impact factor). The author also prepared 7 co-authored chapters of a monograph (see Diškus, Stonis, 2012: The Nepticulidae (Lepidoptera): taxonomy, chorological composition and trophic relationships). In total, the author (together with co-authors) published 8 papers in English and 11 papers in Lithuanian.

STRUCTURE AND SCOPE OF THE DISSERTATION

The dissertation is in Lithuanian and its abstract is in English. The dissertation consists of the following chapters: Introduction, Literature Overview, Methods and Materials, Investigation Results, Conclusions,

13 References, and 3 Annexes: Catalogue of the Western Palaearctic Nep- ticulidae Fauna, Catalogue of genitalia of the Crimean Nepticulidae, and Catalogue of leaf-mines of the Crimean Nepticulidae. The list of references includes 282 entries. The dissertation consists of 333 pages; it includes 188 figures (56 within the text of the dissertation and 132 within Annex- es) and 10 tables.

ACKNOWLEDGEMENTS

I am sincerely grateful to the dissertation supervisors Assoc. Prof. Dr. A. Diškus and Prof. Habil. Dr. J. R. Stonis for their patience, permanent help and valuable advice. I would like to express gratitude to Prof. Dr. V. Sruoga, PhD fellow A Rocienė (LEU), PhD fellow A. Remeikis (NRC, Vilnius), Dr. P. Ivinskis (NRC, Vilnius), K. R. Tuck (BMNH, London), O. Karsholt (ZMUC, Copenhagen), Dr. E. J. van Nieukerken (NMM, Leiden), Dr. D. R. Davis (USNM, Washington), DSc. Sergey Yu. Sinev (ZIN, St. Petersburg). Dr. Yu. I. Budashkin (KRUAS, Karadag) for their contribution, consultations and recommendations. I am thankful to Dr. B. Tamutė (Vytautas Magnus University and the Tadas Iva- nauskas Zoological Museum, Kaunas) and the researchers E. Anisimov, V. Juchnevič, S. Brusokaitė, K. Lensbergaitė, G. Varačinskas, J. Čeputytė, E. Poškaitė, V. Zeleniūtė, D. Aržuolaitytė, V. Dobrynina, J. Prakapa- vičiūtė, S. Šutaitė, R. Mankutė, A. Bobnis, J. Vertelytė and M. Adomaitis for their assistance in investigating the Nepticulidae fauna of Lithuania (or neighbouring countries). I wish to thank D. Mikelaitis (the Fauna Use Control Division at the Environment Protection Agency, Vilnius) for his assistance in getting permits for the exchange of study material with the Russian Academy of Sciences and the Ukrainian Academy of Sciences. I am thankful to the Director of the Regional Park of Vištytis Nerijus Paškauskas for the permission to carry out Nepticulidae research within the territory of the Park. I am grateful to the ecologist senior specialist at the Regional Park of Vištytis Vaidas Leonavičius (Vištytis) and the landscape manager senior specialist Andrius Vaičaitis (Kybartai) for their benevolent cooperation in carrying out the research and preparing the information. I am greatly indebted to Rita and Kęstutis Navickas (Kaunas), Ona Žuliuvienė (Vilkaviškis) and Svajūnas Krivickas for their

14 support and understanding. I am also thankful to Andrius Navickas (Kaunas) for their help in preparing tutorial aids. I would like to express sincere thanks to the Director of the Žemyna Gymansium (Vilnius) Rūta Krasauskienė and the administrative staff of this Gymnasium (Romalda Čižienė, Aldona Šventickienė, Rasa Kazlauskienė, Šarūnė Saikauskienė and Regina Kryžiuvienė) as well as the colleagues from the Methodolog- ical Group of Natural Sciences Gražina Drebickienė, Evelina Macijausk- ienė, Danguolė Miliauskienė, Ana Lavrinovič and Asta Valčiukienė for their sincere involvement, support and permanent assistance during the period of my doctoral studies. I am also thankful to the Research Foundation of the Lithuanian University of Educational Sciences and the Research Foundation of the Research Council of Lithuania for the support in performing this project.

1. LITERATURE OVERVIEW

A detailed literature overview in given the dissertation (pages 18– 48): 1.1. Short survey of the prehistory of the research; 1.2. Survey of the recent investigation of Nepticulidae in Europe and worldwide (1.2.1. Re- cent investigation of Nepticulidae in Europe and neighbouring territories; 1.2.2. Investigation of Nepticulidae in Asia and Australia; 1.2.3. In- vestigation of Nepticulidae in Africa; 1.2.4. Investigation of Nepticulidae in Central and South America); 1.3. Published data about the endobiontic lifestyle within plant tissues; 1.4. Data about characters used for the identification of Nepticulidae taxa; 1.5. Data about the taxonomy and phylogeny of Nepticulidae; 1.6. New taxa described by the Biosystematics Research Group; 1.7. Survey of the investigation of the Nepticulidae of Lithuania and the Crimea (1.7.1. Investigation of the Lithuanian fauna; 1.7.2. Investigation of the Crimean fauna).

15 2. MATERIAL AND METHODS

2.1. RESEARCH METHODS

2.1.1. Rearing of adults from miners’ larvae – the fundamental method applied in the present research Registration of the traces of larval activity (the mines or leaf-miners) was conducted by thorough inspection of damaged parts of host-plants or potential host-plants. Most of the Lithuanian Nepticulidae usually mine leaves, more seldom apple-tree or maple buds or maple samaras (e.g. Etainia sericopeza), and stems and young bark (Ectoedemia atrifrontella) (Puplesis, 1994, Puplesis, Diškus, 2003). Based on our own research experience, some methodological recommendations have been suggested and published. These methodological recommendations are discussed in our monograph with co-authors (Diškus, Stonis, 2012), in the chapter by J. R. Stonis, A. Diškus, and A. Navickaitė: „Nepticulidae investigation methods“(Stonis et al., 2012). During our doctoral studies, following Stonis et al. (2012) and Stonis et al. (unpublished), five groups of mining abundance were accepted according to the mine count (empty and with larvae). The following system of verbal description of species and detection (identification) codes were employed. The occurrence of leaf-miners’ species (common/rare) was determined by summing up the spatial distribution and species mining abundance: Occ = spA + spD. (Occ – species occurrence: common/rare; spA – species mining abundance in an analysed territory: collecting site/locality, region, country, etc.; spD – species spatial distribution in an analysed territory) (Stonis et al., 2012, and unpublished). Following Stonis et al. (2012) and Stonis et al. (unpublished), a species is regarded as very common when Occ is 10–9, common when Occ is 8, not rare when Occ is 7–6, rare when Occ is 5–4, and very rare when Occ is 3–2.

16 2.1.2. Collection of material using light traps This method is most relevant in the countries with hot climate when night temperature oscillates around +25°C – +28ºC (Puplesis, Diškus, 2003). A sheet of white stiff cloth as a screen and ultraviolet spectrum bulbs (125 W and 250 W) were used for attracting insects. It is recommended that on windy nights with suitable temperature, the bulb should be hung lower in dense but low scrubs whereas the upper edge of the screen should be turned down by 90° to as to form a hood (Stonis, Remeikis, 2011). The Nepticulidae specimens were collected using small glass test- tubes (5 mm in diameter and 3–4 mm in depth) plugged with a tight cotton wad, and, following Puplesis (1994) and Stonis (unpublished) kept alive in a dark container, with a few plant leaves added to preserve some moisture. It is best when the collected material is pinned and set (i.e., wings spread) outright (before the collected specimens get dead), or kept alive in a refrigerator at +10ºC and prepared in the first half of the next day. The dead material stored at higher temperatures and dry air conditions loose elasticity in 2–4 hours and crumbles at pinning. For killing collected specimens, no longer acetil acetat, chloroform or organophosphorus insecticides were used, but each test-tube with live specimen was placed individually in hot (just boiled) water for a one-two seconds. Some specimens were placed alive in individual sterile plastic test-tubes with 70 % alcohol and kept in a freezer for later molecular studies.

2.1.3. Preparation of collected specimens The reared under laboratory conditions or freshly collected specimens are pinned under a stereoscopic binocular microscope on a special elastic cushion using 5–10 mm stainless minutien pins. The insect wings are fixed using narrow (1–1.2 mm wide and 8–10 mm long) transparent paper strips pinned with D1 and D2 minutien pins. The mounted material is left to dry for about 14 days. During the fieldwork, a simpler technique for spreading wings can be used: the wings of the fresh individual are spread by air flow (blowing from the caudal end) and left to dry (there is no need to worry about the fixation or regularity of wing setting). The correct wing setting of collected material not always is successful. It also depends on the correct

17 pinning. For taxonomic analysis, this preparation stage can be dismissed, however, when the wings of the examined specimen are spread it is easier to observe and describe the colour of the abdomen and arrangement of androconial scales on the hindwings and forewings; it is also easier to snap off the abdomen for genitalia preparation (Stonis et al., 2012b). Under the laboratory conditions, the specimens are mounted on small (1.5 mm wide and 7–10 mm long) white plastic strips and labelled. The labels contain information about the species, finding locality, habi- tat, date of finding, name of collector, information about the host-plant, fieldwork card number, and, if applicable, and altitude above sea level; the labels of night-trapped insects also may contain the information about the light trap. The collected material is stored in hermetically closed boxes to prevent from pests, moisture and bright light. In the zone of hot and humid climate the boxes also may be supplied with fungicide powder to prevent from formation of mould.

2.1.4. Preparation and examination of temporary micro-mounts of genital structures After Puplesis, Diškus (2003) and Stonis et al. (unpublished), the stages of preparing temporary micro-mount of genital structures in glycerine are as follows. 1. Under stereoscopic microscope with an attached background of sticky white plastic plate and using a handled minutien pin the abdomen of the insect is snapped off by gentle movements up and down. 2. The snapped off abdomen is transferred into a test-tube with a handled minutien pin steeped in glycerine; about 1 ml of potassium hy- droxide (10% KOH) is dripped into the test-tube using a pipette. 3. The test-tube is heated on an open flame (e.g. spirit lamp) or in boiling water; the abdomen is boiled until it becomes transparent; during the boiling, the test-tube should be jolted to prevent the squirting out of air bubbles together with the abdomen. 4. The content of the test-tube is poured out into a clean small Petri dish and then, using a preparation needle, transported to other dish with boiled or distilled water. 5. Gently moving the handled minutien pin, the abdomen is rinsed. 6. A drop of glycerine is dripped on a clean slide which is covered with cover slip in such a way as to leave part of the glycerine drop uncovered. 7. The rinsed mount is transported into the glycerine drop and cautiously pushed under the

18 cover slip with the handled minutien pin; the mount should be thrust between the slide and cover slip ventral part upside; for this purpose, a stereoscopic binocular microscope with magnification of 28–56 times is used. 8. The temporary micro-mount is examined using a study microscope with higher magnification. 9. The temporary micro-mount is stored in supersaturated sugar solution (i.e. in sugar crystal) or in glycerine drop (in a minute test-tube or pit of plastic strip which is covered with another strip of the same kind (Puplesis, 1994; Puplesis, Diškus, 2003). Only the temporary mount provides a possibility to observe and photograph the morphological structures laterally or to image morpho- logically interesting or diagnostically important sclerites from all sides whereas it is impossible to roll and observe or photograph laterally the permanent mount. Yet as the examined temporary mounts are stored in glycerine their structures are not as clearly visible as those of transparent permanent mounts (the quality of their photographs also are worse than those of permanent and highly transparent micro-mounts). Some time ago, drawings or photographs of temporary mounts alone sufficed for documentation of newly discovered species. Nowdays, all Nepticulidae species mainly are described and compared based only on the permanent mounts. However, the preparation of temporary mounts preserves the natural structure of spatial genitalia of Nepticulidae, i.e. it changes but little or does not change altogether. Moreover, temporary mounts offer a possibility of repeated examination and documentation of the spatial characters of structures and sclerite links. For this reason, it is recommended that the initial observation and documentation of the little known (or exotic) fauna is conducted using glycerine as a mounting medium. Euparal should be used during further examination (Diškus, Stonis, 2012).

2.1.5. Preparation, examination and documentation of permanent micro-mounts of genital structures After Puplesis, Diškus (2003) and Stonis et al. (unpublished), the stages of preparing permanent micro-mount of genital structures in Euparol are as follows. 1. For preparation of the permanent micro-mount, the genital armature stored in glycerine is rinsed with boiled or distilled water, then transferred with a handled minutien pin into a pit of clean slide with and

19 30% ethanol (ethyl alcohol) solution and under stereoscopic microscope separated from the abdomen. 2. After the dissection, the genital armature and the abdomen pelt are transferred into a pit of clean slide with 70% ethanol and the mount is cautiously rinsed; the scales adhering to the abdomen pelt are cleaned using a very small and thin brush or(and) a very thin and sharp handled minutien pin. 3. After this partial dehydration, the genital armature and the abdomen pelt are died with spirit solution of Chlo- razol Black. 4. The final dehydration of the genitalia and abdomen pelt is performed, pure ethanol is dripped over and the mount is cautiously rinsed using a handled minutien pin. 5. A small drop of Euparal (if thicker than fresh honey it should be diluted with Euparal essence) is dripped on a clean slide. 6. Using the stereoscopic microscope, the genitalia and abdomen pelt are transferred into the Euparal drop and covered with a very small cover slip. The genital armature is fixed ventral part upside whereas the separate sclerites can be spread or even separated (dissected); sometimes, the abdomen pelt or separate sclerites of genital apparatus (e.g. phallus dissected from the capsule) are fixed under separate cover slip but on the same slide. 7. The mount is labelled (a paper label is tagged on the slide), placed onto a special card or plastic slide tray and dried for 2–3 months at room temperature or for 20 days in a heating oven (at +50º–60º) (Diškus, Stonis, 2012). It is recommended to photograph the permanent genital mounts right after the preparation. Each prepared permanent micro-mount must be numbered; the photographs of the genital structures must be supplied with the same data. In our study, the genital micro-mounts were examined using Leica DM2500 microscope and Leica DFC420 digital camera connected with the microscope and computer. The adults were measured using ocular micrometre and stereoscopic binocular microscope MBS-10. The length of the wingspan was measured from the end of the left wing cilia and the end of the right wing cilia. If the measured specimen individual is not straightened out the length of one of the forewings is indicated. The indicated length is multiplied by two and the thorax width is added. For depiction of the external coloured morphological patterns of adults Faber-Castell watercolour pencils can be used or specimens can be photographed using Leica S6D microscope and computer-linked DFC290 camera.

20 2.1.6. Chorological method The distribution ranges of species (or taxa of other ranks) were chor- ologically analysed. The distribution ranges of all analysed species were thoroughly mapped. Taxa distinguished for the same or comparable distribution range were joined into groups which, in their turn, were named indicating the type of the range. The chorological analysis not only included the congeneric species but also the unrelated taxa. This type of species evaluation requires all available information about the species, including trophic relationships and trophic specializa- tion, distribution of host-plant, and habitats.

2.1.7. Cladistic analysis For proving monophyly (as also for grounding sister clades) as many characters as possible were used including not only the morphological characters but also chorological and ecological ones. In the upper part of the cladograms, the recent specie were registered whereas the clades proving the possible relationship were drawn only dichotomically. Though different numbers of synapomorphic and autapomorphic characters were chosen for different clades the higher number of synapomorphic characters is more reliably indicative of evolutionary links (i.e. the fact of common ancester) between sister taxa and implies older evolutionary processes. Some distinguished clades could be taxonomically ranked yet our analysis confined to already identified and described species groups. In the present dissertation, an attempt was made to relate the phylogenetic clades with taxa distribution ranges in different continents.

2.2. MATERIALS

The author conducted fieldworks during six years (2008–2013) and collected scientific material of Nepticulidae from different Lithuanian regions. The collected and analysed material is stored in the collection of LEU Biosystematics Research Group. In the dissertation, the new data were generalized and the data provided by other researchers were for the first time revised and generalized.

21 The integration of individual data and data published by other researchers offered a possibility for full-scale analysis of the Lithuanian Nepticulidae fauna (Fig. 2.2.1.).

Fig. 2.2.1. Localities of Nepticulidae investigations in Lithuania from which the data obtained served as a basis for the present dissertation.

In 2011, the author, assisted by A. Diškus, J. R. Stonis, G. Varačinskas (LEU) and S. R. Hill from the UK carried out intensive long-term fieldworks in the Crimea: Karadag and Yalta Forest Reserve, Angara Moun- tain Pass, Feodosia, Koktebel, Alushta, and other localities. During the fieldwork, the author collected big original scientific material. Also the author dissected and identified so far unidentified sizea- ble material collected in 1977–1989 in Karadag and its environs by other researchers: S. Yu. Sinev (ZIN), Yu. I. Budashkin (KNURAS), I. Reznik (ZIN), and R. Puplesis (ZIN). This material was submitted for scientific analysis by foreign scientific research institutions. Also for comparative Nepticulidae analysis and validation of species identification, abundant collection material (about 450 specimens) from the neighbouring Central Asia was analysed including typical series of about 40 species (for the first time, permanent micro-mounts were prepared and genital structures were digitally documented). Other comparative material was collected together with the colleagues during expeditions to the Uttarakhand District (India, 2010) but also to Na- ples (Campania, Italy, 2012) and Bergamo (Lombardy, Italy, 2013) (Fig. 2.2.2.).

22 About 3300 Nepticulidae specimens were investigated for precise species identification. More than a half of them were dissected and mounted: temporary or permanent micro-mounts of genital structures were prepared.

Fig. 2.2.2. Localities where in 2008–2013 the author collected material for the present dissertation: A – in Europe (Lithuania (1), the Crimea (2) and Italy (3, 4); B – in the Himalaya (localities where other authors collected material analysed in the present dissertation).

23 3–5. RESULTS

3. GENERAL EVALUATION OF NEPTICULIDAE FAUNA

3.1. INVENTORISATION AND EVALUATION OF NETICULIDAE FAUNA OF THE WORLD

Th e results of the current research work, which included consistent complementation of the database, analysis of the newest data from different biogeographical regions of the world, and generalization of yet unpublished data on the scarcely explored South Africa and totally unex- plored equatorial Africa and other tropic and subtropical regions, allow assuming that so far only about half of all Nepticulidae species (i.e. 850 species) has been identifi ed. According to our data, 449 Nepticulidae species have been registered in the Palaearctic region (Fig. 3.1.1.).

Fig. 3.1.1. Distribution of Nepticulidae species in the biogeographical regions of the world

24 The Stigmella genus is idiosyncratic as it is dominant in the fauna of all regions of the Earth (except Antarctica). Currently 399 species of this genus are known. The trophic relationships of 63% of world fauna species have been established yet biology of almost half of these pygmy moths is unknown (a more thorough survey is given in the dissertation).

3.2. SPECIES GROUPS OF STIGMELLA TROPHICALLY ASSOCIATED WITH QUERCUS AS HOST-PLANT PLANTS

The current dissertation addresses a new species Stigmella acuta (Diškus, Navickaitė & Remeikis, 2013) belonging to the S. hemargyrella species group (the other 13 discovered new species are not discussed). The mentioned species has been found in Himalays. Oak (Quercus acutisima) is its host-plant. This information has been published in an article with co-authors (Stonis et al., 2013b). For the first time, trophic relationships of the S. hemargyrella species group with Quercus has been elucidated. Thus it has been established that seven Stigmella species groups (in- stead of five) are partly or ultimately trophically associated with oaks: caerusifasciella (East Asia and Japan; one species registered); saginella (North and Central America; 10 species); quercipulcella (North and Cen- tral America; five species); ruficapitella (Palaearctic; 29 species); castanopsiella (East and South-East Asia; 6 species), hemargyrella (Palaearctic; a total of 6 species including S. acuta Diškus, Navickaitė & Remeikis) and cornuta (East Asia; one species). A detailed survey of groups of the species groups is given in the dissertation and in the mentioned our article with co-authors (Stonis et al., 2013b).

3.2.1. Taxonomic diagnosis and description of the recently discovered species from the S. hemargyrella group Stigmella acuta Diškus, Navickaitė & Remeikis Stigmella acuta Diškus, Navickaitė & Remeikis, 2013; in Stonis et al., 2013: 202–206. Type material. Holotype: ƃ, Himalaya, India, Uttarakhand, Dehradun District (Mussoorie), 30º27’31’’N, 78º01’46’’E, elevation

25 1980 m, mining larva on Quercus acutissima, 15–18.viii.2010, ex pupa 02.ix.2010, genitalia slide no. AD480 (ZMUC). Paratypes: 1ƃ, the same label data, pupa, genitalia slide no. AD481; 1Ƃ, Tehri Garhwal (Cham- ba), 30º20’39’’N, 78º28’59’’E, mining larva on Quercus acutissima, 24.viii.2010, ex pupa 06.ix.2010, genitalia slide no. RA265 (ZMUC). Diagnosis. A distinctive, fuscous-winged species with a cream to pale orange fascia in the forewing; the pointed valva, M-shaped gnathos, uncus with four small caudal lobes, large acute cornuti, developed manica, androconial scales on male hindwing, also folded accessory sac in female genitalia indicate that it belongs to the Stigmella hemargyrella group. Externally, males of the new species can be easily distinguished from all other currently known S. hemargyrella group species, except the Fagus-feeding S. hemargyrella (Kollar), by the basal patch of dark brown androconia in the hindwing. In male genitalia, S. acuta differs from all Stigmella species by the shape and number of the large and pointed cornuti in the phallus (Fig. 3.2.1.1).

Fig. 3.2.1.1. Male genitalia of the recently discovered and described species Stigmella acuta: A – capsule, B – phallus with well-developed cornuti (published by A. Navickaitė in a co-authored paper by Stonis et al., 2013b).

Fig. 3.2.1.2. A – Female genitalia of the recently discovered and described species Stigmella acuta; B – Stigmella acuta leaf-miner on a Quercus acutisima leaf (published by A. Navickaitė in a co-authored paper by Stonis et al., 2013b).

Bionomics. A mines in leaf (Fig. 3.2.1.2 B). Host-plant: Quercus acutissima Carruth. (Fagaceae). Egg on upper side of the leaf. Larvae mine in August. Contorted or sinuous gallery of mine filled with brown-black frass; the first half of the mine completely filled with frass; in the second half of the mine frass coiled, leaving clear margins. Larva pale green to yellowish, with red-brown intestine and dark brown head capsule. Larval exit slit on upper side of the leaf. Cocoon brown to ochre-brown; length 2.5 mm, maximal width 1.5 mm. Adults fly in September. Distribution. It occurs in mountainous broadleaf and mixed forests of the western Himalaya at altitude about 2 km; possibly common: many empty leaf-mines were observed in different localities. A detailed description of the species is given in the dissertation and in our article with co-authors (Stonis et al., 2013).

3.2.2. Characteristics of species groups of the Stigmella genus trophically associated with Quercus A survey of the groups was published in 2013 in our article with co-authors (Stonis et al., 2013) and explicitly represented in the current dissertation.

27 We have established that the species groups can be diagnosed on combinations of characters of male and female genitalia. It was for the fi rst time demonstrated that diagnostics of the groups also is possible using the individually worked out dichotomous system of specifi c characters.

3.2.3. Diagnostics of the recognized groups of using the dichotomous systems During the current research, dichotomous diagnostic systems based on the characters of male and female genitalia structures were worked out which allow reliable diagnosing of the 6 recognized groups of species of Stigmella trophically associated with Quercus (Figs 3.2.3.1–3.2.3.3).

Uncus

No lobes Bilobed S. saginella group S. caesurifasciella group S. quercipulchella group uncus S. ruficapitella group S. castanopsiella group tegumen S. hemargyrella group S. cornuta group

Uncus

Well separated narrowed long lobes Short broad lobes S. saginella group S. ruficapitella group S. quercipulchella group S. castanopsiella group S. hemargyrella group S. cornuta group

Gnathos

Broadly U-shaped Central plate and two caudal processes S. ruficapitella group S. caesurifasciella group S. saginella group S. quercipulchella group S. castanopsiella group S. hemargyrella group S. cornuta group Fig. 3.2.3.1. Dichotomous diagnostic systems of the species groups based on uncus and gnathos, characters (published by A. Navickaitė in a co-authored paper by Stonis et al., 2013b).

28 Cornuti

Extremely large Small to large S. caesurifasciella group S. cornuta group S. quercipulchella group S. ruficapitella group S. castanopsiella group S. hemargyrella group S. saginella group

Manica

Absent Developed S. caesurifasciella group S. saginella group S. quercipulchella group S. cornuta group S. ruficapitella group (very most) S. castanopsiella group S. hemargyrella group manica

Accessory sac

Very large and spiral Not spiral, sometimes short

S. quercipulchella group S. caesurifasciella group S. saginella group S. ruficapitella group S. castanopsiella group S. hemargyrella group

Signa

Strongly developed Absent S. caesurifasciella group S. saginella group S. quercipulchella group S. ruficapitella group S. castanopsiella group S. hemargyrella group

Fig. 3.2.3.2. Dichotomous diagnostic systems of the species groups based on manica, accesory sac and signa characters (published by A. Navickaitė in a co- authored paper by Stonis et al., 2013b).

29 Gnathos

With single central process Without central process

S. caesurifasciella group S. saginella group S. quercipulchella group S. ruficapitella group S. castanopsiella group S. hemargyrella group S. cornuta group Juxta

X-shaped Absent or not developed

S. saginella group S. caesurifasciella group S. quercipulchella group S. ruficapitella group S. castanopsiella group S. hemargyrella group

Phallus and vesica

Bulbous basally, vesica coiled Not bulbous, vesica not coiled S. caesurifasciella group S. castanopsiella group S. saginella group S. quercipulchella group S. ruficapitella group S. hemargyrella group S. cornuta group

Cornuti

Not numerous and not aggregated into band Numerous, aggregated into band

S. saginella group S. caesurifasciella group S. cornuta group S. quercipulchella group S. ruficapitella group S. castanopsiella group S. hemargyrella group

Fig. 3.2.3.3. Dichotomous diagnostic systems of the species groups based on gnathos, juxta, phallus and cornuti characters (published by A. Navickaitė in a co- authored paper by Stonis et al., 2013b).

3.2.4. Cladistic analysis of the species groups of Stigmella trophically associated with Quercus The species of the Stigmella genus were analysed by cladistic method. Thirty seven apomorphic characters were selected for cladistic analysis. All of them are listed in the dissertation.

30 Fig. 3.2.4.1. Cladogram of the species groups of the Stigmella genus associated with Quercus.

31 Fig. 3.2.4.2. Phylogeny and geographical distribution trends of the species groups of Stigmella associated with Quercus.

32 The cladistic analysis showed that only four of the 7 species groups of Stigmella trophically associated with Quercus comprise a monophyletic clade: S. quercipulchella, S. hemargyrella, S. ruficapitella and S. castanopsiella (Fig. 3.2.4.1). Only for these species groups a possible common ancestor was determined, i.e. 4 apomorphic traits (1–4) of the cladogram (Fig. 3.2.4.1) were chosen as a proof of monophyly. Two phylogenetic clades can be distinguished in this complex of the species groups, yet both of them only are based on one–two apomorphies. S. quercipulchella and S. hemargyrella, as well as S. ruficapitella and S. castanopsiella, are sister clades and obviously independent taxa (each of them is based on 3–6 autapomorphic characters) (Fig. 3.2.4.1). Mean- while, S. saginella, S. caesurifasciella and S. cornuta groups can be characterized by many autapomorphic characters which during the cladistic analysis were in no way associated with the complex of S. quercipulchella–S. castanopsiella groups but addressed to as independent phylogenetic clades. The S. quercipulchella–S. castanopsiella complex was not given taxonomic rank yet; the data of the cladistic analysis could support the description of taxonomic rank of tribe. Linking up of the geographical distribution data with the cladogram (Fig. 3.2.4.2) showed that the two groups of species without the common ancestor only are found in North and Central America whereas all other groups of species (with and without the common ancestor) are spread in the Eastern Hemisphere (mainly in East Asia). Approaching from a different angle, it could be possible to suggest that Stigmella fauna of North and Central America and Eurasia trophically associated with Quercus are not of the same origin but represent a few different evolutionary tendencies (clades).

33 4. THE CRIMEAN NEPTICULIDAE WITHIN THE CONTECT OF THE EUROPEAN FAUNA

4.1. A SURVEY OF THE EUROPEAN NEPTICULIDAE FAUNA AND A REVISED TAXONOMIC NOMENCLATURE

Data analysis showed that currently 264 species of Nepticulidae belonging to 11 genera are known in the European fauna (Fig. 4.1.1). As was pointed out in the survey of literary sources, in the last ten years, the investigations of the European Nepticulidae fauna have be- come very intensive. New possibilities to analyse the geographical distribution and ecological traits of this family of pygmy moths have emerged. During the mentioned time span, the taxonomic nomenclature of Nep- ticulidae of the old continent also has changed. A detailed analysis is given in the dissertation. Th e dissertation also contains the taxonomic nomenclature of all Nepticulidae species known in the European fauna.

Fig. 4.1.1. Th e revised taxonomic composition of the European Nepticulidae fauna.

34 4.2. TROPHIC RELATIONSHIPS OF THE EUROPEAN NEPTICULIDAE

Analysis of the European Nepticulidae fauna showed that trophic relationships had been established for 89% of species. Host-plants of 11% of European species are still unknown. A detailed analysis of host-plants revealed that the European Nep- ticulidae are trophically associated with 97 plant genera. The majority of Nepticulidae species mines plants of Quercus (42), Betula (11) and Salix (10) genera. Yet 47% of plant genera are engaged by single mining Neptic- ulide species. Woody plants are dominant (66%) among the host-plants of the Eu- ropean pygmy moths whereas 34% of the known pygmy moths are trophically associated with grasses. The majority of the pygmy moths of the old continent mine plants of Rosaceae, Fagaceae, Fabaceae, Salicaeae and Betulaceae families. Rosacae is the dominant plant family with which 58 Nepticulidae species have trophic relationships; 17% of Nepticulidae species are trophically associated with Fagaceae family. However, 14 known host-plant families are mined only by one species of leaf-miners. In the current dissertation, 31 host-plant families of the European Nepticulidae are registered.

4.3. RESULTS OF CHOROLOGICAL ANALYSIS OF THE EUROPEAN NEPTICULIDAE

Based on chorological method (see Methods) for the first time applied for analysis of the European Nepticulidae fauna it was established that Nepticulidae species in this part of the world belong to 10 chorological groups: Euro-Nemoral, Euro-Submediterranean, Euro-Medi- terranean, Sub-Mediterranean, Sub-Mediterranean–Mediterranean, West-Palaearctic, Euro-Siberian, Trans-Palaearctic, and Holarctic. The majority of species are related with the Mediterranean and Sub-Mediterranean chorological groups (74%). The largest groups in the European fauna are: Mediterranean (29%), Euro-Mediterranean (12%) and Sub-Mediterranean (6%); also many species belong to the Eu- ro-Nemoral group (9%).

35 Fig. 4.3.1. Chorological groups of the European Nepticulidae by size (relative percentage).

Th e European Nemoral – wooded – region includes the Euro-Boreal, Euro-Atlantic, Central European, Euro-Continental, and Sub-Mediterra- nean provinces (ecoregions). Th e European Nepticulidae fauna analysed in the current dissertation was interpreted as “Euro-Nemoral Nepticul- idae fauna” (in spite that some Nepticulidae species mine woody plants and some other – the smaller part – mine grasses). Th e Sub-Mediterranean province (ecoregion), which, based on the individual investigations conducted in the Crimea, is for the fi rst time elaborated pointing out the characteristic Nepticulidae fauna, represents a transitory zone between the Euro-Nemoral and Mediterranean provinces. Among the 54 Nepticulidae species attributed to the Euro-Submed- iterranean group, Ectoedemia quinquella and Glaucolepis lituanica are the most interesting ones. Th e latter has been recently published as a new species found in a few extremely remote and isolated European localities (including Lithuania). Presumably, this species should be considered as a relict rather than a species spreading from north to south due to the changing climate. Meanwhile, E. quinquella can serve as a typical exam- ple of a spreading species: the species is widely known in the Sub-Med- iterranean province (ecoregion) and, according to the data of our col-

36 leagues (van Nieukerken et al., 2010), has gradually spread in the Atlantic part of France, Benelux countries and, later, in the southern part of GB. In our opinion, the expansion of the distribution range of this species is related to the climate changes. As a rule, species (17) of the Sub-Mediterranean chorological group inhabit very limited areas. Stigmella rhamnella, S. szoecsiella and Ectoe- demia contorta, which have trophic relationships either with Rhamnus (1 species) or with Quercus (2 species) host-plants, make an exception. Among the identified 21 species distributed in the Palaearctic region, two can be classified as Submediterranean Stigmella( paliurella and Glau- colepis melanoptera) and other two as Mediterranean–Asian (Stigmella minusculella and Etainia obtusa). Based on the geographical distribution of flora and fauna (e.g. Acer platanoides, Carpinus betulus, Olea europaea and distribution ranges of other plants), specific features of the European climate and landscape, and individual research data an original map of the European zoogeo- graphical provinces was compiled. The fauna of the Euro-Atlantic province (ecoregion) comprises 123, Central European 141 and Euro-Mediterranean 164 Nepticulidae species. Meanwhile, 151 species of pygmy moths have been identified in the Sub-Mediterranean province. The data about the geographical bounda- ries of the Sub-Mediterranean province and the identified fauna of pygmy moths are original, based on the results of individual field investigations and chorological analysis (Navickaite et al., 2012). The analytical data showed that the greatest number of the European Nepticulidae species is found in the Mediterranean province – 191 species. Yet the identified Nepticulidae fauna is unevenly distributed: some species occur in the area between the Iberian Peninsula and Greece or South-West Turkey (and Cyprus) whereas some of them are bound only to a certain part of the Mediterranean province. The number of Nepticu- lidae species identified in the western part of the Mediterranean province amounts to 109, in the central part 94 and in the eastern part 99. From the taxonomical point of view, the unexpectedly high number of species identified in the Mediterranean province allows assuming that the distribution of some Nepticulidae genera is strongly restricted within the Mediterranean province (ecoregion) as the number of species

37 identified in this province accounts for 93% of the European fauna of Glaucolepis genus. A very strong bond to the Mediterranean province also is characteristic of Acalyptris, Ectoedemia and Parafomoria genera (respectively 89%, 86% and 85% of the total of the species of this genus identified in Europe). The largestNepticulidae genus (Stigmella) represented in Europe by 109 species also is widely distributed in this province (73% of the European Stigmella fauna). A detailed survey is provided in the dissertation.

4.4. CHECKLIST OF NEPTICULIDAE OF THE CRIMEA

4.4.1. Inventorization of Nepticulidae of the Crimea The first material of Nepticulidae was collected in the Crimea in 1977; however, for a long time only a small number of attempts were reported to identify and publicize the fauna of this Lepidoptera family from the Crimea. A comprehensive history of the research is presented in the literature overview section of the dissertation. In 2009–2010, under the “Crimea Project”, the author identified 31 species of Nepticulidae, among which Stigmella ulmiphaga and Ectoe- demia rufifrontella were recorded for the first time from the Crimea; it was reported in our publication (Stonis et al., 2013d). Out of more than 600 examined specimens, only 3 were attributed to the Trifurcula genus: one to T. subnitidella, and two to Trifurcula macedonica, a new species within the fauna of the Crimea. Stigmella szoecsiella, Ectoedemia albifasciella and E. pubescivora were excluded from the list of the Crimean species because of misidentification of these species or insufficient scope of previous research. The author of the dissertation, after having concluded her work in Karadag, Yalta, Feodosia, Sudak, Alushta, Simferopol and other locations in 2011, succeeded in identifying 26 species of Nepticulidae which were new to the fauna of the Crimea. This section presents a revised and fully updated checklist of the Nepticulidae species of the Crimea, chorological and trophic analysis of the species and characteristics of the life-cycles. The checklist includes comprehensive data about 20 species from the Stigmella genus, 1 species respectively from Bohemannia, Ectoedemia and Fomoria genera and

38 3 species from the Trifurcula genus, all of which are recorded for the first time for the fauna of the Crimea.

4.4.2 The novelty of the Crimean fauna After the identification of the scientific material of Nepticulidae (preparation of temporary mounts and permanent slides in Euparal) and accomplishment of fieldwork tasks, we succeeded in considerably updat- ing the list of the Crimean Nepticulidae. 26 new species were identified, i.e. 40% of the whole of the Crimean fauna. One species, Ectoedemia spinosella, was not included into the list because of previous misidentifi- cations having resulted from referring exclusively to leaf-mines (see van Nieukerken et al., 2004); our presumption is that E. mahalebella has been misidentified and mistaken as E. spinosella. At present the fauna of the Crimea includes 64 Nepticulidae species belonging to the Simplimorpha (1), Stigmella (41), Bohemannia (1), Ectoe- demia (10), Fomoria (1), Acalyptris (1), Trifurcula (5), Glaucolepis (2) and Etainia (2) genera. TheBohemannia and Fomoria genera were registered as new within the Crimean Nepticulidae fauna for the first time during our investigation. The greatest number of the new Nepticulidae species were identified within the Stigmella genus (20), which makes 50% of the fauna of this genus. Trifurcula subnitidella and T. josefklimeschi had been the only Trifur- cula species in the Crimea. Additionally, we succeeded to document the distribution of the T. silviae and T. eurema in the Crimea. T. macedonica was identified during the study of the material collected in 1987. Accord- ing to these data, 15% of Europe’s species of the Trifurcula genus occur in the Crimean peninsula. When analysing trophic relationships among Nepticulidae registered in the Karadag Nature Reserve and its surroundings, we identified that the majority of the newly identified species were related to the Rosaceae (35%) and Betulaceae (20%) host-plants. Our study presents Trifurcula macedonica and Ectoedemia mahalebella without describing new subspecies, though the investigated material is of very distinctive nature.

39 4.4.3. Th e updated checklist of Nepticulidae of the Crimea Th e dissertation presents a comprehensive checklist of all the species identifi ed in the Crimea making up 20.5 printed pages. Data about the species are presented in the following order: the name, author and year of the description of the species, the studied material, the description of the general geographical distribution, and notes.

4.5. CHOROLOGICAL ANALYSIS OF NEPTICULIDAE OF THE CRIMEA

4.5.1. Chorological evaluation of the Nepticulidae fauna of the Crimea Th e mapping and analysis of Nepticulidae distribution ranges taking into account the Crimean and general distribution showed that from the chorological point of view, Nepticulidae of the Crimea could be divided into 8 chorological groups. Th ese groups could be aggregated into two larger distribution groups: the European (Euro-Submediterranean + Euro-Mediterranean + Mediterranean + Sub-Mediterranean + Mediter- ranean + Euro-Nemoral) and Palaearctic (Western Palaearctic + Euro-Si- berian + Trans-Palaearctic) (Fig. 4.5.1.). Th ere is a tendency that 72% of species from the Crimea belong to chorological groups that are more or less related to the Mediterranean ecoregion. It leads to a conclusion that these species could have originated in the South and later gradually spread all over the European continent.

Fig. 4.5.1. Chorological composition of the Crimean Nepticulidae fauna deﬁ ned during the chorological analysis.

40 The dissertation presents a detailed survey of chorological groups and also lists and discusses the species attributed to these groups.

4.5.2. Distribution of common species among different chorological groups After having evaluated occurrence (data on geographical distribution and quantity/abundance, see Methods), we identified the common and rare species of the region. Rather a large number of species identified in the Crimea are characterized by wide distribution and moderately abundant mining (19%) or wide distribution and extremely abundant mining (17%). Among the Crimean fauna that has been analyzed, there are a number of species characterized by limited distribution and moderately abundant mining (12%), limited distribution and not abundant mining (12%) and local distribution and not abundant mining (12%). Only those species geographical distribution of which had been rated at no less than 5 points and abundance of mining at no less than 3 points (see Methods) were attributed to the category of common species. Com- mon species can also be divided into common and highly common; the latter are characterized by wide or very wide distribution and very abundant or abundant mining. Common species may be characterized by limited distribution if their mining has been identified as very abundant. The dissertation lists common and highly common species in full detail. The common species identified within the Crimean fauna predom- inantly are of the Euro-Submediterranean chorological group (47%), whereas, contrary to our expectations, the most rare among the common species are those characterized by Sub-Mediterranean/Mediterranean ranges (18% of the common species of the Crimea).

4.5.3. Common species and their host-plants The common Nepticulidae species of the Crimea are trophically associated with 13 plant genera, among which Quercus, Sorbus, Crataegus and Prunus are the most prolific (if estimated by the number of mining Nepticulidae species) (Fig. 4.5.3.1.).

41 Fig. 4.5.3.1. Identified trophic relationships of common and highly common Nepticulidae species.

A more detailed analysis is presented in the dissertation.

4.5.4. Distribution of rare species among diff erent chorological groups Rare species of the region are characterized by limited geographical distribution and not abundant or sporadic mining (see methods). How- ever, in line with the applied methods, some of the widely distributed species were attributed to the rare category if their mining was sporadic; and, on the contrary, rare species may be characterized by abundant or very abundant mining if their geographical distribution has been identifi ed as highly limited or based on solitary fi ndings (according to Stonis et al., unpublished). Among the investigated Crimean fauna, there are species which are characterized by limited distribution and not abundant mining (5% of the entire fauna), solitary fi nding and not abundant mining (2%) and highly limited distribution and sporadic mining (2%). Rare species may also be divided into rare and highly rare; the latter, according to Stonis et al. (unpublished), are characterized by only highly limited distribution or solitary fi nding and not abundant or sporadic mining. Th e dissertation lists the rare and highly rare species of the Crimea in full detail. Th e rare species of the Crimean fauna mainly belong to the Eu- ro-Mediterranean chorological group (36%); a number of rare species

42 belong to the Euro-Submediterranean and Euro-Nemoral groups (18% respectively to both groups) and the Sub-Mediterranean chorological group (14% of rare species). As it could have been predicted, species which are characterized by wide ranges and Sub-Mediterranean/Medi- terranean distribution are the fewest within the category of rare species (Fig. 4.5.4.1.).

Fig. 4.5.4.1. Distribution of rare (rare and very rare) Nepticulidae species among various chorological groups.

4.5.5. Distribution of rare species and their host-plants Among the host-plants which are mined by rare species, the most prominent is the Quercus genus: 23% of the total number of host-plants. Th e remaining rare species are associated with 14 other genera of host- plants (Fig. 4.5.5.1.). To date, there are no known data about the Trifurcu- la macedonica host-plant.

43 Fig. 4.5.5.1. Th e identiﬁ ed trophical relationships between the Nepticulidae species of the Crimea.

Th e dissertation presents a more detailed survey of the rare species and more comprehensive discussion regarding some of the species.

4.6. TROPHIC ANALYSIS OF THE CRIMEAN NEPTICULIDAE

Th e majority of the Crimean Nepticulidae are related to the Rosace- ae (18 species of Nepticulidae) and Fagaceae (12 species) plant families. It should be noted that as far as 54 species are adapted to mine woody plants and the remaining 10 species, herbaceous plants. Table 4.6.1 of the dissertation shows revised and supplemented data on trophic relationships of the Crimean Nepticulidae. Highlighted are host-plants identifi ed and approved during our investigation. Th e analysis of the Crimean Nepticulidae showed that 7 species (11% of the identifi ed fauna) mine cultivated plants: Prunus spinosa, P. ceras- ifera, Malus domestica and Pyrus elaeagnifolia. During the investigation, we succeeded to record for the fi rst time the host-plant of Stigmella irregularis, Pyrus elaeagnifolia, which widely occurs not only throughout the Karadag Reserve but also the whole of the mountainous Crimea.

44 4.7. SPECIFIC FEATURES OF THE LIFE CYCLE OF THE CRIMEAN PYGMY MOTHS

During the present research, it has been observed that larval stage (and the whole life cycle) depends not only on the mined plant but also on the mining season, ambient temperature (sunshine or shade) and biological properties of Nepticulidae genus. During our fieldwork, the mining seasonality of the Crimean species was analysed for the first time (Table 4.7.1.). The precise mining time was established for 39 (61%) known Crime- an Nepticulidae species. Most of adults of these species were reared under laboratory conditions. Meanwhile, such species as Stigmella samiatella, S. atricapitella, Trifurcula silviae, Glaucolepis melanoptera, Ectoe- demia atrifrontella and E. longicaudella were recorded solely applying the light trap method. Thus, the seasonal cycles were established taking into consideration the flying time of adults and published literary data. It can be maintained that almost 50% of Nepticulidae species found in Crimea have two generations in a year and about 30% have one gener- ation. It has been reported that under favourable conditions some species may have even three generations in a year (e.g. Simplimorpha promissa, Stigmella aceris, and Trifurcula eurema). Comparison of the life cycles of Crimean species with the life cycles of the same species in other Europe- an countries show some mismatches which might be predetermined by the geographical position of the Crimean Peninsula and the favourable conditions of the coastal zone entailed by the Crimean mountains, which serve as a barrier for cold air masses.

45 Table 4.7.1. New revised data about the mining time of the Crimean Nepticulidae species. – reliably established (mines with mining caterpillars were collected); z – established during ﬁeldworks according to the empty mines); { – conjectural mining by caterpillars; I–III – month decades. No. Species June July August Septem- October ber I II III I II III I II III I II III I II III 1. Simplimorpha promissa z z z

2. Stigmella confusella zz

3. Stigmella irregularis

4. Stigmella freyella z

5. Stigmella tiliae z

6. Stigmella paliurella zz

7. Stigmella microtheriella z

8. Stigmella glutinosae z{

9. Stigmella alnetella {{ z z

10. Stigmella prunetorum zzzz

11. Stigmella aceris z z { z

12. Stigmella paradoxa

13. Stigmella minusculella {{{{{ {{{

14. Stigmella desperatella z

15. Stigmella torminalis

16. Stigmella crataegella {{

17. Stigmella hahniella {{

18. Stigmella hybnerella {{ {{

19. Stigmella catharticella z

46 Continued of the table 4.7.1. No. Species June July August Septem- October ber I II III I II III I II III I II III I II III 20. Stigmella malella z

21. Stigmella rhamnella z

22. Stigmella viscerella zz

23. Stigmella ulmivora {z

24. Stigmella ulmiphaga {z

25. Stigmella thuringiaca z

26. Stigmella rolandi z z

27. Stigmella trimaculella z

28. Stigmella salicis {z

29. Stigmella obliquella {z

30. Stigmella floslactella {{ {{

31. Stigmella tityrella {zz

32. Stigmella carpinella { z {{{

33. Stigmella lemniscella zz

34. Stigmella incognitella z

35. Stigmella perpygmaeella {{ {{

36. Stigmella splendidissi- mella 37. Stigmella plagicolella { z

38. Stigmella atricapitella {{ {{{

39. Stigmella plagicolella {z

47 Continued of the table 4.7.1. No. Species June July August Septem- October ber I II III I II III I II III I II III I II III 40. Stigmella atricapitella {{ {{{

41. Stigmella basiguttella z {{{

42. Stigmella dorsiguttella z z {{{

43. Stigmella samiatella {{{ {{{

44. Stigmella roborella {z {{{

45. Bohemannia pulverosella {{z z

46. Ectoedemia atrifrontella {{{{{{{{{{{{{

47. Ectoedemia longicaudella {{{{{{{{{{{{{

48. Ectoedemia amani {{{{{{{{

49. Ectoedemia similigena {{{{{{{

50. Ectoedemia turbidella {{{{{{{{{{

51. Ectoedemia subbimaculella {{{{{

52. Ectoedemia rufifrontella {{{{{{{

53. Ectoedemia heringi {{{{{{{

54 Ectoedemia contorta {{{{

55. Ectoedemia mahalebella z {{{

56. Fomoria septembrella {{ {{{{

57. Acalyptris platani {z {{{

58. Trifurcula subnitidella {{{{{

59. Trifurcula josefklimeschi {{{{

48 Continued of the table 4.7.1. No. Species June July August Septem- October ber I II III I II III I II III I II III I II III 60. Trifurcula silviae {{{

61. Trifurcula eurema {{{

62. Trifurcula macedonica {{{{{

63. Glaucolepis melanoptera {{{{

64. Glaucolepis bleonella {{{

65. Etainia sericopeza z

66. Etainia louisella

5. THE LITHUANIAN NEPTICULIDAE FAUNA

5.1. TAXONOMIC COMPOSITION OF THE LITHUANIAN NEPTICULIDAE AND AN ANNOTATED CHECKLIST OF THE FAUNA

Currently, 76 species of pygmy moths are known in the fauna of this country. Most of them belong to Stigmella (56 species) and Ectoedemia (13 species) genera. All Nepticulidae species discovered in Lithuania belong to 7 genera (see the Checklist). Comparison of the taxonomic composition of the Lithuanian Nepticulidae fauna with the European one revealed no major differences or unique trends. The Annotated Checklist contained in the current dissertation includes new data about species distribution patterns, trophic speciali- zation, new host-plants, specified seasonal cycles, and occurrence. The common biogeographical distribution is indicated for the first time.

49 Th is nomenclature was published with co-authors in 2012 (Diškus ir kt., 2012). Th e Annotated Checklist occupies 36 pages of the current dissertation.

5.2. CHOROLOGICAL EVALUATION OF THE LITHUANIAN NEPTICULIDAE FAUNA

5.2.1. Chorological groups of the Lithuanian pygmy moths Analysis of the Lithuanian Nepticulidae fauna in chorological as- pect showed that 76 pygmy moth species belong to several chorological groups: Holarctic, Sub-Baltic, Trans-Palaearctic, Euro-Mediterranean, Euro-Submediterranian, Euro-Siberian, West-Palaearctic and Eu- ro-Nemoral (Fig. 5.2.1.1.). Th e Lithuanian Nepticulidae species are distributed both in vast Holarctic and Palaearctic areas and in geographical- ly restricted areas of Europe (Navickaitė, Stonis, 2012).

Fig. 5.2.1.1. Data of chorological analysis: chorological groups of the Lithuanian fauna of pygmy moth.

Th e dissertation elaborates all chorological groups of Lithuanian fauna and names all species of these groups.

50 5.2.2. Regional distribution of Nepticulidae Based on climate regions of Lithuania and some specifi c features of Lithuanian fauna, 8 distribution regions were recognized (Diškus, Stonis, 2012). Th e greatest number of pygmy moth species has been discovered in South-Eastern (63 species) and South-Western (67 species) regions. Mapping of all seventy six species recorded in the Lithuanian regions and their chorological grouping showed that the Curonian Spit region has unique species diversity. So far, 24 species of pygmy moths, or 32% of the total Lithuanian Nepticulidae fauna, have been discovered in this region. Th e recorded species are from all 8 chorological groups. As distinct from what had been expected, part of the pygmy moth fauna (from 26 to 31%) almost in all Lithuanian regions is represented by Euro-Submediterranean distribution group (Navickaitė, Stonis, 2012). Th e best explored regions (the South-Eastern and South-Western) are predominated by Euro-Nemoral and Euro-Submediterranean species whereas the Coastal region stands out for abundance of species of Eu- ro-Mediterranean group (Fig. 5.2.2.1.).

Fig. 5.2.2.1. Chorological groups of Nepticulidae in the Lithuanian regions.

51 5.3. A SURVEY OF TROPHIC RELATIONSHIPS OF THE LITHUANIAN NEPTICULIDAE

Th e Lithuanian Nepticulidae fauna is predominated by species mining all kinds of plants. According to the newest data, the number of these species of pygmy moths – monophagous – amounts to 48. Th e strict oligophagous, who can mine plants of two or more species from one genus, are represented by 20 Nepticulidae species. Th is group of trophic spe- cialisation prevails over the group of wide oligophagous (represented by 8 species). Th e almost all species of the latter group exclusively mine the plants from the Rosaceae family. According to reliable data, among the Lithuanian pygmy moth species only Enteucha acetosae has three generations in a year. It was established that the fauna of this country mostly is represented by univoltine species including more than 30 species from Stigmella genus. Table 5.3.1. of the dissertation gives elaborated new and revised data about the mining seasonality of Nepticulidae. Th e seasonality of caterpil- lar activity in Lithuania was determined (Fig. 5.3.1.).

Fig. 5.3.1. The determined seasonality of larval activity of the Lithuanian Nepticulidae and two activity periods.

52 The dissertation contains a thorough analysis of other specific mining features of the Lithuanian Nepticulidae (not included in the present summary due to restricted space).

5.3.1. Trophic relationships of the Lithuanian Nepticulidae It has been determined that Nepticulidae species mine host-plants from 15 families and 35 genera. Rosaceae, Salicaceae, Fagaceae and Betulaceae are mainly mined by caterpillars of pygmy mothes whereas 25 Nepticulidae species, i.e. 33% of the total Lithuanian Nepticulidae fauna, are trophically related to Quercus, Betula and Malus genera. Eleven species of the Lithuanian Nepticulidae mine fruit trees including 7 species from 3 genera (Stigmella, Bohemannia and Ectoedemia) which in different seasons mine the leaves of domestic apple tree Malus( domestica). Some Nepticulidae species (Stigmella oxyacanthella, S. plagicolella and S. prunetorum) have accommodated to mining fruit trees of a few species.

5.4. OCCURRENCE OF OCCURRENCE OF THE LITHUANIAN NEPTICULIDAE SPECIES

The evaluation of occurrence of species (common or rare) was fol- lowed by the method described in Diškus, Stonis (2012) and Stonis et al. (unpublished). Among the total of Nepticulidae species recorded in Lithuania, 14 are rather widespread and abundantly mining (e.g. Stigmella splendi- dissimella and Fomoria septembrella). The mentioned category accounts for 1/5 of the known fauna. Bohemannia pulverosella is one of the 10 species which belong to the category of sparse distribution and abundant mining. They account for 13% of the Lithuanian Nepticulidae fauna. Groups of highly limited distribution and very abundant mining (Ectoe- demia turbidella), wide distribution but not abundant mining (Stigmella floslactella), and wide distribution and extremely abundant mining (Ec- toedemia argyropeza) are represented only by single species each. The Lithuanian Nepticulidae fauna is predominated by not rare (37%) and rare (24%) species. It was established that 18 Lithuanian Nepticulidae species belong to common species: including 13 common (they account for 17% of the

53 fauna) and 5 very common (about 7%) species. The latter are especially interesting since they can be regarded as potential pests. In terms of biogeographical distribution of Nepticulidae, most of the common species (the sum of common and very common species) are characterized by Euro-Nemoral or Euro-Submediterranean distribution ranges (Fig. 5.4.1.).

Fig. 5.4.1. Belonging of the rare Lithuanian Nepticulidae species to different chorological groups.

Not abundant mining is characteristic of about half of the rare species (45%) of the Lithuanian Nepticulidae fauna: average abundance mining is characteristic of 38% of rare species and especially sporadic mining of 17% of the rare species. As distinct from the common species, the distribution of the rare species is highly limited (59%), based on solitary findings (34%) or sparse (7%). They are never widespread. The rare species mainly mine plants from Ulmus, Fagus and Betula genera. As distinct from the common species the rare species include miners trophically associated with host-plants belonging to Salix plants. Only two rare species – Stigmella atricapitella and S. samiatella (same as the common ones S. ruficapitella and S. roborella) mine leaves of Quercus robur.

54 CONCLUSIONS

General assessment of the Nepticulidae fauna 1. General assessment of the Nepticulidae fauna showed that currently 850 species have been described (47% from the Stigmella Schrank genus; 53% from the remaining 15 genera). Among the seven biogeographical regions of the Earth, the highest Nepticulidae diversity has been recorded in the Palaearctic region (52% of the world fauna). Mean- while, 10.8% and 16.5% of the world fauna have been registered in the Neotropical and Afrotropical biogeographical regions. 2. The host-plants of most of the Nepticulidae species have been identified, yet trophic relationships of 37% of the world Nepticulidae fauna are still unknown. The recently discovered new species Stigmella( acuta mining Quercus dentata; other our recently discovered Nepticuli- dae species are not included in the current dissertation) for the first time provided reliable evidence that the Eurasian S. hemargyrella group is trophically associated with the Quercus genus. 3. Taxonomic analysis and analysis of diagnostic characters showed that in the world fauna six species groups of Stigmella trophically associated with Quercus can be recognized: S. caesurifasciella, S. saginella, S. quercipulchella, S. ruficapitella, S. castanopsiella ir S. hemargyrella (all these groups can be reliably diagnosed using the original dichotomic systems). It was determined that, in terms of trophic relationships, S. quercipulchella group is the most specific one as its species exclusively mine the plants of red oaks (Quercus: section Lobatae). 4. Cladistic analysis of the species groups of Stigmella trophically associated with Quercus as a host-plant, revealed that S. quercipulchella, S. hemargyrella, S. ruficapitella and S. castanopsiella groups are closely related (their monophyly is proved by at least 4 apomorphic characters) whereas S. saginella, S. caesurifasciella and S. cornuta are three independent phylogenetic clades. However, the phylogenetic data do not coincide with the trends of geographical distribution: S. saginella and S. quercipulchella groups are of limited distribution in North and Central America whereas the remaining all groups are distributed in Eurasia (among them S. castonopsiella, S. caesurifasciella and S. cornuta groups only occur in

55 East Asia). Thus it can be assumed that the species groups of Stigmella trophically associated with Quercus and occurring in the Eastern and Western Hemispheres (e.g. in America) do not share the same origin and represent different evolutionary tendencies.

The Crimean Nepticulidae within the context of the European fauna 5. Taking into consideration the recent taxonomic changes and the data on the newly described species it can be stated that currently the European Nepticulidae fauna is comprises of 264 species which are trophically associated with 97, mostly woody (63%), host-plant genera and 34 plant families (mostly Rosaceae, Fagaceae and Fabaceae, in a lesser degree Betulaceae, Salicaceae, Lamiaceae, Cistaceae, Rhamnaceae, Ulmaceae, Corylaceae, Apiaceae and Anacardiaceae; meanwhile, only solitary (4–5, usually 1–2) Nepticulidae species are trophically associated with other families of host-plants). The trophic relationships of some Eu- ropean Nepticulidae (11%) are unknown. 6. Chorological analysis of the European fauna revealed that the Nepticulidae species of this region belong to 10 chorological groups: 28% Mediterranean, 21% Euro-Submediterranean, 12% Euro-Mediterranean, 10% Euro-Nemoral, 8% West-Palaearctic, 7% Sub-Mediterranean and Trans-Palaearctic and 1% Euro-Siberian and Holarctic each. Analysis of the faunas of different European ecoregions showed that the greatest number (194) of Nepticulidae species is known from the Mediterranean ecoregion. Simplimorpha and Glaucolepis genera (93–100% of species) demonstrate an especially close links with the Mediterranean. The link of Acalyptri, Ectoedemia and Parafomoria to this region is somewhat weaker (85–89% of the “European” species occur in the Mediterranean ecoregion). 7. The material collected during intensive our fieldworks and the material collected by other researchers helped to identify 64 species in the Crimean fauna (40% of this fauna was for the first time identified by the author of the present dissertation). The occurrence of a few species – Stigmella szoecsiella, Ectoedemia albifasciella and E. pubescivora – in the Crimea was not confirmed by reliable data. For this reason, these species were excluded from the list of fauna of the region. 8. Mapping of the Crimean Nepticulidae distribution ranges and chorological data analysis showed that about 72% of species discovered

56 in the Crimea belong to the chorological groups which more or less are related with the Mediterranean ecoregion. Among the 8 chorological groups identified in the Crimean fauna, the species of the Euro-Submed- iterranean distribution range (36% of the known fauna of the peninsula) are dominant. Approximately one fifth of the species (22%) belongs to the Euro-Mediterranean chorological group and one sixth (16%) to the West-Palaearctic group. Meanwhile, the especially wide Euro-Siberian or Trans-Palaearctic ranges are characteristic only of 1% and 5% of species identified in the Crimea respectively. 9. Assessment of the occurrence of the species registered in the Crimean fauna allowed determining the common and rare species of the region; most of them are widespread in the peninsula and characterized by abundant (19% of species) and very abundant (17%) mining. However, among the analysed Crimean fauna there are many species which are characterized by limited distribution and abundant mining (12%) or limited distribution and not abundant mining (12%). 10. The majority of the common Crimean species mines Quercus, Sorbus, Crataegus and Prunus plants and mainly belong to the Euro-Sub- mediterranean chorological group (47%). Meanwhile, as distinct from what had been expected, the least number of common species is characterized by the Sub-Mediterranean Mediterranean distribution ranges. 11. Most of the rare species also mine Quercus (23%) as well as Lotus, Alnus and Acer (9% each) plants and are characterized by Euro-Medi- terranean (36%), Euro-Submediterranean and Euro-Nemoral (18% each) or Sub-Mediterranean (14%) distribution ranges. As could have been expected, the smallest number of rare species is characterized by wide distribution ranges or Sub-Mediterranean–Mediterranean distribution. 12. The data of trophic analysis showed that the majority of the Crimean Nepticulidae fauna is associated with the plants from the Rosaceae and Fagaceae families (18 and 12 species respectively); about 11% of the Crimean species are leaf-miners of cultivated plants (including Stigmella irregularis whose host-plant Pyrus elaeagnifolia was identified for the first time). According to the data of our fieldworks, the mining was reliably registered in the 1st decade of June to the 3rd decade of Au- gust (though it may continue to the 3rd decade of October). The peak of the larval activity is in July.

57 Taxonomic, chorological and trophic analysis of the Lithuanian Nepticulidae Fauna 13. Based on the current data, 76 Nepticulidae species are known in the Lithuanian fauna. The majority of species were discovered in the South-Eastern (65 species) and South-Western (67 species) regions of the country. The majority of the species of the Lithuanian fauna (74%) belongs to Stigmella genus; a rather big part belongs to Ectoedemia and only a few or one species (from 1% to 3%) belong to the remaining 5 genera. 14. Analysis of the distribution showed that from the chorological point of view the Lithuanian Nepticulidae fauna is not homogeneous; it belongs to 8 chorological groups. The most abundant in the Lithua- nian fauna are those of moderately limited European distribution – the Sub-Mediterranean (30%) and Euro-Nemoral (24%). 15. According to the available data, the species of the Euro-Submedi- terranean distribution constitute the major part of the Nepticulidae fauna (from 26% to 31%). Yet in the Coastal region, the number of such species is considerably lower (8%). The Coastal region is predominated by the species of the Euro-Mediterranean distribution (32%) which are less numerous in other Lithuanian regions including the Curonian Spit (they are especially few in the Curonian Spit region where they account for 4% of the regional fauna). From the chorological point of view, the Curonian Spit region is the most distinctive one among the Lithuania’s regions. It is characterized by the species of the Sub-Baltic group which do not occur in other regions of the country. The Coastal region also is rather specific: the species of the Euro-Mediterranean group are occurring in greater abundance than anywhere else in Lithuania. 16. Trophic analysis of the Lithuanian Nepticulidae fauna showed that though there are many oligophagous species in it (28 species; 37%) the majority is represented by species mining one species of host-plants (monophagous account for 63% of the Lithuanian Nepticulidae fauna). Among the oligophagous species strict oligophagous predominate (strict oligophagous account for 27% and broad oligophagous for 11%). The latter almost exclusively are leaf-miners of Rosaceae plants. The majority of Nepticulidae found in Lithuania are bivoltine species other species are univoltine (all of Ectoedemia, Glaucolepis, Bohemannia and about 34% of Stigmella species). According to our field data, Enteucha acetosae is the only polyvoltine species in the Lithuanian fauna.

58 17. During the study carried out in Lithuania, 15 families and 35 genera of host-plants were identified. Most of the Lithuanian Neptic- ulidae is trophically associated with Rosaceae, Salicaceae, Fagaceae and Betulaceae families and about 33% of the fauna is associated with Quer- cus, Betula and Malus genera. The majority of the Lithuanian Nepticul- idae species are trophically associated with woody plants (some of them are pest of cultivated/garden plants) and 17% of the fauna are miners of herbaceous plants. 18. The Lithuanian fauna is predominated by abundantly mining Nepticulidae (37 species or 49% of the total Lithuanian Nepticulidae fauna). It was determined that the most common in the Lithuanian Neptic- ulidae fauna are species of wide distribution and abundant mining (they account for more than one fifth of the total of known fauna). The smallest number is represented by species of highly limited distribution/very abundant mining or wide distribution/not abundant or sporadic mining (slightly over 1% each). 19. About 40% of the Lithuanian Nepticulidae species belong to the category of rare (rare and very rare) and 23% to the category of common species (17% of species are common and 6% very common). Nepticulidae both of common and rare species are characterized by the Euro-Nemoral and Euro-Submediterranean ranges (there are no significant chorological differences between the Lithuanian common and rare species). Yet as distinct from the common species, the rare and very rare species are characterized only by highly limited distribution (about 60% of the rare species) or almost solitary findings (34%) and, sometimes, sparse distribution (7%) yet never by wide or very wide distribution over the territory of the country.

59 SANTRAUKA LIETUVIŲ KALBA

DISERTACIJOS STRUKTŪRA IR APIMTIS

Disertacija yra parašyta lietuvių kalba. Disertaciją sudaro šie skyriai: Įvadas, Literatūros apžvalga, Metodai ir medžiaga, Tyrimo rezultatai, Išvados, Literatūros sąrašas ir 3 priedai: Vakarų Palearkties Nepticulidae faunos katalogas, Kryme aptiktų Nepticulidae rūšių genitalinių struktū- rų katalogas ir Kryme aptiktų Nepticulidae rūšių minų katalogas. Į Lite- ratūros sąrašą įtraukti ir disertacijoje pacituoti 282 šaltiniai. Disertaciją sudaro 333 puslapių; disertacijoje pateikta 10 lentelių ir 188 paveikslai (iš jų 56 disertacijos pagrindiniame tekste, o 132 prieduose).

PROBLEMA IR TYRIMŲ AKTUALUMAS

Dėl aktyvaus natūralių kraštovaizdžių naikinimo, dažnos gyvūnų ar augalų gyvenamųjų vietų fragmentacijos bei destrukcijos sparčiai ėmė nykti biologinė įvairovė. Vis didėjantis pasaulio mokslinės visuomenės susirūpinimas biologinės įvairovės krize paskatino tyrinėtojus aktyviau surašyti pagrindinius Žemės biomus (Navickaitė ir kt., 2011b). Mūsų tyrimų objektas – filogenetiškai vienas primityviausių (ir tuo, teoriniu požiūriu, vienas svarbiausių bei įdomiausių) Lepidoptera būrio taksonų (Nepticuloidea: Nepticulidae), jungiantis 850 rūšių ir 16 genčių: Pectinivalva, Roscidotoga, Simplimorpha, Enteucha, Mononeura, Areticu- lata, Stigmella, Bohemannia, Ectoedemia, Fomoria, Acalyptris, Parafomo- ria, Trifurcula, Glaucolepis, Etainia, Varius. Dauguma šių genčių yra pla- čiai paplitusios visuose žemynuose (išskyrus Antarktį) ir joms priklauso mažiausi pasaulyje mikrodrugiai, kurie išsiskiria ne tik archaiška sandara, bet ir labai didele specializacija. Dėl stenofagijos tendencijų, sėslaus gyvenimo būdo ir endemizmo šie smulkūs vabzdžiai gausiai paplitę visuose sausumos biomuose (pradedant dykumomis ir baigiant atogrąžų miškais) ir yra vieni puikiausių objektų, charakterizuojančių tiriamo biomo biologinės įvairovės turtingumą, kilmę ir gamtinius ryšius (Stonis, 2010, 2011; Navickaitė ir kt., 2011b). Tačiau šių, praktinę ir teorinę reikšmę turinčių, vabzdžių tyrimų duomenų (tiek Lie-

60 tuvos, tiek kitų regionų) nepakanka. Iki disertacijos autorės ir jos kolegų atliktų tyrimų daugelio Žemės regionų fauna buvo nežinoma ir neaprašyta, o duomenys apie Lietuvos Nepticuloidea rūšis reikalavo papildymo. Taigi tyrimų duomenų apie Nepticuloidea ir Tischerioidea (ypač Tischeriidae) stoka, jų svarbi praktinė bei teorinė reikšmė, didėjantis mokslinės visuomenės susirūpinimas biologinės įvairovės krize paskatino parengti šią Euronemoralinės faunos Nepticulidae reviziją; tokio pobūdžio darbų aktualumas buvo pabrėžtas dar 1992 m. Rio de Žaneiro tarptautinėje Biologinės įvairovės konvencijoje.

DARBO TIKSLAS IR UŽDAVINIAI

Darbo tikslas. Atlikus bendrąjį pasaulio ir Europos Nepticulidae faunos įvertinimą, pateikti pirmąją Krymo ir Lietuvos taksonominę, chorologinę ir trofinę charakteristiką. Darbo uždaviniai:

Bendrasis Nepticulidae faunos įvertinimas 1. Atlikus bendrąjį Nepticulidae faunos įvertinimą, nustatyti šiuo metu aprašytos faunos taksonominę sudėtį ir pasiskirstymą Žemės bio- geografiniuose regionuose; 2. Įvertinti pasaulio Nepticulidae faunos trofinių ryšių duomenis, pa- tikslinant Stigmella hemargyrella rūšių grupės trofinius ryšius; 3. Atlikti Stigmella genties rūšių grupių, trofiškai susijusių su Quer- cus, taksonominių ir diagnostinių požymių įvertinimą, pateikiant dicho- tomines rūšių grupių identifikacijos sistemas; 4. Atlikti Stigmella genties rūšių grupių, trofiškai susijusių su Quer- cus, kladistinę analizę.

Krymo Nepticulidae Europos faunos kontekste 5. Įvertinti Europos Nepticulidae taksonominę faunos sudėtį ir trofinius ryšius; 6. Atlikti Europos faunos chorologinę analizę; 7. Atlikus lauko darbus ir kitų autorių surinktos mokslinės kolekci- jos medžiagos identifikaciją, išaiškinti Krymo faunoje aptinkamas Nep- ticulidae rūšis;

61 8. Atlikus Krymo aptinkamų mažųjų gaubtagalvių chorologinę ana- lizę, nustatyti Krymo faunos priklausomybę įvairioms chorologinėms grupėms; 9. Įvertinti Krymo faunoje registruotų rūšių aptinkamumą ir išaiš- kinti dažnąsias ir retąsias rūšis; 10. Analizuoti dažnųjų Krymo rūšių minavimo ypatumus ir jų pri- klausomybę chorologinėms grupėms; 11. Analizuoti retųjų Krymo rūšių minavimo ypatumus ir jų pri- klausomybę chorologinėms grupėms; 12. Atlikti Kryme išaiškintos Nepticulidae faunos trofinę analizę bei nustatyti vikšrų minavimo periodus.

Lietuvos Nepticulidae faunos taksonominė, chorologinė ir trofinė analizė 13. Įvertinti Lietuvos mažųjų gaubtagalvių taksonominę sudėtį ir pateikti taksonominį sąvadą; 14. Atlikti Lietuvos Nepticulidae chorologinę analizę ir nustatyti chorologines grupes; 15. Palyginti atskirų Lietuvos regionų Nepticulidae faunos chorolog- inius duomenis; 16. Išaiškinti Lietuvos Nepticulidae trofines ypatybes ir nustatyti vy- raujančius gyvybinius ciklus; 17. Atlikus chorologinę analizę, nustatyti Lietuvos mažųjų gaubta- galvių mitybinių augalų šeimas ir gentis bei kitas trofinių ryšių ypatybes; 18. Vadovaujantis rūšių paplitimo ir minavimo gausumo įvertinimo metodika, nustatyti Lietuvos mažųjų gaubtagalvių minavimo gausumą; 19. Įvertinti Lietuvos faunoje registruotų rūšių aptinkamumą, išaiš- kinant dažnąsias ir retąsias rūšis, o aptinkamumo duomenis susiejant su chorologinės analizės rezultatais.

DARBO NAUJUMAS

Parengtas pirmasis vakarų Palearkties Nepticulidae faunos sistema- tinis katalogas ir patikrinta visų 850 pasaulio Nepticulidae rūšių takso- nominė pozicija; padaryta 15 naujų taksonominių kombinacijų. Kartu su bendraautoriais parengta originali Stigmella genties Qu- ercus augalus minuojančių Nepticulidae taksonominė analizė remiantis

62 naujai išaiškintais išorės morfologijos ir genitalinių požymių duomenimis. Išsamiai diagnozuotos ir trofiškai analizuotos Stigmella genties rūšių grupės. Pirmą kartą atliktas Lietuvos Nepticulidae faunos chorologinis įver- tinimas, naujais rūšių duomenimis papildytas šalies faunos taksonominis sąvadas, pirmą kartą išsamiai analizuoti Lietuvoje paplitusių rūšių trofiniai ryšiai. Atliekant nuolatinius mažųjų gaubtagalvių tiriamuosius lauko darbus Lietuvoje, pirmą kartą identifikuota Kuršių nerijos Nepticulidae fauna, kuri daugiau pasižymi išskirtinėmis euromediteraninio paplitimo rūšimis nei kontinentinė Lietuvos dalis. Pirmą kartą ištyrinėta ir preparuota iki šiol neidentifikuota Nep- ticulidae medžiaga, surinkta Kryme (Ukraina) ir saugota Rusijos MA Zoologijos institute. Pirmą kartą atlikta Krymo faunos taksonominė ir chorologinė analizė. Vykdant ekspedicinius Nepticulidae lauko darbus Karadaho rezervate bei kitose Krymo vietovėse, registruotos 64 mažųjų gaubtagalvių rūšys, iš kurių 26 (41 proc.) yra naujos Krymui; pirmą kar- tą išnagrinėta visų Kryme aptinkamų rūšių medžiaga, padaryti nauji, reikalavimus atitinkantys, aprašai bei pirmą kartą paruošti originalūs Krymo minų ir genitalinių struktūrų katalogai. Tiriant artimų euronemoralinei faunai regionų Nepticulidae, atrastos 8 naujos mokslui mažųjų gaubtagalvių rūšys (Navickaitė et al., in prep.), priklausančios Stigmella, Enteucha, Ectoedemia ir Acalyptris gentims. Kartu su bendraautoriais aprašyta ir paskelbta nauja, Stigmella hemargyrella rūšių grupei priskirta, Stigmella acuta rūšis.

TEORINĖ IR PRAKTINĖ REIKŠMĖ

Lietuvos Respublikos Seimo ratifikuota tarptautinė Rio de Žaneiro konvencija skatina vykdyti ir plėtoti įvairius biologinės įvairovės tyrimus. Pastaraisiais metais, ištobulėjus technologijoms, siekiant aprašyti kuo daugiau naujų mokslui rūšių, aktyviai pradėti tirti endobiontiniai vabzdžiai. Viena išskirtiniausių šių primityvių Microlepidoptera savy- bių – gyvenimo būdas žaliuosiuose augalo audiniuose yra vienas iš įdo- mesnių ir aktualių vabzdžių ekologijos klausimų. Disertaciniame darbe nagrinėjama Nepticuloidea antšeimiui pri- klausanti Nepticulidae šeima yra plačiai paplitusi įvairiose sausumos

63 eksosistemose, tačiau teoriniu aspektu menkai ištyrinėta. Klimato kaita, intensyvi žmonių migracija ir ekonominiais tikslais vykdomi prekybiniai projektai (ypač su egzotiniais kraštais) sudaro palankias sąlygas atsitik- tinei kenkėjų faunos introdukcijai susidaryti. Dėl minėto Nepticulidae vikšrų endobiontinio gyvenimo būdo asimiliaciniuose augalų lapų, pum- purų, stiebų, jaunos žievės arba vaisių audiniuose daugelis mūsų tirtų or- ganizmų gali būti pripažinti kenkėjais arba potencialiais kenkėjais, todėl svarbūs ūkiniu požiūriu. Manoma, kad endobiontiniai vabzdžiai-oligofa- gai geba gana lengvai adaptuotis ir kituose (naujuose) kraštuose, trofiškai susisieti su jiems nebūdingais mitybiniais augalais (Diškus, Stonis, 2012). Vykdant tyrimus Lietuvoje ir kitose Europos vietovėse pastebėta, kad didžioji dalis Nepticulidae vikšrų minuoja Rosaceae šeimos augalus. Minėta augalų šeima yra viena didžiausių pasaulyje, kuriai priskirta daugiau nei 2800 augalų rūšių ir 95 gentys. Tai leidžia daryti prielaidą, kad daugelis Rosaceae rūšių gali būti potencialūs mažųjų gaubtagalvių mitybiniai augalai. Šiuo metu pasaulyje yra registruota 850 Nepticulidae rūšių, tai sudaro apie 29 proc. Rosaceae šeimos trofinių ryšių atžvilgiu. Tačiau, žinant Nepticulidae prisitaikymą minuoti ir kitų šeimų augalus, galima ateityje tikėtis atrasti žymiai daugiau naujų rūšių mokslui, nei yra užregistruota. Ištyrus galimų (arba potencialių) kenkėjų rūšių gyvybinius ciklus, trofinius ryšius, galimų kenkėjų biologijos ypatumus bei vykdant nuo- latinį bendradarbiavimą su aplinkos apsaugos specialistais, gali būti sudaromos strategiškai veiksmingos priemonės, kurios neleistų kitų kraštų kenkėjams įsivyrauti mūsų faunoje, būtų galima efektyviai taikyti karantino priemones. Nepticulidae, kaip ir kiti primityvūs endobiontiniai Lepidoptera, yra itin parankus objetas įvairiems tyrimams įgyvendinti, todėl mūsų atlikta faunų inventorizacija gali padėti patikslinti teiginius apie evoliucinius procesus, sausumos ekosistemų biogeografiją ir kt.

GINAMIEJI DISERTACIJOS TEIGINIAI

Bendrasis Nepticulidae faunos įvertinimas 1. Dėl daugelio naujų rūšių aprašymų bei kai kurių pavadinimų sinonimizacijos, pasaulio Nepticulidae faunos taksonominė sudėtis pakito,

64 tačiau rūšys ne vienodai pasiskirsčiusios skirtinguose Žemės biogeogra- finiuose regionuose; 2. Daugumos Nepticulidae rūšių mitybiniai augalai yra jau išaiškin- ti, tačiau po naujos rūšies (Stigmella acuta) aprašymo pasikeičia S. hemargyrella rūšių grupės trofinių ryšių samprata; 3. Pasaulio faunoje gali būti pripažintos šešios Stigmella genties rūšių grupės, kurios trofiškai susijusios su Quercus: S. caesurifasciella, S. saginella, S. quercipulchella, S. ruficapitella, S. castanopsiella ir S. hemargyrella; 4. Stigmella quercipulchella, S. hemargyrella, S. ruficapitella ir S. castanopsiella grupės yra artimai giminiškos, tuo tarpu S. saginella, S. caesurifasciella ir S. cornuta yra 3 nepriklausomos filogenetinės klados. Galime teigti, kad tiek Rytų, tiek Vakarų pusrutuliuose (pavyzdžiui, Amerikoje) aptinkamos Stigmella genties rūšių grupės, trofiškai susijusios su Quercus, nėra vienos kilmės, o atstovaujamos keliomis skirtingomis evoliucinėmis šakomis;

Krymo Nepticulidae Europos faunos kontekste 5. Dėl kai kurių naujų rūšių aprašymo bei kai kurių pavadinimų sinonimizacijos, Europos Nepticulidae faunos taksonominė sudėtis ir trofinių ryšių samprata pasikeitė. Šiuo metu Europos faunoje vyrauja rūšys, kurios trofiškai susijusios su sumedėjusiais augalais, daugiausiai priklausantiems Rosaceae ir Fagaceae, mažiau Betulaceae, Salicaceae ir kitoms šeimoms; 6. Europos Nepticulidae fauna nėra vienalytė, šio regiono rūšys priklauso 10 skirtingų chorologinių grupių, iš kurių didžiausios yra medite- raninė ir eurosubmediteraninė; 7. Šiuo metu Krymo fauna jungia 64 rūšis, iš kurių 41 proc. yra pir- mą kartą išaiškintos disertacinio darbo metu; 8. Krymo Nepticulidae fauna nėra vienalytė, šio regiono rūšys priklauso 8 skirtingoms chorologinėms grupėms; didžioji dalis rūšių (apie 72 proc.) priklauso toms chorologinėms grupėms, kurios siejasi su Medi- teranine provincija (ekoregionu); tuo tarpu itin platūs, eurosibiriniai arba transpalearktiniai arealais būdingi tik 1–5 proc. Kryme aptinkamų rūšių; 9. Daugumai Krymo faunos Nepticulidae rūšių būdingas platus paplitimas pusiasalyje ir vidutiniškai gausus arba gausus minavimas;

65 10. Dauguma dažnųjų Krymo rūšių minuoja Quercus, Sorbus, Cra- taegus ir Prunus augalus ir daugiausiai priklauso eurosubmediteraninei chorologinei grupei; 11. Dauguma retųjų rūšių taip pat minuoja Quercus, o taip pat Lotus, Alnus ir Acer augalus ir daugiausiai yra euromediteraninio, eurosubmediteraninio, euronemoralinio arba submediteraninio paplitimo; 12. Daugiausiai Krymo Nepticulidae faunos yra susiję su Roasaceae ir Fagaceae šeimų augalais; apie dešimtadalis Kryme aptiktų rūšių minuoja kultūrinius augalus (tarp jų Stigmella irregularis, kurio mitybinis augalas Pyrus elaeagnifolia išaiškintas pirmą kartą);

Lietuvos Nepticulidae faunos taksonominė, chorologinė ir trofinė analizė 13. Šiuo metu Lietuvos faunoje žinomos 76 mažųjų gaubtagalvių rū- šys, kurios priklauso 7 gentims; 14. Lietuvos Nepticulidae fauna nėra vienalytė, mūsų šalies rūšys priklauso 8 skirtingoms chorologinėms grupėms, tarp kurių pačios gausiausios yra vidutiniškai riboto, europinio, paplitimo grupės – eurosu- bmediteraninė ir euronemoralinė; 15. Nors visuose Lietuvos regionuose svarbiausią mažųjų gaubta- galvių faunos dalį sudaro eurosubmediteraninio paplitimo rūšys, tačiau tokių rūšių yra daug mažiau pajūrio regione. Tarp šalies regionų chorologiniu požiūriu pats savičiausias yra Kuršių nerijos regionas, kuriam būdingos priebaltijinės grupės rūšys, kurių neaptinkama kituose šalies regionuose; 16. Tarp Lietuvos Nepticulidae yra nemažai oligofagų (37 proc.), ta- čiau daugiausiai yra monofagų (63 proc.). Dauguma Lietuvoje aptinkamų Nepticulidae yra biciklės rūšys, kitos – vienaciklės; tik viena Lietuvos faunos rūšis yra policiklė; 17. Lietuvoje mažieji gaubtagalviai minuoja augalus, priklausančius 15 mitybinių augalų šeimų ir 35 genčių (daugiausiai rūšių yra trofiškai susiję su Rosaceae, Salicaceae, Fagaceae ir Betulaceae augalais, tarp jų dauguma minuoja sumedėjusių augalų lapus); 18. Lietuvos faunoje vyrauja vidutiniškai gausus mažųjų gaubtagal- vių minavimas. Pačios gausiausios Lietuvos faunoje yra plataus paplitimo ir vidutiniškai gausaus minavimo mažųjų gaubtagalvių rūšys;

66 19. Lietuvos Nepticulidae faunoje apie 40 proc. rūšių yra priskirtinos retųjų (retų ir labai retų) rūšių kategorijai, o 23 proc. – dažnosioms rūšims. Tiek dažnųjų rūšių, tiek retųjų rūšių kategorijoms priskirtiems Nepticulidae daugiausiai būdingi euronemoraliniai arba eurosubmediteraniniai arealai (reikšmingų chorologinių skirtumų tarp Lietuvos dažnų- jų ir retųjų rūšių nėra).

DARBO APROBACIJA

Svarbiausi rezultatai, sudarantys disertacijos pagrindą, buvo pateikti svarstymui ir aprobuoti šiose konferencijose, seminaruose, mokslinių kolektyvų posėdžiuose: • Šalies mokslinė konferencija „Lietuvos biologinė įvairovė: būklė, struktūra, apsauga: „Vilkaviškio rajono mažųjų gaubtagalvių (Lepi- doptera, Nepticulidae) fauna ir trofiniai ryšiai“. Stendinis pranešimas su bendraautoriumi (Vilnius, 2008). • Konferencija „Ekosistemų vystymosi kryptys ir populiacijų fitoge- ografijos molekuliniai tyrimai Baltijos regione“: „Naujos filogene- tinės hipotezės apie augalus minuojančius primityvius Lepidoptera remiantis molekuliniais duomenimis (Lepidoptera: Nepticuloidea, Tischeriodea, Gelechiodea)“. Žodinis pranešimas su bendraautoriais (Vilnius, LMA, 2009). • Tarptautinis seminaras „Urban Ecology – Introduction: how to work with a gide Nr. 8“ • (Sonderburg, Danija, 2010). • Visuotinis Lietuvos entomologų draugijos susirinkimas: „Indijoje per didįjį musoną. Biosistematikos tyrimų grupės lauko darbai Hima- lajuose, Utarakandas, 2010“. Žodinis pranešimas su bendraautoriais (Vilnius, VU, 2010). • The V International Conference of Naturalists, „From Biotechnology to Environment Protection – interdisciplinary meeting of naturalists“: „First survey of Nepticulidae (Insecta: Lepidoptera) of the Curonian Spit (Baltic Coast of Lithuania)“. Pranešimas su bendraautoriais (Lenkija, Zielona Gora, 2010). • Šalies konferencija „Menas gyventi 2011“: „Himalajai: gamtos paslap- tys ir tyrimų pavojai“. Žodinis pranešimas (Panevėžys, 2011).

67 • VPU Gamtos mokslų fakulteto mokslinė konferencija: „Ar gretai bus pasiektas 500 naujų rūšių rekordas?“. Žodinis pranešimas su bendraautoriais (Vilnius, 2011). • LEU Biologijos katedros posėdžiuose (Vilnius, 2009, 2010, 2011, 2012, 2013, 2014). • First Baltic International Conference: „What is new and most interesting about Nepticulidae of Lithuania and the Crimea“. Pranešimas su bendraautoriumi (Vilnius, 2014). • First Baltic International Conference: „Advanced field methods of Nepticulidae studies: the leaf-mining Lepidoptera could be exciting. Pranešimas su bendraautoriais (Vilnius, 2014).

Dalyvauta Lietuvos mokslo tarybos mokslo fondo finansuojamame mokslininkų grupės projekte ,,Naujos faunos. Taksonominė analizė ir naujos mokslui endobiontinių Lepidoptera rūšys (MIP–049/2011)“ (2010– 2011). Taip pat buvo dalyvauta Lietuvos mokslo tarybos finansuojamame MTEP veiklos projekte „Lietuvos endobiontinės faunos taksonominė identifikacija“ (2012).

TYRIMŲ REZULTATŲ PUBLIKAVIMAS

Disertacijoje nagrinėjamų tyrimų rezultatai paskelbti 19 darbų, kurie išspausdinti moksliniuose žurnaluose, tęstiniuose ir vienkartiniuose leidiniuose. Iš jų 2 straipsniai paskelbti Acta Zoologica Lituanica, 1 – Eco- logy and Zoology, 2 – Zootaxa (su svorio koeficientu) moksliniuose peri- odiniuose leidiniuose ir kt. Taip pat svarbiausi tyrimų rezultatai (kartu su bendraautoriais) apibendrinti septyniuose monografijos „Lietuvos Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika“ skyriuose (Diškus, Stonis, 2012). Iš viso autorė kartu su bendraautoriais paskelbė: 8 darbus anglų kalba ir 11 darbų lietuvių kalba.

68 PADĖKOS

Nuoširdžiai dėkoju darbo vadovams doc. dr. A. Diškui ir prof. habil. dr. J. R. Stoniui už kantrybę, pastabas, nuolatinę pagalbą ir vertingus pa- tarimus ruošiant darbą. Už bendradarbiavimą, konsultacijas ir rekomen- dacijas dėkoju LEU Gamtos, matematikos ir technologijų fakulteto dar- buotojams: dekanui, prof. dr. V. Sruogai, doktorantei A. Rocienei (LEU), doktorantui A. Remeikiui (GTC). Už dalykinę pagalbą ir mokslinį bendradarbiavimą esu dėkinga Lie- tuvos ir užsienio kolegoms: dr. P. Ivinskiui (GTC), K. R. Tuck (BMNH, Londonas), O. Karsholt (ZMUC, Kopenhaga), dr. E. J. van Nieukerken (NNM, Leidenas), dr. D. R. Davis (USNM, Vašingtonas), dr. DSc. Sergey Yu. Sinev (ZIN, Sankt Peterburgas), dr. Yu. I. Budashkin (KRUAS, Kara- dahas, Ukraina), prof. dr. S. Podėnui (GTC, Vilnius). Už bendradarbiavimą tiriant Lietuvos (arba kaimyninių kraštų) Nepticulidae fauną disertacijos autorė dėkoja dr. B. Tamutei (Vytauto Didžiojo universitetas ir Kauno Tado Ivanausko zoologijos muziejus, Kaunas), tyrėjams E. Anisimovui, V. Jukchnevič, S. Brusokaitei, K. Lens- bergaitei, G. Varačinskui, J. Čeputytei, E. Poškaitei, V. Zeleniūtei, D. Ar- žuolaitytei, V. Dobryninai, J. Prakapavičiūtei, S. Šutaitei, R. Mankutei, A. Bobnis, J. Vertelytei, M. Adomaičiui. Už pagalbą išduodant leidimus tyrimų medžiagos mainams su Rusi- jos MA ir Ukrainos MA dėkoju D. Mikelaičiui (Aplinkos apsaugos agen- tūra, Gyvūnijos naudojimo kontrolės skyrius). Už suteiktą galimybę atlikti Nepticulidae tyrimus Vištyčio regioninio parko teritorijoje dėkoju direktoriui Nerijui Paškauskui (Vištytis). Už geranorišką bendradarbiavimą atliekant tyrimus bei ruošiant informaciją esu dėkinga Vištyčio regioninio parko vyr. specialistui ekologui Vaidui Leonavičiui (Vištytis) ir vyr. specialistui kraštotvarkininkui Andriui Vaičaičiui (Kybartai). Už monografijos „Lietuvos Nepticulidae faunos taksonominė, cho- rologinė ir trofinė charakteristika“ (kaip skyrių bendraautorė) dėkoju pirminio rankraščio recenzentams dr. A. Petrašiūnui (VU, Vilnius) ir dr. J. Bašilovai (VU, Vilnius), o už paruošimus spaudai – I. Oranskytei ir K. Saveljevai („Lututė“, Kaunas) bei T. Auškalniui (Lietuvos nacionalinė Martyno Mažvydo biblioteka, Vilnius).

69 Taip pat disertacijos autorė dėkoja LEU Mokslo fondui ir Lietuvos mokslo tarybos Mokslo fondui už paramą vykdant šį projektą. Už lietuviškų mažųjų gaubtagalvių (Nepticulidae) vardų bei su en- dobiontiniais vabzdžiais susijusių terminų aiškinimus esu dėkinga Ter- minologijos centro vadovei dr. A. Auksoriūtei (Lietuvių kalbos institutas, Vilnius). Taip pat esu labai dėkinga Ritai ir Kęstučiui Navickams, O. Žuliu- vienei (Vilkaviškis) ir Svajūnui Krivickui už palaikymą ir supratingumą. Taip pat dėkoju Andriui Navickui (Kaunas) už pagalbą ruošiant metodi- nes priemones. Už doktorantūros studijų metu išreikštą supratingumą, palaikymą ir nuolatinę pagalbą nuoširdžiai dėkoju Vilniaus Žemynos gimnazijos direktorei Rūtai Krasauskienei ir administracijai (Romaldai Čižienei, Al- donai Šventickienei, Rasai Kazlauskienei, Šarūnei Saikauskienei, Reginai Kryžiuvienei) bei gamtos mokslų metodinės grupės kolegėms Gražinai Drebickienei, Evelinai Macijauskienei, Danguolei Miliauskienei, Anai Lavrinovič ir Astai Valčiukienei.

70 TYRIMŲ REZULTATAI (IŠVADOS)

Bendrasis Nepticulidae faunos įvertinimas 1. Atlikus bendrąjį Nepticulidae faunos įvertinimą, nustatyta, kad šiuo metu aprašyta 850 rūšių (tarp kurių 47 proc. priklauso Stigmella Schrank genčiai; likusios priklauso kitoms 15 genčių). Tarp 7 Žemės biogeografinių regionų, didžiausia Nepticulidae rūšių įvairovė nustatyta Palearkties regione (52 proc. pasaulio faunos), tuo tarpu Neotropiniame ir Afrotropiniame regionuose šiuo metu registruota 10,8 ir 16,5 proc. pasaulio faunos. 2. Daugumos Nepticulidae rūšių mitybiniai augalai yra jau išaiš- kinti, tuo tarpu 37 proc. pasaulio Nepticulidae faunos trofiniai ryšiai yra nežinomi. Naujos mokslui rūšies atradimas (Stigmella acuta minuojanti Quercus dentata; kitos mūsų atrastos naujos mokslui Nepticulidae rūšys disertacijoje yra neaprašomos) pirmą kartą pateikė duomenų, kad Eura- zijoje paplitusi S. hemargyrella grupė yra trofiškai susijusi ir su Quercus gentimi. 3. Taksonominė ir diagnostinių požymių analizė parodė, kad pasaulio faunoje gali būti pripažintos šešios Stigmella genties rūšių grupės, kurios trofiškai susijusios suQuercus : S. caesurifasciella, S. saginella, S. quercipulchella, S. ruficapitella, S. castanopsiella ir S. hemargyrella (visos šios grupės gali būti patikimai diagnozuotos pasinaudojant origina- liomis dichotominėmis sistemomis). Nustatyta, kad trofiniu požiūriu pati specifiškiausia yra S. quercipulchella rūšių grupė, kurios rūšys minuoja išimtinai raudonųjų ąžuolų sekcijos (Quercus: Lobatae) augalus. 4. Kladistinė Stigmella genties rūšių grupių, trofiškai susijusių su Quercus, analizė parodė, kad S. quercipulchella, S. hemargyrella, S. ruficapitella ir S. castanopsiella grupės yra artimai giminiškos (jų monofiliją paremia mažų mažiausiai 4 apomorfiniai požymiai), tuo tarpuS. saginella, S. caesurifasciella ir S. cornuta yra 3 nepriklausomos filogenetinės klados. Tačiau filogenetiniai duomenys nesutampa su geografinio paplitimo tendencijomis: S. saginella ir S. quercipulchella grupės yra riboto paplitimo Šiaurės ir Centrinėje Amerikoje, o visoms likusioms rūšių grupėms būdingas paplitimas Eurazijoje (iš jų S. castonopsiella, S. caesurifasciella ir S. cornuta grupės aptinkamos tik Rytų Azijoje). Taigi galime teigti, kad tiek Rytų, tiek Vakarų pusrutuliuose (pavyzdžiui, Amerikoje) ap-

71 tinkamos Stigmella genties rūšių grupės, trofiškai susijusios suQuercus , nėra vienos kilmės, o atstovaujamos keliomis skirtingomis evoliucinėmis šakomis.

Krymo Nepticulidae Europos faunos kontekste 5. Įvertinus pastarųjų metų taksonominius pakitimus ir naujų rūšių aprašymų duomenis, išaiškinta, kad šiuo metu Europos Nepticulidae fau- ną sudaro 264 rūšys, kurios trofiškai susijusios su 97 mitybinių, daugiausiai sumedėjusių (63 proc.) augalų gentimis ir 34 mitybinių augalų šeimo- mis (daugiausiai – Rosaceae, Fagaceae ir Fabaceae, mažiau su Betulaceae, Salicaceae, Lamiaceae, Cistaceae, Rhamnaceae, Ulmaceae, Corylaceae, Apiaceae ir Anacardiaceae, tuo tarpu su likusiomis mitybinių augalų šeimomis trofiškai susijusios tik pavienės (4–5, dažniausiai 1–2) Nepticu- lidae rūšys; nedidelės Europos Nepticulidae dalies (11 proc.) mitybiniai ryšiai yra nežinomi. 6. Chorologinė Europos faunos analizė parodė, kad šio regiono Nepticulidae rūšys priklauso 10 chorologinių grupių: 28 proc. – mediteraninei, 21 proc. – eurosubmediteraninei, 12 proc. – euromediteraninei, 10 proc. – euronemoralinei, 8 proc. – vakarų palearktinei, 7 proc. – submediteraninei-mediteraninei, po 6 proc. – submediteraninei ir transpalearktinei ir po 1 proc. eurosibirinei ir transpalearktinei. Analizuojant atskirų Europos provincijų (ekoregionų) faunas, nustatyta, kad daugiausiai Nepticulidae rūšių (194) aptinkama Mediteraninėje provincijoje; ypač didelį faunos prieraišumą Mediteraninei provincijai demonstruoja Simplimorpha ir Glaucolepis gentys (93–100 proc. rūšių), kiek mažiau – Acalyptris, Ectoedemia ir Parafomoria gentys (85–89 proc. šių genčių „europinių“ rūšių aptinkama Mediteraninėje provincijoje). 7. Atlikus intensyvius lauko darbus ir ištyrus kitų tyrėjų surinktą mokslinę kolekcinę medžiagą, Krymo faunoje aptiktos 64 rūšys (40 proc. šios faunos yra pirmą kartą išaiškinta disertacinio darbo metu). Kelių rūšių, Stigmella szoecsiella, Ectoedemia albifasciella and E. pubescivora, aptikimas Kryme nebuvo patvirtintas jokiais patikimais tyrimų duomenimis, todėl šios rūšys išbrauktos iš regiono faunos sąrašo. 8. Krymo mažųjų gaubtagalvių rūšių arealų kartografavimas ir cho- rologinė duomenų analizė parodė, apie 72 proc. rūšių, aptiktų Kryme, priklauso toms chorologinėms grupėms, kurios daugiau ar mažiau siejasi

72 su Mediteranine provincija (ekoregionu). Tarp 8 chorologinių grupių, nustatytų Krymo faunoje, vyrauja eurosubmediteraninio paplitimo rū- šys (36 proc. išaiškintos pusiasalio faunos); maždaug penktadalis rūšių (22 proc.) priskiriamos euromediteraninei chorologinei grupei, šeštadalis (16 proc.) – vakarų palearktinei grupei. Tuo tarpu itin platūs, eurosibiriniai arba transpalearktiniai arealais būdingi tik (atitinkamai) 1 ir 5 proc. Kryme aptinkamų rūšių. 9. Įvertinus Krymo faunoje registruotų rūšių aptinkamumą, išaiš- kintos dažnosios ir retosios regiono rūšys; daugumai būdingas platus paplitimas pusiasalyje ir vidutiniškai gausus (19 proc. rūšių) arba gausus (17 proc. rūšių) minavimas. Tačiau tarp analizuotos Krymo faunos yra nemažai rūšių, kurioms būdingas ribotas paplitimas ir vidutiniškai gausus minavimas (12 proc.) arba ribotas paplitimas ir negausus minavimas (taip pat 12 proc.). 10. Dauguma dažnųjų Krymo rūšių minuoja Quercus, Sorbus, Cra- taegus ir Prunus augalus ir daugiausiai priklauso eurosubmediteraninei chorologinei grupei (47 proc.), tuo tarpu, skirtingai negu buvo tikėtasi, mažiausiai tarp dažnųjų rūšių yra tokių, kurioms būdingi submeditera- niniai-mediteraniniai arealai. 11. Dauguma retųjų rūšių taip pat minuoja Quercus (23 proc.), o taip pat Lotus, Alnus ir Acer (po 9 proc.) augalus ir yra euromediteraninio (36 proc.), eurosubmediteraninio, euronemoralinio (po 18 proc.) arba submediteraninio (14 proc.) paplitimo. Tuo tarpu, kaip buvo galima ir tikė- tis, mažiausiai tarp retųjų rūšių yra tokių, kurioms būdingi platūs arealai arba submediteraninis-mediteraninis paplitimas. 12. Trofinės analizės duomenys parodė, kad daugiausiai Krymo Nepticulidae faunos yra susiję su Roasaceae ir Fagaceae šeimų augalais (atitinkamai 18 ir 12 rūšių); apie 11 proc. Kryme aptiktų rūšių minuoja kultūrinius augalus (tarp jų Stigmella irregularis, kurio mitybinis augalas Pyrus elaeagnifolia išaiškintas pirmą kartą). Remiantis vien mūsų lauko darbų duomenimis, vikšrų minavimas patikimai registruotas nuo birže- lio I dekados iki rugpjūčio III dekados (nors gali tęstis iki spalio III dekados), tuo tarpu vikšrų aktyvumo pikas būdingas liepos mėnesiui.

73 Lietuvos Nepticulidae faunos taksonominė, chorologinė ir trofinė analizė 13. Remiantis tyrimų duomenimis, šiuo metu Lietuvos faunoje ži- nomos 76 mažųjų gaubtagalvių (Nepticulidae) rūšys; daugiausiai rūšių išaiškinta šalies pietryčių (65 mažųjų gaubtagalvių rūšys) ir pietvakarių (67 rūšys) regionuose. Dauguma Lietuvos faunos rūšių (74 proc.) priklauso Stigmella genčiai, gana didelė dalis – Ectoedemia genčiai, tik po kelias ar vieną rūšį (arba nuo 1 iki 3 proc.) – kitoms 5 gentims. 14. Arealų analizė parodė, kad chorologiniu požiūriu Lietuvos ma- žųjų gaubtagalvių (Nepticulidae) fauna nėra vienalytė; iš viso nustatytos 8 skirtingos mūsų šalies Nepticulidae chorologinės grupės. Pačios gausiausios Lietuvos faunoje yra vidutiniškai riboto, europinio, paplitimo grupės – eurosubmediteraninė (30 proc.) ir euronemoralinė (24 proc.). 15. Remiantis šiuo metu žinomais duomenimis, beveik visuose Lietuvos regionuose svarbiausią mažųjų gaubtagalvių faunos dalį (nuo 26 iki 31 proc.) sudaro eurosubmediteraninio paplitimo rūšys, tačiau tokių rūšių yra daug mažiau pajūrio regione (8 proc.). Pajūrio regione daugiausiai yra euromediteraninio paplitimo rūšių (32 proc.), kurių kituose Lietuvos regionuose (įskaitant ir Kuršių neriją) yra kur kas mažiau (ypač jų mažai Kuršių nerijos regione; jos sudaro tik 4 proc. regiono faunos). Tarp šalies regionų chorologiniu požiūriu pats savičiausias yra Kuršių nerijos regionas. Jam būdingos priebaltijinės grupės rūšys, kurių neaptinkama kituose šalies regionuose. Palyginti savitas ir pajūrio regionas, kuriame, kaip niekur kitur Lietuvoje, aptinkama daug euromed- iteraninės faunos. 16. Trofinė Lietuvos Nepticulidae faunos analizė parodė, kad nors mūsų šalies faunoje yra nemažai oligofagų (28 rūšys, 37 proc.), tačiau daugiausiai yra rūšių, kurios minuoja tik vienos rūšies augalus (mono- fagai sudaro 63 proc. šalies mažųjų gaubtagalvių faunos). Tarp oligofagų vyrauja siauresnės mitybinės specializacijos rūšys: siaurųjų oligofagų – 27 proc., plačiųjų oligofagų – 11 proc. Pastarieji mūsų šalyje beveik iš- imtinai yra Rosaceae augalų minuotojai. Dauguma Lietuvoje aptinkamų Nepticulidae yra biciklės rūšys, kitos (visos Ectoedemia, Glaucolepis, Bo- hemannia ir apie 34 proc. Stigmella rūšių) – vienaciklės; Lietuvos faunoje vienintelė Enteucha acetosae, remiantis mūsų lauko tyrimų duomenimis, yra policiklė rūšis.

74 17. Lietuvoje atliktų tyrimų metu nustatyta 15 mažųjų gaubtagalvių mitybinių augalų šeimų ir 35 mitybinių augalų gentys. Daugiausiai Lietu- vos mažųjų gaubtagalvių rūšių yra trofiškai susiję su Rosaceae, Salicace- ae, Fagaceae ir Betulaceae augalais, o su Quercus, Betula ir Malus gentimis – apie 33 proc. faunos. Daugiausiai šalies faunoje aptinkamų mažųjų gaubtagalvių rūšių yra trofiškai susijusios su sumedėjusiais augalais (dalis jų – kultūrinių/sodo augalų kenkėjai), o apie 17 proc. faunos yra žolinių augalų minuotojai. 18. Lietuvos faunoje vyrauja vidutiniškai gausus mažųjų gaubtagal- vių minavimas (tokių rūšių Lietuvoje yra 37, t. y. apie 49 proc. visos mūsų šalies Nepticulidae faunos). Vadovaujantis rūšių paplitimo ir minavimo gausumo vertinimais nustatyta, kad pačios gausiausios Lietuvos faunoje yra plataus paplitimo ir vidutiniškai gausaus minavimo mažųjų gaub- tagalvių rūšys (jos sudaro daugiau nei penktadalį šiuo metu žinomos faunos); mažiausiai yra lokalaus paplitimo, bet gausaus minavimo arba plataus paplitimo ir negausaus arba itin gausaus minavimo (po kiek daugiau nei 1 proc.). 19. Lietuvos Nepticulidae faunoje 40 proc. rūšių yra priskirtinos retųjų (retų ir labai retų) rūšių kategorijai, o 23 proc. dažnosioms rūšims (17 proc. yra dažnos, o apie 6 proc. – labai dažnos). Tiek dažnųjų rūšių, tiek retųjų rūšių kategorijoms priskirtiems Nepticulidae daugiausiai būdingi euronemoraliniai arba eurosubmediteraniniai arealai (reikš- mingų chorologinių skirtumų tarp Lietuvos dažnųjų ir retųjų rūšių neaptikta). Tačiau, skirtingai negu dažnosioms rūšims, retoms ir labai retoms rūšims būdingas tik lokalus paplitimas šalyje (apie 60 proc. retųjų rūšių faunos) arba labai lokalus paplitimas (34 proc.), tik kartais ribotas (7 proc.), bet niekada platus arba labai platus paplitimas šalies teritorijoje.

75 LIST OF PUBLICATIONS CONTAINING MATERIALS OF THE DISSERTATION

CHAPTERS OF MONOGRAPH 1. Stonis, J. R., Diškus, A., Navickaitė, A. 2012. Nepticulidae tyrimų metodai. In: Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabz- džiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Lututė, Kaunas. 38–59 p. 2. Stonis, J. R., Diškus, A., Navickaitė, A., Remeikis, A., Rocienė, A. 2012. Morfologiniai Nepticulidae požymiai, naudojami identi- fikuojant taksonus ir paremiantys šiuolaikinę šeimos sistemą.In : Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticuli- dae faunos taksonominė, chorologinė ir trofinė charakteristika. Lutu- tė, Kaunas. 60–72 p. 3. Diškus, A., Navickaitė, A., Stonis, J. R. 2012. Taksonominė Lietuvos Nepticulidae sudėtis ir papildytas faunos sąvadas. In: Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Lututė, Kaunas. 73–100 p. 4. Navickaitė, A., Stonis, J. R. 2012. Chorologinis Lietuvos Nepticulidae faunos įvertinimas. In: Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofi- nė charakteristika. Lututė, Kaunas. 101–111 p. 5. Diškus, A., Navickaitė, A., Stonis, J. R. 2012. Lietuvos Nepticulidae trofinė apžvalga.In : Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Lututė, Kaunas. 112–126 p. 6. Diškus, A., Navickaitė, A., Stonis, J. R. 2012. Nepticulidae rūšių api- būdinimas pagal gyvybinės veiklos pėdsakus – minas. In: Diškus, A., Stonis, J. R. Lietuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Lututė, Kaunas. 127–167 p. 7. Stonis, J. R., Navickaitė, A., Diškus, A. 2012. Lietuvos mažųjų gaub- tagalvių rūšių aptinkamumo grupės. In: Diškus, A., Stonis, J. R. Lie- tuvos endobiontiniai vabzdžiai. Nepticulidae faunos taksonominė, chorologinė ir trofinė charakteristika. Lututė, Kaunas. 168–174 p.

76 PAPERS IN SCIENTIFIC PERIODICAL JOURNALS 8. Stonis, J. R., Diškus, A., Remeikis, A., Navickaitė, A., Rocienė, A. 2013. Description of new species of oak leaf-miners (Lepidoptera: Nepticulidae), with notes on the species groups of Stigmella Schrank associated with Quercus as a host-plant. Zootaxa, 3737 (3): 201–222. ISSN 1175-5326 (print), ISSN 1175-5334 (online). 9. Navickaitė, A., Diškus A., Stonis J. R. 2014. An updated checklist of Nepticulidae (Lepidoptera) of the Crimea, Sub-Mediterranean SE Europe. Zootaxa: accepted. 10. Navickaitė, A., Diškus, A., Stonis, J. R., Dobrynina, V. 2011. Taxo- nomic catalogue of the world Nepticuloidea and Tischerioidea (Lepi- doptera) described by members of the Biosystematics Research Gro- up (Lithuania) up to 2009. Acta Zoologica Lituanica, 21 (2): 113–132. ISSN 1648-6919 11. Navickaitė, A., Diškus, A., Stonis, J. R. 2011. Review of Nepticulidae (Insecta: Lepidoptera) occurring in the Curonian Spit (Baltic Coast of Lithuania). Acta Zoologica Lituanica, 21 (3): 221–231. ISSN 1648- 6919 12. Diškus, A., Navickaitė, A., Stonis, J. R. 2011. New records of four rare species of Nepticulidae (Lepidoptera) discovered in western Lithuania. New and Rare for Lithuania Insect Species. Records and descriptions, 23: 57–60. ISSN 1648-8555 13. Stonis, J. R., Navickaitė, A., Rocienė, A., Remeikis, A., Diškus, A. 2013. A provisional checklist of the Nepticulidae (Insecta, Lepidop- tera) of the Crimea. Zoology and Ecology, 23 (1): 20–28. ISSN 2165- 8005 (print), ISSN 2165-8013 (online)

PAPERS AND ABSTRACTS OF SCIENTIFIC CONFERENCES 17. Navickaitė, A., Diškus, A., Stonis, J. R. 2010. First survey of Nepti- culidae (Insecta: Lepidoptera) of the Curonian Spit (Baltic Coast of Lithuania). V International Conference of Naturalists “From Biotech- nology to Environment Protection”. 31 p. University of Zielona Gora Publishers, Zielona Gora. 18. Stonis, J. R., Remeikis, A., Diškus, A. & Navickaitė, A. 2014. Field methods of Nepticulidae research: the studies leaf-mining Lepidop- tera could be exciting. In: Stonis, J. R., Diškus, A., Valainis, U., Jac-

77 kowski, J., Auškalnis, T. & Jurkonytė, A. (eds). Abstracts and papers of the First Baltic International Conference on Field Entomology and Faunistics. Edukologija Publishers, Vilnius. 19. Navickaitė, A., Diškus, A. & Stonis, J. R. 2014. What is new and most interesting about the Nepticulidae of Lithuania and the Crimea. In: Stonis, J. R., Diškus, A., Valainis, U., Jackowski, J., Auškalnis, T. & Jurkonytė, A. (eds). Abstracts and papers of the First Baltic Interna- tional Conference on Field Entomology and Faunistics. Edukologija Publishers, Vilnius.

OTHER PUBLICATIONS 20. Navickaitė, A., Diškus, A. 2008. Vilkaviškio rajono mažųjų gaub- tagalvių (Lepidoptera, Nepticulidae) fauna ir trofiniai ryšiai [Nepti- culidae (Lepidoptera) of Vilkaviškis district (Lithuania): fauna and trophic relationships]. Lietuvos biologinė įvairovė (būklė, struktūra, apsauga), 3: 79–85. ISSN 1822-2781 21. Stonis, J. R., Diškus, A., Navickaitė, A., Dobrynina, V. 2011. Naujų faunų paieškos nuo Baltijos iki Ramiojo vandenyno krantų. Ar grei- tai bus pasiektas 500 naujų rūšių rekordas? Mokslas ir gyvenimas, 5/6: 48–51. ISSN 0134-3084 22. Stonis, J. R., Diškus, A., Navickaitė, A. 2012. Lauko darbai ant Vezu- vijaus ugnikalnio šlaito. Žurnalas apie gamtą, 5: 17–19. ISSN 1648- 8938

78 CURRICULUM VITAE

Name: Asta Navickaitė

Date and place of birth: 9 November 1983, Vilkaviškis District, Lithuania

E-mail: [email protected]

Education: 2003–2007 B.Sc., Biology, Lithuanian University of Educational Sciences 2007–2009 M.Sc., Biology, Lithuanian University of Educational Sciences 2009–2014 Ph.D. fellow, Department of Biology, Faculty of Science and Technology, Lithuanian University of Educational Sciences

Research visits abroad and fieldwork: December, 2009: the Zoological Institute of the Russian Academy of sciences, St. Petersburg August, 2010: Uttarakhand State, India July–August, 2011: Karadag Nature Reserve, Ukraine October, 2012: Campania Region, Italy October, 2013: Lombardy Region, Italy

Employment: 2008 – up to date, Biology teacher, Vilnius Žemyna Gymnasium

79 Asta Navickaitė

TAXONOMY, CHOROLOGICAL COMPOSITION AND TROPHIC RELATIONSHIPS OF THE EURONEMORAL FAUNA OF NEPTICULIDAE (INSECTA, LEPIDOPTERA)

Summary of Doctoral Dissertation Biomedical Sciences, Biology (01 B): Entomology and Plant Parasitology, B–250

Maketavo Laura Petrauskienė

SL 605. 5 sp. l. Tir. 75 egz. Užsak. Nr. 14- Išleido ir spausdino Lietuvos edukologijos universiteto leidykla T. Ševčenkos g. 31, LT-03111 Vilnius Tel. +370 5 233 3593, el. p. [email protected]