Arūnas Diškus - VPU biblioteka - Vilniaus pedagoginis universitetas

VILNIUS UNIVERSITY
Arûnas Diðkus
THE NEPTICULOIDEA & TISCHERIOIDEA:
STRATEGIC REGIONAL REVISIONS
WITH A GLOBAL REVIEW
(INSECTA: LEPIDOPTERA)
Summary of doctoral dissertation
Biomedical Sciences, Zoology (05B)
Vilnius, 2005
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The work was carried out in 1990–2004 at Vilnius Pedagogical University
Consultant:
Prof. Habil. Dr. Sigitas Podënas (Vilnius University, Biomedical Sciences,
Zoology – 05B)
The defence of the doctoral dissertation is held at Vilnius University Zoology
science council
Chairman:
Prof. Habil. Dr. Rimantas Rakauskas (Vilnius University, Biomedical Sciences,
Zoology – 05B)
Members:
Fellow Lith. Acad. Sci., Habil. Dr. Vytautas Kontrimavièius (Institute of Ecology
of Vilnius University, Biomedical Sciences, Zoology – 05B)
Prof. Habil. Dr. Sigitas Podënas (Vilnius University, Biomedical Sciences,
Zoology – 05B)
Prof. Dr. Sergejus Oleninas (Klaipëda University, Biomedical Sciences, Ecology
and Environmental Sciences – 03B)
Doc. Dr. Algimantas Paulauskas (Vytautas Magnus University, Biomedical
Sciences, Ecology and Environmental Sciences – 03B)
Opponents:
Habil. Dr. Vincas Bûda (Institute of Ecology of Vilnius University, Biomedical
Sciences, Ecology and Environmental Sciences – 03B)
Prof. Dr. Remigijus Noreika (Vilnius Pedagogical University, Biomedical
Sciences, Zoology – 05B)
The official defence of the dissertation will take place on April 22, 2005, at 14
p.m. in the Faculty of Natural Sciences of Vilnius Pedagogical University, in the
Auditorium A 11
Address: Studentø str. 39, Vilnius, LT–08106, Lithuania
The abstract of doctoral dissertation was sent on March 21, 2005
The dissertation is available in the libraries of Vilnius University and Institute
of Ecology of Vilnius University

VILNIAUS UNIVERSITETAS
Arûnas Diðkus
NEPTICULOIDEA IR TISCHERIOIDEA:
TAKSONOMINËS STRATEGINIØ REGIONØ REVIZIJOS
IR PASAULIO FAUNOS APÞVALGA
(INSECTA: LEPIDOPTERA)
Daktaro disertacijos santrauka
Biomedicinos mokslai, zoologija (05B)
Vilnius, 2005
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Disertacija parengta 1990–2004 metais Vilniaus pedagoginiame universitete.
Disertacija ginama eksternu.
Mokslinis konsultantas
prof. habil. dr. Sigitas Podënas (Vilniaus universitetas, biomedicinos mokslai,
zoologija – 05B)
Disertacija ginama Vilniaus universiteto Zoologijos mokslo krypties taryboje:
Pirmininkas
prof. habil. dr. Rimantas Rakauskas (Vilniaus universitetas, biomedicinos
mokslai, zoologija – 05B)
Nariai:
akad. habil. dr. Vytautas Kontrimavièius (Vilniaus universiteto Ekologijos
institutas, biomedicinos mokslai, zoologija – 05B)
prof. habil. dr. Sigitas Podënas (Vilniaus universitetas, biomedicinos mokslai,
zoologija – 05B)
prof. dr. Sergejus Oleninas (Klaipëdos universitetas, biomedicinos mokslai,
ekologija ir aplinkotyra – 03B)
doc. dr. Algimantas Paulauskas (Vytauto Didþiojo universitetas, biomedicinos
mokslai, ekologija ir aplinkotyra – 03B)
Oponentai:
habil. dr. Vincas Bûda (Vilniaus universiteto Ekologijos institutas, biomedicinos
mokslai, ekologija ir aplinkotyra – 03B)
prof. dr. Remigijus Noreika (Vilniaus pedagoginis universitetas, biomedicinos
mokslai, zoologija – 05B)
Disertacija bus ginama vieðame posëdyje, kuris ávyks 2005 m. balandþio 22 d.
14 val. Vilniaus pedagoginio universiteto Gamtos mokslø fakulteto A 11 auditorijoje.
Adresas: Studentø g. 39, Vilnius, LT–08106, Lietuva
Disertacijos santrauka iðsiuntinëta 2005 m. kovo 21 d.
Disertacijà galima perþiûrëti Vilniaus universiteto ir VU Ekologijos instituto
bibliotekose

1. INTRODUCTION
1.1. Relevance of the study. Growing international concern over the biodiversity
crisis together with the provisions of the 1992 Rio Convention on Biological
Diversity have imparted a new urgency to the need for taxonomic revisions of
diverse groups and the provision of identification manuals, particularly for tropical
areas. Hence this dissertation.
Nepticuloidea (Nepticulidae and Opostegidae) and Tischerioidea
(Tischeriidae) are very specialized, isolated superfamilies (families) of primitive
monotrysian Microlepidoptera with a worldwide distribution. The minute size of
the adults, the concealed mining life-style of the larvae (predominantly in leaves),
and the difficulty of rearing imagines goes some way towards explaining why these
moths are still poorly studied in many regions. Only the northern European
nepticuloid fauna can be considered to have been exhaustively studied (e.g.,
Johansson et al., 1990). Studies in other regions of the world fall some way short
of reflecting the actual diversity of the group although coverage compares well
with that of other groups of Microlepidoptera. The most exhaustive revisions or
detailed taxonomic papers have been published for the western Palaearctic
(Johansson, 1971; Nieukerken, 1983, 1985a, 1985b, 1986a; Johansson et al., 1990;
Nieukerken & Puplesis, 1991, etc.), Central and Eastern Palaearctic, including
Far-eastern Russia and Japan (Kemperman & Wilkinson, 1985; Puplesis, 1994;
Puplesis & Diðkus, 1995, 1996a, 1996b, 1996c; Puplesis et al, 1996, 1997), South
Africa (Scoble, 1978a, 1978b, 1980a, 1980b, 1983), Nearctic (Davis, 1978; Wilkinson,
1979, 1981; Wilkinson & Scoble, 1979; Wilkinson & Newton, 1981; Newton &
Wilkinson, 1982), Neotropics (Puplesis, Robinson, 2000; Puplesis et al., 2002),
Oriental Region (Puplesis, Robinson, 1999), New Zealand (Donner & Wilkinson,
1989), Australia (Hoare et al., 1997; Hoare, 2000) and recently – as a summary of
the currently known fauna of the world – Puplesis, Diðkus, 2003 and the present
thesis. Long neglected, the diverse Australian fauna of nepticulids is being further
studied by Robert Hoare; the Chinese and Japanese fauna is being revised by Erik
van Nieukerken (NNM); a revision of the Central Asiatic nepticulids is currently
in preparation by R. Puplesis and A. Diðkus (VPU) and a revision of the New
World opostegids is currently in a final stage of preparation by D. Davis and R.
Puplesis (NMNH, VPU).
1.2. Goal and objectives. The goal of our study were taxonomic revisions or
reviews of these phylogenetically primitive Microlepidoptera (Nepticuloidea and
Tischerioidea) and a review of the currently known fauna of the World.
The following objectives were set for achieving the main goal:
• An extensive field collecting in Lithuanian to eludidate the occuring species,
trophical relationship and seasonal peaks for nepticulid larval activity;
An extensive field collecting in Central Asia, a review of data on nepticulid
bionomics and species diversity (incl. new and published ones);
A study of collection material from Oman, India and Nepal, description of
new taxa;
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A re-assessment of the Mandshurian fauna of the Nepticulidae (East
Asia);
A review of the Nepticulidae of Central and South America, with
description of new species from the Andes and Amazon rainforest;
A study of collection material from southern Africa, with description of
new Tischeriidae taxa;
A catalogue of the world Nepticuloidea and Tischerioidea, together with
revised data on species distribution, host-plants and a review of taxa
geographical distribution and history of description;
A morphological study and review on biology of tischeriids to support a
polytypic concept of the family, together with a cladogram and new
classification of Tischeriidae.
1.3. Novelty of the study. As a result of specified field collecting of 1990–2003 in
Cetral Asia and study of a collection material from various regions (weakly studied
or fully unstudied by previous researchers) 44 new species of Nepticulidae and 36
new species of Tischeriidae are described in co-authorship with Prof. Dr. R.
Puplesis, Dr. G. S. Robinson or other colleagues (80 new species in total) (fig. 1).
The new species described fall into 9 genera (fig. 2). One new genus in tischeriids
(Astrotischeria Puplesis & Diðkus, 2003) is described. A phylogeny and new
classification of Tischeriidae is given.
Primary types of 42 species of Nepticulidae or Tischeriidae (deposited at
BMNH, ZMUC or other museums) have been examined and illustrated for the
first time.
Larvae rearing data have introduced about 66 new plant-plant species; for the
Neotropical Nepticulidae also three new host-plant genera and three plant families:
Psidium (Myrtaceae), Acalypha (Euphorbiaceae) and Rubus (Rosaceae) and a few
host-plant genera for the Afrotropical Tischeriidae.
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Species number
60
55
50
45
40
35
30
25
20
15
10
5
0
1995 1996 1997 1998 2002 2003*
sp. n.
Fig. 1. Eighty new species of Nepticulidae and Tischeriidae are described by the author in
co-authorship with other colleagues. Additionally, two new species are described a family
not belonging to the Nepticuloidea or Tischerioidea – i.e., in Bucculatricidae (Bucculatrix
multicornuta ir B. macrognathos)

The first catalogue of the world Nepticulidae or Tischeriidae is presented. A
few new species-groups are newly designated in this catalogue and 126 new
combinations are made.
Species number
25
20
15
10
5
0
Enteucha
Stigmella
Ectoedemia
Fomoria
Acalyptris
Etainia
Tischeria*
Astrotischeria*
Coptotriche*
sp. n.
Fig. 2. The new species described in the thesis fall into 9 genera (* – genera of the Tischeriidae
(Tischerioidea); all remaining genera belong to the Nepticulidae (Nepticuloidea))
1.4. Significance of the results. One of the most fundamental challenges for
mankind in the 21 st century is to document the extent and distribution of global
biodiversity, and to understand the ecological processes that generate and maintain
it. Such information will be essential to inform and guide efforts to safeguard the
natural ecosystems that provide earth’s life support systems in the face of escalating
threats from human habitat destruction and modification.
The conducted studies have significantly broadened knowledge of biodiversity
of leaf-mining moths. The results of the study can provide the wherewithal for
users to identify representatives of groups for which there was previously no
‘entry-point’. The present study is noteworthy, since few have studied these tiny
moths, yet many species remain to be discovered and described. Without the
baseline data of providing names and means of identification for the species in a
region, no one can go further and properly plan their conservation in the case of
endemic species, or their control in the case of introduced species damaging
endemic vegetation, or recognise if a species is newly introduced and a potential
pest. Many species of Nepticuloidea and Tischerioidea are also of economic
importance.
Also leaf-mining insects (like Nepticuloidea and Tischerioidea) have proved
outstandingly well suited for inquiries into a number of fundamental problems in
functional and evolutionary ecology.
We hope that the present thesis and the reviews of regional faunas and the
currently known fauna the world will stimulate further studies of these intriguing
but much-neglected groups – Nepticuloidea and Tischerioidea.
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1.5. The main statements to be defended:
The occurrence of 73 Nepticulidae species (12 of these species are new)
are reported to the Lithuanian fauna; they are trophically related with 15
plant families and 32 genera.
Only the genera Stigmella and Fomoria (Nepticulidae) can be recognized as
having truly cosmopolitan distributions. Acalyptris, in contrary to previous
knowledge, appears also to be widely distributed and unexpectedly occurs
in rain forest habitats (in the Neotropics the genus predominates,
accounting for up to 50% of the nepticulid fauna).
World Nepticuloidea & Tischerioidea are trophically associated with 66 plant
families: most species occur on Rosaceae. In the light of host-plant data, the
evolution of the Nepticuloidea or Tischerioidea probably occurred by way of
trophic adaptation to 7 plant subclasses from the 12 recognized; four of these
host-plant subclasses are common to all three studied moth families:
Hamamelididae, Dilleniidae, Rosidae and Lamiidae.
Morphology and biology of Tischeriidae well support the polytypic concept
of the family. The recently erected Emmetia is synonymized, but a new
genus (Astrotischeria Puplesis & Diðkus) is described by us from Northern
and Southern America for tischeriid species with striking genitalia and
feeding characters.
Three main lineages of generic rank can be recognized within Tischeriidae.
The first (Tischeria) may be the best characterized by the development of a
juxta in the male genitalia, and an antrum in the female genitalia; the
second (Astrotischeria) by the development of a dorsal lobe to the valva, the
uncus overlaid dorsally by lobes of the pseuduncus, utilization of new
host-plant families (particularly Asteraceae), and the third (Coptotriche)
by the development of transtilla, spines on the diaphragma, a ‘tulip-shaped’
aedeagus and by the great enlargement of the membranous half of the
ductus spermathecae in the female genitalia.
1.6. Presentation and approval of the results. The results of the research were
presented at:
International Russian – Finnish symposium on insect biogeography (St.
Petersburg, 1993) (2 presentations, with co-authors);
IXth Congress of the European Lepidopterological Society (SEL)
(Lednice, Czech Republic, 1994);
Conference of Lithuanian Entomological Society (Vilnius, 1996);
Conference „Biodiversity in Lithuania“ (Vilnius, 1997);
Department meetings at Zoology Department of Vilnius Pedagogical
University (Vilnius, 2001 and 2003);
The Natural History Museum, London (Microlepidoptera division)
(London, 2002);
Conference „Biodiversity in Lithuania“ (Vilnius, 1997) (2 presentations);
Department meetings at Zoology Department of Vilnius University
(Vilnius, 2004 and 2005).
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The results of the doctoral thesis were published in 21 publication (incl. 8
papers in western European scientific journals). The main materials of the thesis
were published in a monograph (in co-authorship with the former supervisor):
Puplesis, Diðkus, 2003. The Nepticuloidea & Tischerioidea (Lepidoptera) – a global
review, with strategic regional revisions. 512 pp. „Lututë Publishers“, Kaunas. In
total, 11 papers published in English, 1 – in Russian, 9 – in Lithuanian language.
1.7. Structure of the thesis. The disertation includes an Introduction, Methods
and Material, Results (2 chapters and 10 sub-chapters), Conclusions, Literature
(482 references), List of papers published by the author, English Summary and 3
supplements.
In total, the presented study contains 388 pages (207 pages in the main part of
the dissertation, 181 pages in the supplements), 382 figures (109 figures in the
dissertation, 273 figures in the supplements), 16 tables. The original language of
the dissertation (and the supplement) is Lithuanian.
1.8. Acknowledgements. The work was carried out in 1990–2004 at Vilnius
Pedagogical University. The scientific supervisors of the present dissertation Prof.
Dr. Rimantas Puplesis (VPU, Vilnius) and DSc. Gaden S. Robinson (BMNH,
London) (figs 3, 4) provided the initial stimulus for the study together with
generous and enormous support during its course.
Figs 3, 4. Scientific supervisors of the present study: 3 – Prof. Dr. Rimantas Puplesis, VPU,
Vilnius (supervised the study from 1990 to 2004); 4 – DSc. Gaden S. Robinson, BMNH,
London (supervised the study from 2000 to 2003)
We also thank Dr. Don R. Davis (USNM, Washington), Prof. Dr. Niels
P. Kristensen, Ole Karsholt (ZMUC, Copenhagen), Dr. Erik van Nieukerken
(NNM, Leiden), Dr. Malcolm Scoble (BMNH, London), Cees van den Berg
(Holand), Dr. W. J. Schoorl (Amsterdam), Dr. S. Yu. Sinev, Dr. S. Baryshnikova,
Dr. A. L. Lvovskyi (ZIN, St. Petersburg), Dr. Martin Kruger (TMSA, Pretoria),
Dr. W. Mey (ZMHB, Berlin), Dr. Yu. Budashkin (Karadag, the Crimea), Dr.
Owen Lewis (Oksford), Simon R. Hill (London) and many other colleagues for
the loan of material and for providing valuable information.
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A. Diðkus thanks the trustees and the keepers of Entomology of the Natural
History Museum (BMNH) and the Zoological Museum, University of Copenhagen
(ZMUC), for study facilities and access to collections; the author is grateful to Kevin R.
Tuck (BMNH) for generous support during the course of study visits to the BMNH.
Special thanks are due to Lina Jasiukonytë and Birute Noreikienë (Vilnius)
for the Indian-ink drawings of adults and leaf-mines of some species. For other
help (incl. leaf-mine collecting) Kristina Ðarakauskienë (Alytus), Romas
Kavaliauskas (Lazdijai), Jolita Þvironienë (Vilnius) and especially Modestas Jocius
(Vilnius) are thanked.
The author is indebted to the members of the defence council, opponents
and to the colleagues from Zoology Department of Vilnius University and Institute
of Ecology of Vilnius University and particularly to the colleagues from Zoology
Department of Vilnius Pedagogical University. Special thanks are due to the
Dean of the Natural Sciences Faculty Doc. Dr. B. Ðalkus and the former Dean
Doc. Dr. A. Vilkas, Director in Chief of Finance Department of VPU Mrs Vida
Gulbinienë and many others.
Part of this study was conducted with a support given from the International
Science Fundation (USA, 1993–1994), the Prof Hering Memorial Research Fund
(England, 1993), the EU SYS-RESOURCE Programme administrated at the BMNH
(2002) and the Lithuanian State Science and Study Foundation (2002–2004).
2. METHODS AND MATERIAL
Larvae were collected and reared, and adults were collected at light. Mined
leaves (or other plant parts) were placed in Petri dishes which were then checked
regularly for emerged adults. The rearing of adults is conducted under both
natural and laboratory (indoors) conditions. For this purpose, leaves (or other
organs of a plant) are carefully checked, and infested parts with a larva are taken
and placed in a Petri dish or special container. Detailed description of the method
is given in the thesis or in published version in Puplesis, Diðkus, 2003.
Adult moths were collected by attracting them to mercury-vapour light from a
lamp suspended slightly above eye-level and 5–10 cm in front of a white screen. A
Honda EX 350 generator was used as a power-source. As many different habitats
and sites as possible were sampled. Moths attracted to the screen were collected
into small glass tubes and pinned after killing with ethyl acetate.
Genitalia were prepared following the method described by Robinson (1976).
After maceration of the abdomen in 10% KOH and subsequent cleaning, male
genital capsules were removed from the abdomen and mounted ventral side
uppermost. Where the genital armature was particularly complicated, the genitalia
were studied and sketched in glycerin before permanent mounting. The aedeagus
was removed and mounted alongside the genital armature except in the case of
some paratypes or where its removal would jeopardise study of fine structure and
where it did not obscure other sclerites. Female genitalia were removed entirely
from the abdomen, cleaned and mounted ventral side uppermost. Genitalia and
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abdominal pelts of both sexes were stained with Chlorazol Black (Direct Black 38/
Azo Black) or, occasionally, mercurochrome, and mounted in Euparal.
Fig. 5. Study and photography of tischeriid genital slides at the Zoological Museum,
University of Copenhagen (1995–1996)
Illustrations of adults were drawn in Indian ink by Mrs Lina Jasiukonytë
(VPU) using preliminary sketches and notes by the author and with additional
observations using a stereoscopic microscope. In a few cases, they could be
described as reconstructions from damaged specimens. Some species were
photographed (fig. 5).
Genitalia and wing venation drawings were made using a camera lucida, mainly
from permanent slides, but occasionally from temporary glycerin mounts (see above).
Types of all newly described species are deposited in the institution from
which the specimen was received.
This study was conducted as a part of 10 research projects of Biosystematics
Division at Zoology Department of Vilnius Pedagogical University (fig. 6) with a
financial support given from the International Science Fundation (USA, 1993–
1994), the Prof Hering Memorial Research Fund (England, 1993), the EU SYS-
RESOURCE Programme administrated at the BMNH (2002), the Lithuanian
State Science and Study Foundation (2002–2004) and other foundations.
Substantial unidentified, identified or type material was available to the author
of the dissertation, collected during the last thirty years and deposited mainly in
European and American institutions (together with material that we have collected
in Central Asia and Lithuania): BMNH – The Natural History Museum, London,
Great Britain; HNHM – Hungarian Natural History Museum, Budapest, Hungary;
VUIE – Institute of Ecology of Vilnius University, Vilnius, Lithuania; NMM –
National Museum of Namibia, Namibia; NMW – Naturhistorisches Museum Wien,
Austria; NNM – Nationaal Natuurhistorisch Museum (Naturalis), Leiden, The
Netherlands; TMSA – Transvaal Museum, Pretoria, Republic of South Africa;
USNM – National Museum of Natural History, Washington DC, USA; VPU –
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Vilnius Pedagogical University, Vilnius (incl. the material collected by Prof. Dr.
R. Puplesis in 1982–2000), Lithuania; ZIN – Zoological Institute, Russian Academy
of Sciences, St. Petersburg, Russia; ZMUC – Zoological Museum, University of
Copenhagen, Denmark; ZMHB – Zoologisches Museum der Humboldt-
Universität, Berlin, Germany.
Fig. 6. This study was conducted as a part of research projects of Biosystematics Division at
Zoology Department of Vilnius Pedagogical University in co-operation with Prof. Dr. R. Puplesis
(VPU), DSc. G. S. Robinson (BMNH), Dr. D. Davis (USNM) and other colleagues
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RESULTS AND DISCUSSION
3. STRATEGIC REGIONAL REVISIONS
3.1. REVISED FAUNA OF THE NEPTICULIDAE OF LITHUANIA
Intensive field work in Lithuania from 1996 to 2003 confirmed the occurrence
of 73 Nepticulidae species; 12 of these species are new to the Lithuanian fauna:
Stigmella nivenburgensis, S. crataegella, S. ulmivora, S. benanderella, S. continuella, S.
poterii, S. ulmariae, S. hemargyrella, Bohemannia pulverosella, Ectoedemia turbidella,
E. klimeschi, E. subbimaculella (Table 1). A further species new to the Lithuanian
fauna – Tischeria dodonea – was among the Tischeriidae.
A few species, reported in the past by other authors, have been excluded by us
from the revised list of Lithuanian nepticulids. Two species (Stigmella mespilicola
and Ectoedemia spinosella) appear to have been recorded originally from incorrectly
identified mines. We have been unable to confirm the occurance of three species
(Stigmella sakhalinella, S. sanguisorbae and Ectoedemia angulifasciella) either by
field-work or from data on material in existing collections.
The author’s research has demonstrated 15 plant families and 32 plant genera
as host-plants for Nepticulidae in Lithuania. Most of the species in the Lithuanian
fauna are trophically associated with plants from Rosaceae, Salicaceae, Fagaceae
and Betulaceae; the first – Rosaceae – strongly predominates and is the hostplant
family for 24 nepticulid species (fig. 7).
Two seasonal peaks for nepticulid larval activity (mining) were recognized:
from mid-June to mid-July (when 36 nepticulid species may be found mining),
and from mid-September to mid-October (when larvae of 49 species may be
found) (fig. 8).
Table 1. Revised list of Lithuanian Nepticulidae together with the data about nepticulid
larval activity in Lithuania
Genera and species
Period of mining
(1 – first half of a month, 2 – second half of a month)
V V VI VII VIII IX X XI
1 2 1 2 1 2 1 2 1 2 1 2 1 2 1
Enteucha l l l l l l l
1. acetosae l l l l l l l
Stigmella l l l l l l l l l l l l
2. lapponica l l
3. confusella l l l l
4. freyella l l l m
5. tiliae l l l l l
6. microtheriella l l l l l l
7. betulicola l l l l l l l
8. alnetella l l m l l
9. luteella l l l l l l
10. glutinosae l l m l l
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Genera and species
Period of mining
(1 – first half of a month, 2 – second half of a month)
V V VI VII VIII IX X XI
1 2 1 2 1 2 1 2 1 2 1 2 1 2 1
11. nivenburgensis * m m l l l
12. prunetorum l l l l
13. aceris l l l l l
14. nylandriella l l l
15. oxyacanthella l l l
16. minusculella l l l m m
17. desperatella l l l l l
18. pyri m m l l l l l
19. crataegella * l l
20. magdalenae l l l
21. hybnerella l l l l l l l l
22. anomalella l l l l l
23. catharticella l l l l l
24. malella l l l l l l l
25. viscerella l l
26. ulmivora * m l l
27. trimaculella l l l l l
28. assimilella m l l l l
29. salicis l l l l l
30. myrtillella l l l l l
31. obliquella m l l
32. zelleriella m m m l
33. benanderella * l l l m m
34. floslactella l l l l l l l
35. tityrella l l l l
36. carpinella l l l l
37. lemniscella l l l l l l l
38. continuella * l l
39. incognitella l l
40. aurella m m l
41. aeneofasciella m l l l l
42. splendidissimella l l l l
43. poterii * l l l l
44. pretiosa l l l l
45. lediella l l
46. ulmariae* l l l l
47. hemargyrella * l l l
48. lonicerarum l l l
49. plagicolella l l l l l l l l l
50. sorbi l l l l
51. atricapitella m m l
52. ruficapitella l l l l l l
53. samiatella m m l
54. basiguttella m l l l l l
55. roborella l l l l l l
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Genera and species
Period of mining
(1 – first half of a month, 2 – second half of a month)
V V VI VII VIII IX X XI
1 2 1 2 1 2 1 2 1 2 1 2 1 2 1
Bohemannia l l l l
56. pulverosella * l l l l
Ectoedemia m m m m m l l l l l l l l
57. atrifrontella m m m m m m m m m
58. argyropeza l l l l l l l l
59. turbidella * m l l l l l
60. intimella m l l l
61. hannoverella m l l l l
62. klimeschi * m l l l l l
63. subbimaculella * m l l
64. albifasciella m l l l
65. arcuatella l l l l l l
66. atricollis l l l l l
67. rubivora l l
68. occultella m l l l l l
69. minimella l l l
Fomoria l l l l l l l l l l m m
70. septembrella l l l l l l
71. weaveri l l l l m m
Glaucolepis m m
72. headleyella m m
Etainia l l l l l m m m m m
73. sericopeza l l l l l m m m m m
l – author‘s collecting data; m – predicted larval activity according other authors
research in neighbouring countries; * – species newly recorded for Lithuanian fauna
Species number
24
22
20
18
16
14
12
POLYGONACEAE
FAGACEAE
BETULACEAE
CORYLACEAE
HYPECICACEAE
ERICACEAE
SALICACEAE
TILIACEAE
ULMACEAE
ROSACEAE
ACERACEAE
RHAMNACEAE
CAPRIFOLIACEAE
CONVOLVULACEAE
LAMIACEAE
10
8
6
4
2
0
Fig. 7. Trophical relationships of the Lithuanian Nepticulidae
15

16
Species number
IV(1)
60
50
40
30
20
10
0
IV(2)
V(1)
V(2)
VI(1)
VI(2)
VII(1)
VII(2)
VIII(1)
VIII(2)
IX(1)
IX(2)
X(1)
X(2)
Fig. 8. Seasonal peaks for nepticulid larval activity (mining) in Lithuania. Dark colour –
author‘s collecting data, shown in Table 1 as black dots l; white colour – the predicted
seasonal peaks shown in Table 1 as white dots m
3.2. NEPTICULIDAE OF CENTRAL ASIA – A REVIEW OF THE FAUNA
In Central Asia, 90 nepticulid species were recognized; most of them (54
species) belong to Stigmella, 16 to Acalyptris, 11 to Ectoedemia, 4 to Fomoria, 2 to
Glaucolepis, 2 to Etainia and 1 to Trifurcula. Fifteen species were described by the
author as new to science (Table 2).
Western Kopetdag and the southern slopes of the Gissar Ridge are recognized
as the areas in the region most diverse in nepticulid species. However, about onethird
of the Kopetdag fauna is represented by species with a Euro-Asian
distribution, in contrast to the Gissar nepticulid fauna which characterized by
93% of endemic species. In total, the endemism of the Central Asiatic Nepticulidae
fauna is very high (71%) (figs 9, 10).
A review of published data on nepticulid bionomics, heavily modified updated
with the author’s original larval-rearing data, shows that about half (53%) the
Central Asiatic species with a known host-plant are trophically associated with
Rosaceae (Aflatunia, Cerasus, Crataegus, Malus, Prunus, Pyrus, Rosa, Rubus, Spiraea)
(fig. 11); 11% occur on Salicaceae (Populus, Salix), 8% on Ulmaceae (Celtis, Ulmus),
Aceraceae (Acer) or Rhamnaceae (Paliurus, Rhamnus, Ziziphus), and 6% on
Betulaceae (Betula). Plant families such as Hypericaceae, Moraceae and Fagaceae
are known as host-plant families for just a single nepticulid species.
XI(1)

Table 2. A taxonomic list of Nepticulidae of Central Asia with species distribution data: AM –
south-eastern areas of Armenia, AZ – south-eastern areas of Azerbaijan, IR – Iran (except the
southern territories), TM – Turkmenistan, UZ – Uzbekistan, TJ – Tajikistan, AF – northern
provinces of Afghanistan, KZ – western and southern parts of Kazakhstan, KG – Kyrgyzstan
Genera and species
AM AZ IR TM UZ TJ AF KZ KG
Stigmella Schrank l l l l l l m l l
1. maloidica Puplesis, 1991 l
2. paliurella (Klimesch, 1940) m l m l
3. abaiella Klimesch, 1979 l
4. ficulnea Puplesis & Krasilnikova, 1994 l l
5. turbatrix Puplesis, 1994 m l l m
6. luteella (Stainton, 1857) l
7. nivenburgensis (Preissecker, 1942) m l
8. betulifoliae Puplesis & Diškus, 2003 l
9. pamirbetulae Puplesis & Diškus, 2003 l
10. excelsa Puplesis & Diškus, 2003 l
11. semiaurea Puplesis, 1988 l l m
12. bicolor Puplesis, 1988 l l m l m
13. acerna Puplesis, 1988 m l
14. inopinata Laštuvka & Laštuvka, 1990 l
15. juryi Puplesis, 1991 m l
16. montana Puplesis, 1991 l l
17. malifoliella Puplesis, 1991 l
18. regiella (Herrich-Schäffer, 1855) l
19. crataegella (Klimesch, 1936) m m l
20. crataegi Gerasimov, 1937 m l
21. aurora Puplesis, 1984 l
22. caspica Puplesis, 1994 l
23. lanceolata Puplesis, 1994 l
24. hissariella Puplesis, 1994 l
25. hybnerella (Hübner, 1796) l m l
26. anomalella (Göze, 1783) l m
27. spinosissimae (Waters, 1928) l
28. klimeschi Puplesis, 1988 l l
29. armeniana Puplesis, 1994 l
30. kopetdagica Puplesis, 1994 m l
31. ulmiphaga (Preissecker, 1942) m l
32. kazakhstanica Puplesis, 1991 m l m l
33. muricatella (Klimesch, 1978) m m l
34. rolandi van Nieukerken, 1990 l
35. trisyllaba Puplesis, 1992 l l l
36. polymorpha Puplesis & Diškus, 2003 m l
37. fasciola Puplesis & Diškus, 2003 l
38. longispina Puplesis, 1994 l m l
39. aiderensis Puplesis, 1988 l
40. kondarai Puplesis, 1988 l
41. juratae Puplesis, 1988 l
42. flavescens Puplesis, 1994 l m l m
43. johanssoni Puplesis & Diškus, 1996 l
44. talassica Puplesis, 1992 l
45. fuscacalyptriella Puplesis, 1994 l
46. divina Puplesis, Diškus, & Nieukerken, 1997 m l
47. aurella (Fabricius, 1775) l
48. lurida Puplesis, 1994 l m m
49. motiekaitisi Puplesis, 1994 l m l
50. kuznetzovi Puplesis, 1994 l
51. subsorbi Puplesis, 1994 m l
17

Genera and species
AM AZ IR TM UZ TJ AF KZ KG
52. cerasi Puplesis & Diškus, 1996 m l
53. aflatuniae Puplesis & Diškus, 1996 l m
54. basiguttella (Heinemann, 1862) l
Ectoedemia Busck m l l l m l m l m
55. amani Svensson, 1966 l
56. albida Puplesis, 1994 m l
57. arcuatella (Herrich-Schäffer, 1855) l
58. atricollis (Stainton, 1857) l
59. spinosella (Joannis, 1908) m m l
60. ingloria Puplesis, 1988 l
61. insignata Puplesis, 1988 l
62. petrosa Puplesis, 1988 l
63. tadshikiella Puplesis, 1988 l m l m
64. rosiphila Puplesis, 1992 m l
65. albiformae Puplesis & Diškus, 2003 l
Fomoria Beirne m l m l m l m
66. septembrella (Stainton, 1849) m l m l
67. asiatica Puplesis, 1988 l m l
68. lacrimulae Puplesis & Diškus, 1996 m l
69. flavimacula Puplesis & Diškus, 1996 l m
Acalyptris Meyrick m l l l m l
70. vittatus (Puplesis, 1984) l
71. repeteki (Puplesis, 1984) l
72. lvovskyi (Puplesis, 1984) l l l m l
73. falkovitshi (Puplesis, 1984) l l m
74. turcomanicus (Puplesis, 1984) l
75. pallens (Puplesis, 1984) l l m
76. galinae (Puplesis, 1984) l l l
77. arenosus (Falkovitsh, 1986) l l m
78. shafirkanus (Puplesis, 1984) m l l l m m
79. desertellus (Puplesis, 1984) l l m
80. kizilkumi (Falkovitsh, 1986) m l l l m l
81. piculus Puplesis, 1990 l
82. brevis Puplesis, 1990 m l m l m
83. egidijui Puplesis, 1990 l l m
84. vannieukerkeni Puplesis, 1994 l
85. argyraspis Puplesis & Diškus, 1995 l m
Trifurcula Zeller m l l m l
86. puplesisi van Nieukerken, 1990 m l l m l
Glaucolepis Braun l m m l l l l l l
87. raikhonae Puplesis, 1985 l l l l l
88. melanoptera (Nieukerken & Puplesis, 1991) l m m l
Etainia Beirne m l
89. leptognathos Puplesis & Diškus, 1996 m l
90. obtusa Puplesis & Diškus, 1996 l
18

Species number
50
45
40
35
30
25
20
15
10
5
0
Armenia
Host-plants unknown
48%
Azerbaijan
Rosaceae 28%
Iran
Turkmenistan
Fig. 9. Currently known species number of nepticulids per countries of Central Asia:
Armenia (south-eastern areas only), Azerbaijan (south-eastern areas only), Iran (except the
southern territories), Turkmenistan, Uzbekistan, Tajikistan, Afghanistan (northern provinces
only), Kazakhstan (western and southern parts), Kyrgyzstan
Species number
35
30
25
20
15
10
5
0
Kopetdag Deserts &
oases
Uzbekistan
Tajikistan
Afghanistan
Fig. 11. Trophical relationships of Nepticulidae of Central Asia
Kazakhstan
Kyrgyzstan
Gissar Tyan Shan'
Asian endemic Species occuring from Asia to Europe
Fig. 10. Main natural areas of Central Asia and nepticulid species number
Salicaceae 6%
Ulmaceae 4%
Rhamnaceae 4%
Aceraceae 4%
Betulaceae 3%
Oth. families 3%
19

3.3. CONTRIBUTION TO KNOWLEDGE OF THE NEPTICULIDAE OF OMAN,
INDIA AND NEPAL
Study of material collected from Oman and the Indian subcontinent yielded
34 species of Nepticulidae. Two of them are probably boreal (occuring in the high
Himalaya) while others are tropical. Twelve species from the 34 recognized in
total are described as new to science (Table 3).
The discovery of such species as Stigmella hoplometalla and Acalyptris
melanospila, occuring on the western slopes of the Himalaya but previously known
only from Bombay suggests a much wider distribution of these species in the
Indian subcontinent.
Three closely related species (i. e., Acalyptris melanospila, A. auratilis and A.
nigripexus), found in the same habitat of Himalayan tropical montane forest and
characterized by a similar adult activity period are assumed to be of sympatric
origin.
It is interesting that the newly described Stigmella tenebrica from the high
Himalaya morphologically resembles more the European S. sorbi, than the species
of the same species-group occuring in the geographically closer territories of
Central Asia.
Table 3. Nepticulidae species in countries of South Asia: OM – Oman, IN – India, NP –
Nepal, LK – Sri Lanka
Genera and species
OM IN NP LK
Enteucha Meyrick l
1. diplocosma (Meyrick, 1921) l
Stigmella Schrank l l l l
2. isochalca (Meyrick, 1916) l
3. sruogai Puplesis & Diškus, 2003 l
4. nepali Puplesis & Diškus, 2003 l
5. omani Puplesis & Diškus, 2003 l
6. xystodes (Meyrick, 1916) l
7. liochalca (Meyrick, 1916) l
8. ipomoeella (Gustafsson, 1976) l
9. himalayai Puplesis & Diškus, 2003 l
10. fibigeri Puplesis & Diškus, 2003 l
11. elegantiae Puplesis & Diškus, 2003 l
12. maculifera Puplesis & Diškus, 2003 l
13. skulei Puplesis & Diškus, 2003 l
14. tenebrica Puplesis & Diškus, 2003 l
15. polydoxa (Meyrick, 1911) l
16. homophaea (Meyrick, 1918) l
17. argyrodoxa (Meyrick, 1918) l
18. neodora (Meyrick, 1918) l
19. aeriventris (Meyrick, 1932) l
20. hoplometalla (Meyrick, 1934) l l
21. elachistarcha (Meyrick, 1934) l
22 oligosperma (Meyrick, 1934) l
20

Genera and species
OM IN NP LK
23. longicornuta Puplesis & Diškus, 2003 l
Fomoria Beirne l
24. glycystrota (Meyrick, 1928) l
Acalyptris Meyrick l l l
25. sporadopa (Meyrick, 1911) l
26. psammophricta Meyrick, 1921 l
27. acontarcha (Meyrick, 1926) l
28. heteranthes (Meyrick, 1926) l
29. melanospila (Meyrick, 1934) l l
30. clinomochla (Meyrick, 1934) l l
31. auratilis Puplesis & Diškus, 2003 l
32. nigripexus Puplesis & Diškus, 2003 l
Glaucolepis Braun l
33. rusticula (Meyrick, 1916) l
34. „Nepticula” oritis Meyrick, 1910 l
3.4. RE-ASSESSMENT OF THE MANDSHURIAN FAUNA OF THE NEPTICULIDAE
In total, 105 species of Nepticulidae are reviewed by the author in this thesis;
63 of them were recorded only from the eastern part of the Mandshurian region
(Primorskiy Krai). One species – Stigmella auricularia – is described as new to
science (Table 4, fig. 12).
Fifteen species were recognized as distributed from Europe to the
Mandshurian region: Stigmella betulicola, S. luteella, S. sakhalinella, S. aurora, S.
anomalella, S. assimilella, S. salicis, S. obliquella, S. continuella, S. lediella, Bohemannia
quadrimaculella, Ectoedemia intimella, E. preisseckeri, E. occultella and Fomoria weaveri;
most of them are trophically associated with plants from Salicaceae (more rarely
Rosaceae and Ericaceae).
Table 4. Taxonomic list of Nepticulidae species occuring in Mandshurian region and
neighbouring countries (l – confirmed by collection data; m – occurence is predicted)
Genera and species Eastern part of
the
Mandshurian
region
(Primorskiy
Kray)
Sakhalin
and Kuril
Islands
Japan China
and
Korea
Stigmella Schrank l l l l
1. caesurifasciella Kemperman & Wilkinson, 1985 l m
2. dissona (Puplesis, 1984) l
3. mirabella (Puplesis, 1984) l
4. kurilensis Puplesis, 1987 l
5. sashai Puplesis, 1984 l
6. regina Puplesis, 1984 l
7. betulicola (Stainton, 1856) l
8. luteella (Stainton, 1857) l
9. sakhalinella Puplesis, 1984 l l
10. attenuata Puplesis, 1985 l
11. cathepostis Kemperman & Wilkinson, 1985 l
12. conchyliata Kemperman & Wilkinson, 1985 l
21

Genera and species Eastern part Sakhalin Japan China
of the and Kuril
and
Mandshurian
region
(Primorskiy
Kray)
Islands
Korea
13. oplismeniella Kemperman & Wilkinson, 1985 l
14. populnea Kemperman & Wilkinson, 1985 l
15. titivillitia Kemperman & Wilkinson, 1985 l
16. ultima Puplesis, 1984 l m
17. tegmentosella Puplesis, 1984 l m
18. monella Puplesis, 1984 l l m
19. kozlovi Puplesis, 1984 l
20. orientalis Kemperman & Wilkinson, 1985 l
21. nostrata Puplesis, 1984 l
22. aurora Puplesis, 1984 l
23. micromelis Puplesis, 1985 l
24. crataegivora Puplesis, 1985 l
25. alaurulenta Kemperman & Wilkinson, 1985 l
26. chaenomelae Kemperman & Wilkinson, 1985 l
27. honshui Kemperman & Wilkinson, 1985 l
28. sorbivora Kemperman & Wilkinson, 1985 l
29. zumii Kemperman & Wilkinson, 1985 l
30. anomalella (Göze, 1783) l
31. taigae Puplesis, 1984 l
32. kurotsubarai Kemperman & Wilkinson, 1985 l
33. palionisi Puplesis, 1984 l
34. amuriella Puplesis, 1985 l
35. alisa Puplesis, 1985 l
36. nakamurai Kemperman & Wilkinson, 1985 l
37. nireae Kemperman & Wilkinson, 1985 l
38. auricularia Puplesis, Diškus & Juchneviè, 2003 l
39. assimilella (Zeller, 1848) l
40. salicis (Stainton, 1854) l
41. obliquella (Heinemann, 1862) l
42. tranocrossa Kemperman & Wilkinson, 1985 l l
43. continuella (Stainton, 1856) l
44. gimmonella (Matsumura, 1931) l l
45. zelkoviella Kemperman & Wilkinson, 1985 l
46. lediella (Schleich, 1867) l
47. palmatae Puplesis, 1984 l
48. acrochaetia Kemperman & Wilkinson, 1985 l
49. alikurokoi Kemperman & Wilkinson, 1985 l
50. ichigoiella Kemperman & Wilkinson, 1985 l
51. sesplicata Kemperman & Wilkinson, 1985 l
52. spiculifera Kemperman & Wilkinson, 1985 l
53. oa Kemperman & Wilkinson, 1985 l
54. monticulella Puplesis, 1984 l
55. dentatae Puplesis, 1984 l l l
56. aladina Puplesis, 1984 l l l
57. omelkoi Puplesis, 1984 l l l
58. fervida Puplesis, 1984 l l
59.
22
fumida Kemperman & Wilkinson, 1985 l l

Genera and species Eastern part Sakhalin Japan China
of the and Kuril
and
Mandshurian
region
(Primorskiy
Kray)
Islands
Korea
60. clisiotophora Kemperman & Wilkinson, 1985 l m
61. circumargentea Nieukerken & Liu, 2000 l
62. kao Nieukerken & Liu, 2000 l
63. castanopsiella (Kuroko, 1978) l m
64. kurokoi Puplesis, 1984 l l m
65. lithocarpella Nieukerken & Liu, 2000 l
66. vandrieli Nieukerken & Liu, 2000 l
67. egonokii Kemperman & Wilkinson, 1985 l
68. boehmeriae Kemperman & Wilkinson, 1985 l
Bohemannia Stainton l l m
69. quadrimaculella (Boheman, 1853) m
70. manschurella Puplesis, 1984 l l m
71. ussuriella Puplesis, 1984 l
72. suiphunella Puplesis, 1984 l
73. nubila Puplesis, 1985 l l
74. piotra Puplesis, 1984 l
Ectoedemia Busck l l l
75. amani Svensson, 1966 l l
76. admiranda Puplesis, 1984 l
77. sivickisi Puplesis, 1984 l
78. laura Puplesis, 1985 l
79. intimella (Zeller, 1848) l
80. arisi Puplesis, 1984 l
81. christopheri Puplesis, 1984 l
82. philipi Puplesis, 1984 l
83. preisseckeri (Klimesch, 1941) l
84. chasanella Puplesis, 1984 l
85. scoblei Puplesis, 1984 l l
86. aligera Puplesis, 1985 l
87. ermolaevi Puplesis, 1985 l
88. maculata Puplesis, 1987 l
89. olvina Puplesis, 1984 l l l
90. ornatella Puplesis, 1984 l
91. ivinskisi Puplesis, 1984 l
92. pilosae Puplesis, 1984 l l
93. picturata Puplesis, 1985 l
94. occultella (Linnaeus, 1767) l l
95. sinevi Puplesis, 1985 l
96. insularis Puplesis, 1985 l
Fomoria Beirne l m l
97. weaveri (Stainton, 1955) m m l
98. hypericifolia Kuroko, 1982 l l
99. permira Puplesis, 1984 l
Glaucolepis Braun l
100. sinica (Yang, 1989) l
Etainia Beirne l l
101. trifasciata (Matsumura, 1931) l
23

24
Species number
70
60
50
40
30
20
10
0
Genera and species Eastern part
of the
Mandshurian
region
(Primorskiy
Primorskiy
Kray
Sakhalin &
Kuril Islands
Kray)
102. capesella (Puplesis, 1985) l
103. tigrinella (Puplesis, 1985) l
104. peterseni (Puplesis, 1985) l
105. sabina (Puplesis, 1985) l
Sakhalin
and Kuril
Islands
Japan China
Fig 12. Main areas (or countries) of East Asia and nepticulid species number; the fauna of
Korea remains unstudied
3.5. NEPTICULIDAE OF CENTRAL AND SOUTH AMERICA
Japan China
and
Korea
The recently published updated check-list of the Nepticulidae of the region (Puplesis
et al., 2002) includes 74 species; the author participated in the original description of
16 of these species together with the thesis’ supervisors Prof. Dr. R. Puplesis and Dr.
G. S. Robinson (i. e., Enteucha acuta, E. guajavae, Stigmella austroamericana, S.
montanotropica, S. nubimontana, S. rubeta, Fomoria repanda, F. tabulosa, Acalyptris
ecuadoriana, A. onorei, A. basihastatus, A. pseudohastatus, A. articulosus, A. rotundus, A.
amazonius, A. insolentis) (Table 5).
Study of Neotropical material has shown a high level of endemism of the
Nepticulidae fauna in general, and also demonstrated the phenomenon of Acalyptris
predomination: among the currently recognized species in Belize, 48% belong to
the genus Acalyptris, in the western part of the Amazon basin this genus comprises
about 50% of the fauna. Previously, Acalyptris was known with a highly restricted
distribution, that excluded the Neotropics (or, indeed, any tropical area) and also
was not known as very diverse in species.

Table 5. Taxonomic check-list of currently known Nepticulidae taxa in the Neotropical
Region with species distribution data to Central and South American countries: US –
Florida and Arizona only (USA), MX – tropical regions of Mexico, BZ – Belize, DM –
Dominica, GY – Guyana, VE – Venezuela, CO – Colombia, EC – Ecuador, PE – Peru, AR –
Argentina, CL – Chile
Genera and species
US MX BZ DM GY VE CO EC PE AR CL
Enteucha Meyrick l l l l l
1. cyanochlora Meyrick, 1915 l
2. gilvafascia (Davis, 1978) l
3. hilli Puplesis & Robinson, 2000 l
4. contracolorea Puplesis & Robinson, 2000 l
5. terricula Puplesis & Robinson, 2000 l
6. snaddoni Puplesis & Robinson, 2000 l
7. acuta Puplesis & Diškus, 2002 l
8. guajavae Puplesis & Diškus, 2002 l
Manoneura Davis l l l l l
9. basidactyla (Davis, 1978) l l l l
10. trinaria Puplesis & Robinson, 2000 l
Stigmella Schrank l l l l l l l l l
11. plumosetaeella Newton & Wilkinson, 1982 l l
12. barbata Puplesis & Robinson, 2000 l
13. austroamericana Puplesis & Diškus, 2002 l
14. kimae Puplesis & Robinson, 2000 l
15. eurydesma (Meyrick, 1915) l
16. albilamina Puplesis & Robinson, 2000 l
17. fuscilamina Puplesis & Robinson, 2000 l
18. johannis (Zeller, 1877) l
19. epicosma (Meyrick, 1915) l
20. cuprata (Meyrick, 1915) l
21. andina (Meyrick, 1915) l
22. olyritis (Meyrick, 1915) l
23. rudis Puplesis & Robinson, 2000 l l
24. marmorea Puplesis & Robinson, 2000 l
25. peruanica Puplesis & Robinson, 2000 l
26. schoorli Puplesis & Robinson, 2000 l
27. hamata Puplesis & Robinson, 2000 l
28. imperatoria Puplesis & Robinson, 2000 l
29. montanotropica Puplesis & Diškus, 2002 l
30. nubimontana Puplesis & Diškus, 2002 l
31. rubeta Puplesis & Diškus, 2002 l
32. gossypii (Forbes & Leonard, 1930) l
33. hylomaga (Meyrick, 1931) l
34. costalimai (Bourquin, 1962) l
35. guittonae (Bourquin, 1962) l
36. pruinosa Puplesis & Robinson, 2000 l
37. ovata Puplesis & Robinson, 2000 l
Ectoedemia Busck l l l
38. mesoloba Davis, 1978 l
39. reneella Wilkinson, 1981 l
40. helenella Wilkinson, 1981 l
25

26
Genera and species
US MX BZ DM GY VE CO EC PE AR CL
41. species 291058 l
42. fuscivittata Puplesis & Robinson, 2000 l l
Fomoria Beirne l l l
43. molybditis (Zeller, 1877) l
44. diskusi Puplesis & Robinson, 2000 l
45. species 29122 l
46. repanda Puplesis & Diškus, 2002 l
47. tabulosa Puplesis & Diškus, 2002 l
Acalyptris Meyrick l l l
48. latipennata (Puplesis & Robinson, 2000) l
49. dividua Puplesis & Robinson, 2000 l
50. ecuadoriana Puplesis & Diškus, 2002 l
51. onorei Puplesis & Diškus, 2002 l
52. bicornutus (Davis, 1978) l
53. tenuijuxtus (Davis, 1978) l
54. bovicorneus Puplesis & Robinson, 2000 l
55. martinheringi Puplesis & Robinson, 2000 l
56. fortis Puplesis & Robinson, 2000 l
57. hispidus Puplesis & Robinson, 2000 l
58. novenarius Puplesis & Robinson, 2000 l
59. lascuevella Puplesis & Robinson, 2000 l
60. bifidus Puplesis & Robinson, 2000 l
61. trifidus Puplesis & Robinson, 2000 l
62. unicornis Puplesis & Robinson, 2000 l
63. laxibasis Puplesis & Robinson, 2000 l
64. species 29135 l
65. platygnathos Puplesis & Robinson, 2000 l
66. species 29140 l
67. basihastatus Puplesis & Diškus, 2002 l
68. pseudohastatus Puplesis & Diškus, 2002 l
69. articulosus Puplesis & Diškus, 2002 l
70. rotundus Puplesis & Diškus, 2002 l
71. amazonius Puplesis & Diškus, 2002 l
72. insolentis Puplesis & Diškus, 2002 l
Glaucolepis Braun l l
73. aerifica (Meyrick, 1915) l
74. argentosa Puplesis & Robinson, 2000 l
3.6. CONTRIBUTION TO THE TISCHERIIDAE OF SOUTHERN AFRICA
Ten species of Tischeriidae are now known from southern Africa, i. e., the same
number as from Europe with a two hundred year history of investigation. Nine of
the ten African species were described by the author (in cooperation with Prof. Dr.
R. Puplesis and Dr. W. Mey) as new to science: Tischeria sparmanniae, T. martinkrugeri,
T. antilope, Coptotriche zimbabwiensis, C. africana, C. basipectinella; three other species
were not formerly named due to insufficient material (Table 6).
The southern African tischeriids are associated with host-plants of at least of
4 families and 6 genera: Rhamnaceae (Scutia), Tiliaceae (Grewia, Sparmannia,
Triumfeta), Anacardiaceae (Rhus) and Combretaceae (Terminalia).

Table 6. A check-list of currently known Tischeriidae taxa in southern Africa with species
distribution data to countries: NA – Namibia, ZA – Republic of South Africa, ZW – Zimbabwe
Genera and species
NA ZA ZW
Tischeria Zeller l l l
1. zestica Meyrick, 1911 l
2. antilope Puplesis, Diškus & Mey, 2003 l
3. sparmanniae Puplesis & Diškus, 2003 l l l
4. martinkrugeri Puplesis & Diškus, 2003 l
5. species 0219 l
6. species 0220 l
7. species 6547 l
Coptotriche Walsingham l l l
8. zimbabwiensis Puplesis & Diškus, 2003 m l
9. africana Puplesis & Diškus, 2003 l l m
10. basipectinella Puplesis & Diškus, 2003 m l l
27

4. A GLOBAL REVIEW OF THE NEPTICULOIDEA AND
TISCHERIOIDEA AND CATALOGUE OF THE WORLD FAUNA
After revision of all published records (including the author’s personal research
data), 1027 species of Nepticuloidea & Tischerioidea are currently recognized for
the world fauna. Most of them, 780 species, belong to the Nepticulidae, 130 to the
Opostegidae 117 to the Tischeriidae.
These species are included in the first catalogue of the world Nepticuloidea &
Tischerioidea, together with revised data on species distribution and host-plants
(see Supplements of the dissertation or the published version in Diðkus, Puplesis,
2003).
A few new species-groups are designated in the catalogue. Also 126 new
combinations are made and a few species previously omitted from taxonomic
revisions by other authors (dealing with Africa or other regions) are listed.
It is estimated that 124 authors described nepticuloids and tischerioids. Peaks
of species description occurred in 1911–1915, 1976–1980 and particularly in 1984–
1985 and 2000.
The author of this dissertation described (on his own or with co-authors) 44
new species of Nepticulidae and 36 species of Tischeriidae (figs 1, 2).
28
Australian
7%
Oriental
9%
Afrotropical
15%
Palaearctic
44%
Nearctic
14%
Neotropical
11%
Fig. 13. Currently known number of species of Nepticuloidea and Tischerioidea and
biogeographical regions
Most Nepticuloidea and Tischerioidea (445 species or about 44%) are known
from the Palaearctic (fig. 13). In the Nepticulidae, only the genera Stigmella and
Fomoria can be recognized as having truly cosmopolitan distributions. Acalyptris,
in contrary to previous knowledge, appears also to be widely distributed and
unexpectedly occurs in rain forest habitats. During this study an interesting feature
of Acalyptris distribution was discovered: in the Neotropical region the genus
predominates, accounting for up to 50% of the nepticulid fauna of the tropical
areas of the region. Otherwise, Stigmella is the dominant nepticulid genus in most
geographical regions and in non-tropical areas of the Neotropical region.

Despite very high species endemism, endemism at generic level varies from
region to region: in the Palaearctic and Afrotropical regions endemic taxa make up
20% of the fauna, in the Neotropics 17%, in Australia about 50%.
North & South America are supposed to be the centre of highest diversity of
the Tischeriidae and, possibly, their centre of origin. The Neotropical region is
remarkable in the taxonomic composition of its Tischeriidae: as well as Coptotriche
and Tischeria (genera which occur in almost every biogeographical region), there
is the third genus – Astrotischeria – which comprises 74% of the currently known
species of the area (fig. 14).
58%
62%
Coptotriche (3)
13%
Coptotriche (14)
Coptotriche (28)
Palaearctic Region
Nearctic Region
Tischeria (10)
Fig. 14. Predomination of Astrotischeria in the Neotropical Region (% and number of
species known)
9%
Neotropical Region
Tischeria (3)
13%
42%
Tischeria (4)
74%
Astrotischeria (13)
29%
Astrotischeria (17)
29

According to updated and newly reviewed host-plant data, the world
Nepticuloidea & Tischerioidea are trophically associated with 66 plant families:
most species occur on Rosaceae (123 species) and Fagaceae (93 species); to a
lesser extent, but still widely utilised, are such families as: Rhamnaceae (32 species),
Asteraceae (31 species), Fabaceae (31 species), Salicaceae (28 species) and
Betulaceae (27 species). In the light of host-plant data, the evolution of the
Nepticuloidea or Tischerioidea probably occurred by way of trophic adaptation to
7 plant subclasses from the 12 recognized; four of these host-plant subclasses are
common to all three studied moth families: Hamamelididae, Dilleniidae, Rosidae
and Lamiidae. Detailed review is given in the thesis or in published version in
Puplesis, Diðkus, 2003.
Fig. 15. Cilia-like sensilla chaetica on male antenna of Tischeria ekebladella (scale – 10 µm)
30
NEW CLASSIFICATION TO THE TISCHERIIDAE
Morphology and biology
Characters which have most diagnostic importance for family recognition are:
1. Frontal tuft projecting over triangular (or trapezoid) face smoothly covered
with scales. 2. Numerous very long, distinct, cilia-like sensilla chaetica on male
antenna (fig. 15). 3. Scales on enlarged scapus projecting as a modified pecten
over eye. 4. Strongly narrowed aedeagus, usually bifurcated or with spines at apex
(fig. 16, see the structure in the middle). 5. Dark, short, strongly thickened, stout
peg setae on female ovipositor (usually visible even without dissection) (figs 18,
19). 6. Four to five apophyses pairs in female genitalia (fig. 22). For a detailed
description and a review on biology – see the doctoral dissertation or published
version in Diðkus, Puplesis, 2003) (figs 15–23).

Figs 16, 17. Morphology of male genitalia of Tischeriidae: 16 – Astrotischeria pallens Puplesis
& Diðkus, Argentina; 17 – Coptotriche japoniella Puplesis & Diðkus, Japan (scale – 0,1 mm)
Figs 18, 19. Abdominal segments of female of Coptotriche angusticolella: 18 – lateral view;
19 – ventral view
31

Figs 20–23. Morphology of female genitalia of Tischeriidae: 20 – Tischeria ekebladoides,
antrum, Tunisia; 21 – Coptotriche berberella, ductus spermathecae, Spain; 22 – C.
heinemanni, apophyses from dorsal side, Ukraine; 23 – C. angusticolella, corpus bursae
and ductus spermathecae, Tunisia
32

Phylogeny
Morphological or trophic differences among tischeriids have prompted the
erection of a few sections or species-groups within the family. Five sections have been
designated in the Nearctic fauna by Braun (1972). These were not followed (at least
in most cases) by workers on the European fauna. Until recently all known tischeriid
species in the world were treated as belonging to a single genus (i. e., Tischeria Zeller,
1839). However, the monotypic concept of the family was contradicted by Leraut
(1993), who erected a new genus Emmetia. Our personal studies of the genitalia of
some tropical species and particularly studies of chromosomes and gonads (Puplesienë
& Puplesis, manuscript/unpublished) well support the polytypic concept of the family.
Unfortunately, Leraut (1993) did not notice that his newly erected genus completely
corresponded with Coptotriche of Walsingham (1890), a genus designated for Northern
American tischeriids, which for the last hundred years has been commonly treated as
a junior synonym of Tischeria Zeller, 1839. Further, Leraut (1993) associated the
name Coptotriche with Tischeria (s. str.), not with Emmetia, suggesting that he had no
idea what Coptotriche of Walsingham actually represented.
Tischeria complanoides Frey & Boll, 1873 was designated originally by
Walsingham (1890) as the type-species for Coptotriche; however complanoides is a
commonly recognized junior synonym of T. zelleriella Clemens, 1859. Irrespective
of whether complanoides is recognized as a separate species from zelleriella or not
(we recognize it as a synonym), the type-species of Coptotriche possesses the same
phylogenetically important characters as the species of Emmetia, including the
type-species Tischeria marginea Haworth, 1828: the broadened valva, tulip-shaped
and spined aedeagus, presence of transtilla, absence of a juxta, diaphragma with
spines, greatly extended membranous half of ductus spermathecae, etc.
Interestingly, one of most striking characters of the genus, the broadened valva,
was illustrated by Walsingham 113 years ago (Walsingham, 1890: fig. c).
The name Coptotriche can not be treated as a nomen nudum; it was used by
Braun (1972) and twice by Leraut (1993).
Therefore, the recently erected Emmetia has to be synonymized (Diðkus,
Puplesis, 2003). But a new genus (Astrotischeria Puplesis & Diðkus) is described by
us from Northern and Southern America for tischeriid species with striking
genitalia and feeding characters.
Three main lineages of generic rank can be recognized within Tischeriidae (fig.
24). The first (Tischeria) may be the best characterized by the development of a juxta
in the male genitalia, and an antrum in the female genitalia; the second (Astrotischeria)
by the development of a dorsal lobe to the valva, the uncus overlaid dorsally by lobes
of the pseuduncus, utilization of new host-plant families (particularly Asteraceae),
and the third (Coptotriche) by the development of transtilla, spines on the
diaphragma, a ‘tulip-shaped’ aedeagus and by the great enlargement of the
membranous half of the ductus spermathecae in the female genitalia.
The cladogram of the Tischeriidae (fig. 24) is based on 34 apomorphies listed
in the disertation or published version in Diðkus, Puplesis, 2003.
33

Fig. 24. A cladogram of the Tischeriidae (for a list of used apomorphies 1–34 see the
dissertation or published version in Diðkus, Puplesis, 2003.)
34
Taxonomic composition
We recognize three genera among the Tischeriidae: Tischeria Zeller (27
described species for the World fauna), Astrotischeria Puplesis & Diðkus (30 species)
and Coptotriche Walsingham (57 species). For detailed diagnostic descriptions of
the recognized genera – see the doctoral thesis or published version in Diðkus,
Puplesis, 2003).
MAIN CONCLUSIONS
Intensive field work in Lithuania confirmed the occurrence of 73
Nepticulidae species (12 new to the fauna, 5 excluded) and demonstrated
15 plant families and 32 plant genera as host-plants, where most of
Lithuanian Nepticulidae species are trophically associated with plants from
Rosaceae, Salicaceae, Fagaceae and Betulaceae.

In Central Asia, 90 nepticulid species were recognized; in total, the
endemism of the Central Asiatic Nepticulidae fauna is very high (71%)
and about half (53%) the Central Asiatic species with a known host-plant
are trophically associated with Rosaceae.
Study of material collected from Oman and the Indian subcontinent yielded
34 species of Nepticulidae (including twelve new to science).
In total, 105 species of Nepticulidae are reviewed for East Asia (63 of them
were recorded only from the eastern part of the Mandshurian region,
with one species as new to science and 15 species distributed from Europe
to the Mandshurian region.
Study of Neotropical material (74 earlier recorded species and 16 new)
has shown a high level of endemism of the Nepticulidae fauna in general,
and also demonstrated the phenomenon of Acalyptris predomination.
Ten species of Tischeriidae (including 9 new ones) are now known from
southern Africa, i. e., the same number as from Europe with a two hundred
year history of investigation; the southern African tischeriids are associated
with host-plants of at least of 4 families and 6 genera: Rhamnaceae (Scutia),
Tiliaceae (Grewia, Sparmannia, Triumfeta), Anacardiaceae (Rhus) and
Combretaceae (Terminalia).
New data and revision of all published records confirmed 1027 species of
Nepticuloidea & Tischerioidea currently recognized for the world fauna;
most of them, 780 species, belong to the Nepticulidae, 130 to the
Opostegidae 117 to the Tischeriidae. In the Neotropical region the
nepticulid genus Acalyptris predominates (accounting for up to 50% of the
nepticulid fauna); Stigmella is the dominant nepticulid genus in most
geographical regions and in non-tropical areas of the Neotropical region.
The world Nepticuloidea & Tischerioidea are trophically associated with
66 plant families: most species occur on Rosaceae (123 species) and
Fagaceae (93 species); to a lesser extent, but still widely utilised, are such
families as: Rhamnaceae (32 species), Asteraceae (31 species), Fabaceae
(31 species), Salicaceae (28 species) and Betulaceae (27 species).
In the light of host-plant data, the evolution of the Nepticuloidea or Tischerioidea
probably occurred by way of trophic adaptation to 7 plant subclasses from the 12
recognized; four of these host-plant subclasses are common to all three studied
moth families: Hamamelididae, Dilleniidae, Rosidae and Lamiidae.
Three main lineages of generic rank can be recognized within Tischeriidae.
The first (Tischeria) may be the best characterized by the development of a
juxta in the male genitalia, and an antrum in the female genitalia; the
second (Astrotischeria) by the development of a dorsal lobe to the valva, the
uncus overlaid dorsally by lobes of the pseuduncus, utilization of new
host-plant families (particularly Asteraceae), and the third (Coptotriche)
by the development of transtilla, spines on the diaphragma, a ‘tulip-shaped’
aedeagus and by the great enlargement of the membranous half of the
ductus spermathecae in the female genitalia.
35

SANTRAUKA
Problemos aktualumas. Dël aktyvaus natûraliø kraðtovaizdþiø naikinimo, daþnos
gyvûnø ar augalø gyvenamøjø vietø fragmentacijos bei destrukcijos sparèiai
ëmë nykti biologinë ávairovë. Vis didëjantis pasaulio mokslinës visuomenës susirûpinimas
biologinës ávairovës krize paskatino tyrinëtojus aktyviau inventorizuoti
pagrindinius Þemës biomus. Iðaiðkinti pasaulinës biologinës ávairovës apimtis –
didelis ir ilgalaikis sistematikø uþdavinys.
Mûsø tyrimø objektas – filogenetiðkai vieni primityviausiø (ir tuo, teoriniu
poþiûriu, vieni svarbiausiø bei ádomiausiø) Lepidoptera bûrio taksonai (Nepticuloidea
ir Tischerioidea), jungiantys tris giminiðkas ðeimas: Nepticulidae, Opostegidae
ir Tischeriidae. Ðios ðeimos plaèiai paplitusios visuose þemynuose (iðskyrus
Antarktá) ir jungia maþiausius pasaulyje mikrodrugius, kurie iðsiskiria ne tik archaiðka
sandara, bet ir labai didele specializacija.
Dël stenofagijos tendencijø, sëslaus gyvenimo bûdo ir endemizmo ðie smulkûs
vabzdþiai gausiai paplitæ visuose sausumos biomuose (pradedant dykumomis
ir baigiant atogràþø miðkais) ir yra vieni puikiausiø objektø, charakterizuojanèiø
tiriamo biomo biologinës ávairovës turtingumà, kilmæ ir gamtinius ryðius. Taèiau
ðiø, praktinæ ir teorinæ reikðmæ turinèiø, vabzdþiø tyrimø duomenø (tiek Lietuvos,
tiek pasaulio atogràþø regionø) nepakanka. Iki disertacijos autoriaus ir jo kolegø
atliktø tyrimø daugelio Þemës regionø fauna buvo visiðkai neþinoma ir neapraðyta,
o duomenys apie Lietuvos Nepticuloidea ir Tischerioidea rûðis buvo nepatikimi
ir pasenæ.
Taigi tyrimø duomenø apie Nepticuloidea ir Tischerioidea (ypaè Tischeriidae)
stoka, jø svarbi praktinë bei teorinë reikðmë, didëjantis mokslinës visuomenës
susirûpinimas biologinës ávairovës krize paskatino parengti ðià pasaulio Nepticuloidea
ir Tischerioidea revizijà; tokio pobûdþio darbø aktualumas buvo pabrëþtas
dar 1992 m. Rio de Þaneiro konvencijoje.
Tyrimø tikslas ir uþdaviniai. Disertacinio darbo tikslas – taksonominë primityviø
Microlepidoptera (Nepticuloidea ir Tischerioidea) pasaulio faunos apþvalga.
Ðiam tikslui pasiekti reikëjo ágyvendinti ðiuos uþdavinius:
atlikti Nepticulidae lauko tyrimus Lietuvoje ir iðaiðkinti gyvenanèias rûðis,
jø trofinius ryðius ir vikðrø minavimo sezoninius periodus;
atlikti ekspedicinius tyrimus Centrinëje Azijoje, iðaiðkinti ir apraðyti iki ðiol
neþinomas rûðis, sudaryti regiono Nepticulidae taksonominá sàraðà, nustatyti
svarbiausius rûðiø ávairovës centrus bei Centrinës Azijos Nepticulidae
trofinius ryðius;
iðtirti kolekcinæ medþiagà ið Omano, Indijos ir Nepalo ir apraðyti naujus
mokslui taksonus;
revizuoti duomenis apie Rytø Azijos Nepticulidae faunà, papildyti Mandþiûrijos
regiono Nepticulidae faunos sàraðà naujais tyrimø rezultatais;
revizuoti Centrinës ir Pietø Amerikos Nepticulidae, apraðyti naujas mokslui
rûðis, aptiktas Amazonës baseine ir Andø kalnuose, iðaiðkinti bendràsias
neotropinës faunos geografinio paplitimo ypatybes;
36

iðtirti Tischeriidae kolekcinæ medþiagà ið Pietø Afrikos ir apraðyti naujus
mokslui taksonus;
sudaryti pirmàjá Nepticuloidea ir Tischerioidea pasaulio faunos taksonominá
katalogà ir parengti originalià faunos apþvalgà, nustatant rûðiø kieká pasaulyje,
rûðiø apraðymo istorinæ raidà, bendruosius Nepticuloidea ir Tischerioidea
geografinio paplitimo bei trofiniø ryðiø ypatumus;
nustatyti autapomorfinius poþymius, árodanèius Tischeriidae monofiletinæ
kilmæ, sudaryti originalià ðeimos filogenetinæ kladogramà ir pateikti
naujà Tischeriidae klasifikacijà.
Mokslinis naujumas. Atrastos, apraðytos ar ávardytos ir parengtos apraðymui
(su bendraautoriais arba tik disertanto) 44 naujos mokslui Nepticulidae ir 36
naujos mokslui Tischeriidae rûðys (1, 2 pav.) bei viena nauja Tischeriidae gentis ið
iki ðiol maþai tirtø ar visai netyrinëtø kraðtø: Pirënø pusiasalio, Tuniso, Turkijos,
Turkmënistano, Tadþikistano, Rusijos Tolimøjø Rytø, Japonijos, Tailando, Nepalo,
Indijos, Omano, Pietø Afrikos Respublikos, Namibijos, Zimbabvës, Belizo,
Ekvadoro (Amazonës baseino ir Andø kalnø), Peru, Èilës ir Argentinos. Apraðytos
dar dvi naujos Bucculatricidae ðeimos rûðys (Bucculatrix multicornuta ir B.
macrognathos). Taip pat patikrinta visø 1027 nagrinëtø rûðiø taksonominë pozicija
ir padarytos 126 naujos taksonominës kombinacijos.
Pirmà kartà iðtyrinëta, preparuota ir apraðyta iki ðiol neidentifikuota Nepticulidae
ir Tischeriidae medþiaga, saugojama Britø muziejuje, Kopenhagos universitete,
Leideno gamtos muziejuje ir Smitsono centre (JAV).
Pirmà kartà iðtyrinëta tipinë 42 rûðiø medþiaga ir padaryti nauji apraðai, atitinkantys
keliamus reikalavimus.
Parengta originali Tischeriidae morfologinë ir taksonominë charakteristika,
remiantis naujaisiais duomenimis.
Kartu su bendraautoriais pirmà kartà taksonomiðkai revizuoti strateginiai
Þemës regionai (áskaitant Centrinæ ir Pietø Amerikà, Pietø Afrikà, Centrinæ Azijà
ir Lietuvà). Pirmà kartà pateikti duomenys apie ðiol neþinomà Nepticulidae faunà
Omane ir Nepale. Pateikta nauja Mandþiûrijos Nepticulidae apþvalga.
Pirmà kartà apibendrinti Nepticuloidea ir Tischerioidea pasaulio faunos rûðiø
apraðymo, trofiniø ryðiø ir geografinio paplitimo ypatumai.
Parengtas pirmas sistematinis Nepticuloidea ir Tischerioidea pasaulio faunos
katalogas, ávardijantis 1027 Nepticuloidea ir Tischerioidea rûðis, 224 mitybiniø
augalø gentis, priklausanèias 66 augalø ðeimoms. Atliekant ilgalaikius ekspedicinius
lauko darbus Lietuvoje, Kaukaze ir Centrinëje Azijoje, pirmà kartà iðaiðkinti
46 rûðiø mitybiniai augalai.
Rengiant disertacijà, buvo patobulintos smulkiø Microlepidoptera medþiagos
auginimo ið vikðrø ir genitaliniø mikropreparatø darymo metodikos.
Teorinë ir praktinë reikðmë. Viena svarbiausiø primityviø Microlepidoptera specializacijos
apraiðkø – jø entobiontinis gyvenimo bûdas þaliuosiuose augalo audiniuose.
Toks gyvenimo bûdas dar vadinamas minavimu, o augalø asimiliaciniuose
audiniuose padaryti paþeidimai vadinami minomis. Darbe nagrinëjami Nepticuloi-
37

dea ir Tischerioidea yra svarbûs kultûriniø ar miðko augalø kenkëjai (Kuznetzov,
Puplesis, 1994; Puplesis, Diðkus, 2003). Daugelis Nepticuloidea ir Tischerioidea
rûðiø – dël jø vikðrø entobiontinio gyvenimo bûdo asimiliaciniuose augalø lapø,
pumpurø, stiebø, jaunos þievës ar vaisiø audiniuose – yra svarbûs ûkiniu poþiûriu
(kaip kenkëjai ar potencialûs kenkëjai). Patikslinti duomenys apie mitybinius Nepticuloidea
ir Tischerioidea augalus ar iðaiðkinti iki ðiol neþinomi trofiniai ðiø vabzdþiø
ypatumai leidþia ne tik tiksliau apibrëþti atskirø kenkëjø minuotojø ekologines niðas,
bet ir parinkti efektyvesnius naikinimo ar gausumo reguliavimo kelius ir bûdus.
Suintensyvëjus Lietuvos prekybiniams mainams su egzotiðkaisiais kraðtais (ar
net ðiaip padidëjus keliautojø srautams), þymiai iðaugo atsitiktinës svetimø kraðtø
kenkëjø faunos introdukcijos pavojus. Jau pastebëta, kad rûðys oligofagës kartu
su mitybiniais augalais ar vien tik diapauzuojanèiais kokonais, patekusios á naujà
kraðtà, gali pradëti maitintis iki ðiol joms nebûdingais vietos kultûriniais augalais.
Ðiuo atveju gali bûti ypaè pavojingos tos kenkëjø rûðys, kurios trofiðkai nëra kraðtutinai
specializuotos ir kuriø mitybinë specializacija susijusi su Rosaceae ðeima,
vienijanèia obelis, kriauðes, slyvas, abrikosus, persikus, erðkëèius ar roþes bei kt.
Tokiø rûðiø, kurios trofiðkai susijusios su minëtais Rosaceae, daug yra pasaulio, o
ypaè Palearkties faunoje. Atsitiktinio kenkëjø áveþimo prevencijai bei karantino
priemoniø efektyviam taikymui yra bûtina þinoti ne tik ðiuo metu registruotø
kenkëjø rûðis ir jø biologijà, bet ir visø potencialiai pavojingø rûðiø geografiná
paplitimà bei trofinës specializacijos ypatybes. Disertacijos priede pateikiamas
pirmasis Nepticuloidea ir Tischerioidea pasaulio faunos katalogas, kuriame ne tik
aptariami revizuoti ir iðsamûs duomenys apie ðiø vabzdþiø mitybinius augalus, bet
ir padedama ávertinti potencialiø kenkëjø ratà, jø dabartinius arealus ir kilmës
centrus. Iðsamûs Nepticuloidea ir Tischerioidea rûðiø apraðai bus taip pat naudingi
augalø apsaugos, miðkø ûkio, parkø ar dekoratyviniø þeldiniø prieþiûros
specialistams, kad teisingai bûtø diagnozuojamos ðiø kenkëjø rûðys.
Kita vertus, primityvûs Microlepidoptera – neatskiriama kiekvienos sausumos
ekosistemos dalis. Minuojantys Lepidoptera yra labai „patogus“ objektas bendriems
biogeografiniams tyrimams. Jie paplitæ beveik visose svarbiausiose sausumos
ekosistemose, yra labai ávairûs, labai sëslûs ir palyginti lengvai pastebimi ir
tyrinëjami.
Darbo aprobacija. Svarbiausi rezultatai, sudarantys disertacijos pagrindà, buvo
pateikti svarstymui ir aprobuoti ðiose konferencijose ar moksliniø kolektyvø
posëdþiuose:
Tarptautiniame Rusijos – Suomijos biogeografiniame simpoziume (Sankt
Peterburgas, 1993) (2 praneðimai su bendraautoriais);
IX Europos entomologø kongrese (Lednice, Èekija, 1994);
Kopenhagos universiteto Zoologijos muziejaus Entomologijos laboratorijos
lepidopterologø seminare (Kopenhaga, 1996) (su bendraautoriumi);
Lietuvos entomologø draugijos konferencijoje (Vilnius, 1996);
Respublikinëje konferencijoje „Lietuvos bioávairovë (bûklë, struktûra, apsauga)“
(Vilnius, 1997);
38

VPU Zoologijos katedros posëdþiuose (Vilnius, 2001 ir 2003);
Britø muziejaus entomologijos laboratorijos Microlepidoptera sektoriuje
(Londonas, 2002);
Respublikinëje konferencijoje „Lietuvos biologinë ávairovë (bûklë, struktûra,
apsauga)“ (Vilnius, 2003) (2 praneðimai);
VU Zoologijos katedros posëdþiuose (Vilnius, 2004 ir 2005);
Tyrimø rezultatø publikavimas. Disertacijoje nagrinëjamø tyrimø rezultatai
paskelbti 21 darbe, kurie iðspausdinti moksliniuose þurnaluose, tæstiniuose ir vienkartiniuose
leidiniuose. Ið jø – 8 moksliniai straipsniai paskelbti Vakarø Europos
moksliniuose þurnaluose: Phegea (Belgija), Tijdschrift voor Entomologie (Olandija),
Bulletin of the Natural History Museum (Anglija) ir kt.
Svarbiausi tyrimø rezultatai apibendrinti kartu su bendraautoriumi mokslinëje
monografijoje (Puplesis R., Diðkus A. 2003. Nepticuloidea ir Tischerioidea
(Lepidoptera) pasaulio ir Lietuvos faunoje. The Nepticuloidea & Tischerioidea
(Lepidoptera) – a global review, with strategic regional revisions. 512 pp. „Lututë“,
Kaunas).
Autorius paskelbë: 9 darbus lietuviø kalba, 1 – rusø, 11 – anglø kalba.
Disertacijos sandara. Sudaryta ið ávado, tyrimø apþvalgos, metodø ir medþiagos,
rezultatø (2 skyriai ir 10 poskyriø), 29 iðvadø, literatûros sàraðo (482 ðaltiniai),
autoriaus disertacijos tema skelbtø darbø sàraðo (21 pozicija), angliðkos
santraukos ir 3 priedø. Ið viso – 388 puslapiai (207 – disertacijoje, 181 – prieduose),
382 paveikslø (109 – disertacijoje, 273 – prieduose), 16 lenteliø. Disertacija
paraðyta lietuviø k.
Padëkos. Nuoðirdi padëka skiriama ilgameèiams darbo vadovams (3, 4 pav.):
dr. Gaden S. Robinson (2000-2003, The Natural History Museum, London),
kurio talentas ir darbðtumas buvo ákvepiantis ir ypaè prof. habil. dr. Rimantui
Puplesiui (1990–2004, kuris ne tik iðmokë svarbiausiø tiriamojo darbo metodø ir
profesionaliai koordinavo disertacinio darbo procesà, bet ir skatino vykdomus
tyrimus, padëjo ásitraukti á tarptautinius mokslinius projektus, suteikë darbui didelio
patrauklumo ir naujumo bei padëjo pelnyti tarptautiná pripaþinimà. Autorius
dëkoja Vilniaus pedagoginio universiteto kolegoms bei vadovybei (ypaè GMF
Dekanui doc. dr. B. Ðalkui ir buv. Dekanui doc. dr. A. Vilkui, VPU vyr. buhalterei
Vidai Gulbinienei bei Mokslo ir Personalo skyriams) ir visiems Zoologijos katedros
darbuotojams, o taip pat katedros vedëjui doc. dr. Vytautui Semaðkai. Uþ
leidimà naudotis moksline medþiaga raðant disertacijà autorius dëkoja dr. Donald
R. Davis (USNM, Vaðingtonas), prof. dr. Niels P. Kristensen ir Ole Karsholt
(ZMUC, Kopenhaga), Kevin R. Tuck (BMNH, Londonas), dr. S. Yu. Sinev, dr.
S. Baryshnikova, dr. A. L. Lvovskyi (ZIN, Sankt Peterburgas), dr. Martin Kruger
(TMSA, Pretorija), dr. Yu. Budashkin (Karadagas, Krymas) dr. Owen Lewis
(Oksfordas, Anglija) ir Simon R. Hill (UW, Londonas). Autorius nuoðirdþiai
dëkoja dailininkei biologei Linai Jasiukonytei uþ Nepticuloidea ir Tischerioidea
morfologiniø struktûrø iliustracijas, o uþ lingvistiná darbà – Jolitai Þvironienei
(VPU).
39

TYRIMØ METODAI IR MEDÞIAGA
Minuojantiems vikðrams auginti buvo naudojamos chemiðkai ðvarios Petri
lëkðtelës su miðko paklotës imitacija. Monociklinës rûðys ar bicikliniø rûðiø rudeninë
medþiaga buvo laikoma ðaldytuve +10°C – +2°C temperatûroje iki sausio ar
geguþës mën., o tada reaktyvuojamos. Autoriaus darbe vidutinis medþiagos mirtingumas
buvo apie 30% (detalus apraðymas pateiktas disertacijoje).
Ðviesinëm gaudyklëms naudotos bedroselinës 125W ar droselinës 250W galingumo
DRL ir LRF lempos ir portatyvus elektros generatorius Honda EX-350.
Ekspedicijose po Centrinæ Azijà lauko darbai buvo atliekami naudojant dichlofosà,
kuris ne tik patikimai marina Nepticuloidea ir Tischerioidea suaugëlius, bet dël
ápakavimo yra patogus transportuoti karðto oro sàlygomis. Dichlofoso neigiama
átaka tolesniam medþiagos preparavimui nepastebëta. Kitos marinimo medþiagos
nepasiteisino. Surinkti kolekciniai egzemplioriai buvo smaigstomi minucijomis
(5–10 mm ilgio nerûdijanèio plieno miniatiûriniais smeigtukais).
Genitaliniams poþymiams tirti naudojome preparavimo metodikà, apraðytà
þemiau (5 pav.): 1. Po stereoskopiniu mikroskopu, virð balto kibaus plastiko padëklëlio,
su mikropreparavimo adatële atliekant ðvelnius judesius aukðtyn ir þemyn,
nulauþiamas iðdþiovinto drugio pilvelis. 2. Nulauþtas pilvelis preparavimo
adatële, pamirkyta glicerine, perkeliamas á mëgintuvëlá ir pipete álaðinama apie
1ml 10% KOH tirpalo. 3. Mëgintuvëlis kaitinamas ant atviros liepsnos (pvz., spiritinës
lemputës) arba verdanèiame vandenyje ir pilvelis virinamas tol, kol jis pasidaro
skaidrus; verdant pilvelá, mëgintuvëlá bûtina nuolat kratyti ir neleisti susidarantiems
oro purslams „iððauti“ kartu su ruoðiamu preparatu. 4. Mëgintuvëlio
turinys iðpilamas á ðvarià nedidelæ Petri lëkðtelæ ir preparavimo adatële perkeliamas
á kità lëkðtelæ su virintu arba distiliuotu vandeniu. 5. Atsargiai judinant adatëlæ,
preparatas nuplaunamas. 6. Ant labai kruopðèiai nuvalyto objektyvinio stiklelio
su duobute uþlaðinamas glicerino laðas ir uþdedamas dengiamasis stiklelis taip,
kad dalis glicerino laðo liktø neuþdengta. 7. Vandenyje nuplautas preparatas perkeliamas
á glicerinà ir atsargiai pakiðamas labai plona preparavimo adatële po
dengiamuoju stikleliu; preparatas turi bûti áspraustas tarp objektyvinio stiklelio ir
dengiamojo stiklelio ventraline puse á virðø; tam naudojamas stereoskopinis binokuliarinis
mikroskopas, didinantis 28–56 kartus. 8. Paruoðtas laikinasis mikropreparatas
stebimas galingesnio didinimo tiriamuoju mikroskopu. 9. Laikinasis mikropreparatas
saugojamas uþpylus cukraus persotintu tirpalu (t. y. cukraus kristale)
arba glicerino laðe (mini mëgintuvëlyje ar korekso juostos duobutëje, kuri ið
virðaus pridengiama tokios pat juostos plokðtele). 10. Ruoðiant pastovøjá mikropreparatà,
genitalinis aparatas dar kartà perplaunamas virintame arba distiliuotame
vandenyje, o po to perkeliamas ant ðvariai nuvalyto objektyvinio stiklelio su
duobute ir 30% etilo alkoholio tirpalu bei atskiriamas nuo pilvelio. 11. Toliau ið
preparato ir atskirto pilvelio paðalinamas vanduo uþpilant 70% etilo alkoholio ir
atsargiai paskalaujant ruoðiamà preparatà; prie pilvelio prikibæ þvyneliai nuvalomi
labai maþu plonu teptuku ar (ir) plona ir labai nusmailinta mikropreparavimo
adatële; tam naudojamas stereoskopinis binokuliarinis mikroskopas. 12. Prepa-
40

atas ir pilvelis, ið dalies paðalinus vandená, daþomas Chlorazol Black (retai –
merkurochromo) daþø spiritiniu tirpalu, uþlaðinant labai maþà ðiø daþø laðelá ant
ruoðiamo preparato. 13. Vandens paðalinimas ið genitalijø preparato ir pilvelio
uþbaigiamas uþpilant grynu etilo alkoholiu ir atsargiai paskalaujant preparatà
mikroadatële. 14. Ant kito, ðvariai nuvalyto objektyvinio stiklelio (autoriø darbe
valymui buvo panaudoti suspausto oro purðkikliai), uþlaðinamas nedidelis euparolio
laðelis (jeigu jis tirðtesnis nei ðvieþias medus, reikia atskiesti euparalio esencija).
15. Po stereoskopiniu mikroskopu ruoðiamos genitalinës struktûros bei
pilvelio iðnara perkeliami á euparalio laðà ir uþdengiami labai maþu dengiamuoju
stikleliu; genitalinis aparatas fiksuojamas ventraline puse á virðø, o atskiri skleritai
gali bûti praskleidþiami ar net iðmontuojami; kartais pilvelis ar atskiri genitalinio
aparato skleritai (pvz., ið kapsulës iðimtas aedeagus) fiksuojami po atskiru dengiamuoju
stikleliu, bet bûtinai ant to paties objektyvinio stiklelio. 16. Preparatas etiketuojamas
(ant objektyvinio stiklelio prilipinama popierinë etiketë), o paskui
dþiovinamas apie 2–3 mën. kambario temperatûroje arba apie 20 dienø kaitinimo
krosnelëje (+50–60°C temperatûroje) ant labai lygaus pagrindo.
Kad teisingai bûtø perteiktos preparatø proporcijos, genitalijø ir sparno gyslotumo
pieðiniai buvo daromi naudojant rusiðkà veidrodinæ pieðimo kamerà RA-
4. Suaugëliø iðorës morfologijai ar genitalinëms struktûroms tirti bei iliustruoti
buvo pasinaudota Kopenhagos universiteto skenuojanèiu mikroskopu „Jeol“
(JSM-840). Iðsamûs medþiagos preparavimo ir apraðymo metodai yra pateikti
disertacijoje ir autoriaus publikacijoje (Diðkus, Puplesis, 2003).
Dël Nepticuloidea ir Tischerioidea menko iðtirtumo pasaulio mokslo tyrimø
centruose yra labai nedideli ðiø primityviø Microlepidoptera rinkiniai. Todël viena
ið svarbesniø organizaciniø problemø buvo gauti bei sutelkti reikalingà medþiagà.
Ðiame darbe tyrinëta medþiaga tapo prieinama paskutiná deðimtmetá VPU Biosistematikos
laboratorijos inicijuotø ir iðvystytø moksliniø projektø, uþsienio fondø
paramos ir intensyvaus bendradarbiavimo su uþsienio institucijomis dëka (6 pav.).
Dalis medþiagos (rûðiø tipai, identifikuoti seniau apraðytø taksonø egzemplioriai
ar neidentifikuota medþiaga) buvo perduota ið Britø muziejaus (Londono Gamtos
muziejaus – BMNH, Anglija), Kopenhagos universiteto (ZMUC, Danija),
Kijevo universiteto bei Ukrainos MA Zoologijos instituto (Ukraina), Leideno
Gamtos muziejaus (NNM, Olandija), Rusijos Zoologijos instituto (ZIN, Sankt
Peterburgas), Pretorijos muziejaus (TMSA, Pietø Afrikos Respublika), Namibijos
nacionalinio muziejaus (NMN, Namibija) ir Smitsono centro (USNM, JAV).
Dalá labai svarbios medþiagos ið ekspediciniø rinkiniø, surinktø Pietø Amerikoje,
Nepale, Rytø Azijoje, Afrikoje, perdavë O. Karsholt, E. S. Nielsen, W. Mey ir kt.
kolegos. Dalá labai svarbios medþiagos, sukauptos ekspedicijose pateikë O. Karsholt
ir dr. E. Nielsen (rinkiniai ið Pietø Amerikos), prof. habil. dr. R. Puplesis ir
Simon Hill (rinkiniai ið Belizo, Centrinës Amerikos; Ekvadoro, Pietø Amerikos).
Pasinaudota prof. R. Puplesio rinkiniais ið Tolimøjø Rytø, Nepalo ir ávairiø Centrinës
Azijos regionø. Kità medþiagà autorius pats surinko ekspediciniø tyrimø
metu Lietuvoje, Kaukaze (Abchazijoje) ir Centrinëje Azijoje (Turmënistane: Ko-
41

petdage, Tedþene bei Tadþikistane: Varzobo kanjone ir Tavildaroje). Lietuvoje A.
Diðkus Nepticulidae ávairovæ daugiausia tyrë Klaipëdos, Ðilutës, Trakø, Kaiðiadoriø,
Alytaus ir Vilniaus rajonuose bei Kurðiø nerijoje, taikydamas suaugëliø auginimo
ið vikðrø metodà.
Ið viso darbe buvo iðtyrinëta daugiau nei 7900 Nepticuloidea ir Tischerioidea
egzemplioriø; ið jø – apie 1350 egzemplioriø buvo paruoðti laikinieji arba pastovieji
genitalijø mikropreparatai. Autoriaus iðtirta medþiaga atskleidë 80 naujø mokslui
rûðiø (1, 2 pav.). Atrastø ir apraðytø naujø rûðiø tipinë medþiaga yra saugojama
arba VPU, arba tose mokslo institucijose, ið kur medþiaga buvo gauta.
TYRIMØ REZULTATAI
STRATEGINIØ REGIONØ TAKSONOMINËS REVIZIJOS
Autoriui pavyko iðauginti 93% ðiuo metu iðaiðkintos Lietuvos faunos imagø ir
taip atskleisti daug naujø Nepticulidae biologijos ypatybiø bei 12 naujø Lietuvos
faunai rûðiø (1 lentelë). Tyrimais nustatyti Nepticulidae trofiniai ryðiai (7 pav.) ir
vikðrø minavimo pikai Lietuvoje (8 pav.).
Duomenys Centrinës Azijos apie rûðiø paplitimà ir trofinius ryðius buvo þymiai
praplësti (2 lentelë), kadangi pavyko pirmà kartà ið minuojanèiø vikðrø iðauginti
daug rûðiø suaugëliø. Atskleistas faunos endemizmas yra labai didelis (71%),
o tai tik ið dalies gali bûti siejama su nepakankamu Nepticulidae iðtyrimu kaimyniniuose
kraðtuose (9, 10 pav.)
Centrinëje Azijoje (daugiausia Tadþikistane) buvo registruota nemaþai S. betulicola
rûðiø grupës atstovø. Tai rodo didelæ ðios rûðiø grupës diferenciacijà Azijoje;
ir nors paprastai ðios grupës rûðys yra sunkiai diagnozuojamos, naujos rûðys
pasiþymi gana specifiniais morfologijos poþymiais, rodanèiais didelæ filogenetinæ
tarpusavio izoliacijà ir galbût ilgà ðiø rûðiø amþiø.
Paaiðkëjo, kad rytinë Centrinës Azijos dalis (Turkestano ar Tian Ðanio provincijos)
ir regiono vakaruose esantis Kopetdago kalnagûbris (Persijos provincija)
apgyvendintas gana skirtinga Nepticulidae fauna. Dël Irano – Turano aridinio
„koridoriaus“, skirianèio mezofilinës floros (o kartu ir ant jos gyvenanèios faunos)
arealus, tik keletas Nepticulidae rûðiø uþima arealus nuo vakarø Turkmënistano
iki Tadþikistano. Kai kurios rûðys (pvz., labai giminiðkos Ectoedemia petrosa
Puplesis ir E. albiformae Puplesis & Diðkus), matyt, gali bûti vertinamos kaip
vikarijuojanèios. Jos greièiausiai yra alopatrinës rûðys, kilusios ið vieno artimo
protëvio, bet dabar grieþtai pasiskirsèiusios regionais (viena aptinkama tik Turkestano,
kita – tik Persijos provincijoje).
Beveik pusë (53%) rûðiø, kuriø trofiniai ryðiai jau iðaiðkinti, yra susijusios su
Rosaceae ðeima (11 pav.). Tarp rûðiø, kuriø mitybiniai augalai dar nenustatyti,
vyrauja Acalyptris genties atstovai; ðioje gentyje kol kas në viena Centrinës Azijos
neptikulidø rûðis nëra trofiðkai iðtirta ir todël ateityje gauti duomenys gali labai
pakeisti sampratà apie trofiniø ryðiø ávairovæ Centrinëje Azijoje.
Iðtyrus Omane ir Indostane surinktà naujà medþiagà, atogràþø regione – nuo
Arabijos pusiasalio iki Himalajø atogràþiniø priekalniø – buvo nustatytos 32 Nep-
42

ticulidae rûðys; dar dvi neatogràþinës rûðys iðaiðkintos Himalajø aukðtikalnëse (3
lentelë).
Naujausi kolekciniai duomenys ið Mandþiûrijos leido revizuoti ir papildyti regiono
Nepticulidae sistematiná sàraðà (4 lentelë, 12 pav.). Registruota 15 rûðiø,
bûdingø ir Europos faunai. Tai leidþia patvirtinti rytø ir vakarø Palearkties istorinio
giminiðkumo koncepcijà.
Naujai apþvelgta Centrinës ir Pietø Amerikos Nepticulidae fauna (5 lentelë).
16 Nepticulidae rûðiø apraðytos kaip naujos mokslui. Nauji duomenys leido iðaiðkinti
kelis naujus mitybinius augalus. Nustatytas labai didelis Neotropinës Nepticulidae
faunos endemizmas. Visos ðiuo metu þinomos Centrinës ir Pietø Amerikos
neptikulidø rûðys nëra þinomos uþ Neotropinës srities ribø; kelios rûðiø grupës
ir viena gentis (Manoneura), matyt, taip pat yra Neotropinio regiono endeminiai
taksonai.
Galbût pats netikëèiausias atskleistas fenomenas – Acalyptris genties vyravimas
neotropinëje Nepticulidae faunoje. Ið visø Belize iðaiðkintø neptikulidø rûðiø
15 jø priklauso Acalyptris genèiai, o tai sudaro apie 48% ðio kraðto iðaiðkintos
faunos. Ádomu tai, kad beveik tiek pat procentø (apie 50%) Acalyptris genties
neptikulidø buvo pakartotinai nustatyta ir Amazonës baseino faunoje (Puplesis,
Diðkus, 2003). Dar gali bûti paminëta tai, kad Neotropinëje srityje atrastos Acalyptris
rûðys pasiþymi labai didele morfologiniø struktûrø ávairove.
Disertacinio darbo metu nustatyta 10 Tischeriidae rûðiø ið Pietø Afrikos (6
lentelë), todël galima spëti, kad tik pradëta apraðinëti Pietø Afrikos regiono fauna
yra labai ávairi. Didelæ regiono Tischeriidae biologijos ávairovæ parodë ir trofiniai
ryðiai.
PASAULIO NEPTICULOIDEA IR TISCHERIOIDEA TAKSONOMINË APÞVALGA
Apibendrinus literatûrinius duomenis, iðtyrus daugelio anksèiau apraðytø rûðiø
tipinæ medþiagà bei susumavus autoriaus taksonominiø tyrimø rezultatus,
nustatyta, kad ðiuo metu Þemëje þinomos 1027 Nepticuloidea ir Tischerioidea
rûðys (ið jø Nepticulidae – 780 rûðiø, Opostegidae – 130 rûðiø ir Tischeriidae – 117
rûðiø). Visos ðios rûðys átrauktos á pirmà kartà sudarytà Nepticuloidea ir Tischerioidea
pasaulio faunos katalogà, kuriame patikslinta 126 rûðiø taksonominë pozicija,
o taip pat nurodomos 224 mitybiniø augalø gentys.
Disertacijoje plaèiai tirtø ðeimø ir genèiø geografinio paplitimo ypatybës (13,
14 pav.), endemizmo fenomenas, o taip pat nacionalinës faunos.
Ðiuo metu ið 1027 dabar þinomø pasaulio Nepticuloidea ir Tischerioidea faunos
rûðiø yra nustatyti 580 rûðiø mitybiniai augalai. Sprendþiant pagal tas rûðis,
kuriø trofiniai ryðiai yra þinomi, pasaulio Nepticuloidea ir Tischerioidea minuoja
ant 66 ðeimø augalø: Nepticulidae nustatyta ant 62 ðeimø augalø, Opostegidae –
6, Tischeriidae – 15. Taèiau vienà augalø ðeimà minuojanèiø rûðiø skaièius yra labai
skirtingas (nuo 1 iki 123 rûðiø).
Trofiniø Nepticuloidea ir Tischerioidea ryðiø analizë pagal augalø aukðtesniuosius
taksonus (augalø eiliø ar poklasiø rango) teikia maþai informacijos apie
galëjusià vykti trofiniø ryðiø genezæ.
43

Iðskirti Tischeriidae morfologiniai poþymiai (15–23 pav.), parodantys taksonà, kaip
monofiletinës kilmës sistematiná vienetà. Atnaujintas Coptotriche genties taksonominis
statusas, o anksèiau apraðytos Emmetia Leraut (1993) genties pavadinimas nurodytas
kaip vëlesnysis Coptotriche genties sinonimas. Apraðyta nauja Astrotischeria gentis.
Remiantis 34 apomorfiniais poþymiais, iðvardytais disertacijoje, ir parengta
kladograma (24 pav.), atskleistos trys Tischeriidae ðeimõs evoliucinio vystymosi
kryptys arba filogenetinës ðakos.
GINAMOS IÐVADOS
Revizuota Lietuvos Nepticulidae fauna
1. Po autoriaus 1996–2003 m. atliktø lauko tyrimø ávairiuose Lietuvos rajonuose
nustatytos 73 Nepticulidae rûðys; 12 ið jø – naujos Lietuvos faunoje: Stigmella
nivenburgensis, S. crataegella, S. ulmivora, S. benanderella, S. continuella, S.
poterii, S. ulmariae, S. hemargyrella, Bohemannia pulverosella, Ectoedemia turbidella,
E. klimeschi, E. subbimaculella.
2. Keletas neptikulidø rûðiø, apie kuriø paplitimà Lietuvoje nurodë ankstesni
autoriai, iðbrauktos ið Lietuvos Nepticulidae sàraðo: dvi rûðys ankstesniø tyrëjø
buvo paskelbtos remiantis klaidingai apibûdintomis minomis (Stigmella mespilicola,
Ectoedemia spinosella), trys rûðys (Stigmella sakhalinella, S. sanguisorbae ir Ectoedemia
angulifasciella) anksèiau buvo minimos nesiremiant iðauginta medþiaga ar
minomis, surinktomis mûsø ðalyje.
3. Autentiðkais tyrimais nustatyta 15 Lietuvos Nepticulidae mitybiniø augalø
ðeimø ir 32 mitybiniø augalø gentys. Daugiausia Lietuvos neptikulidø rûðiø yra
trofiðkai susietos su Rosaceae, Salicaceae, Fagaceae ir Betulaceae augalais, taèiau
pirmoji mitybiniø augalø ðeima (Rosaceae) yra vyraujanti; ant ðios ðeimos augalø
autorius rado 24 neptikulidø rûðis.
4. Nustatyti du Nepticulidae vikðrø minavimo pikai Lietuvoje: pirmasis minavimo
aktyvumo laikotarpis – nuo birþelio vidurio iki liepos vidurio, kai vienu metu
minuoja 36 neptikulidø rûðys, antrasis – nuo rugsëjo vidurio iki spalio vidurio, kai
vienu metu minuoja 49 rûðys.
Revizuota Centrinës Azijos Nepticulidae fauna
5. Ið viso Centrinëje Azijoje iðaiðkinta 90 Nepticulidae rûðiø, ið kuriø dauguma
(54 rûðys) priklauso Stigmella genèiai, 16 – Acalyptris, 11 – Ectoedemia, 4 –
Fomoria, po dvi – Glaucolepis ir Etainia bei viena – Trifurcula genèiai.
6. Centrinëje Azijoje atrasta ir autoriaus apraðyta 15 naujø neptikulidø rûðiø
(Stigmella betulifoliae, S. pamirbetulae, S. excelsa, S. polymorpha, S. fasciola, S. johanssoni,
S. divina, S. cerasi, S. aflatuniae, Ectoedemia albiformae, Fomoria lacrimulae,
F. favimacula, Acalyptris argyraspis, Etainia leptognathos, E. obtusa).
7. Nustatyta, kad vakarø Kopetdagas ir pietiniai Gisaro kalnagûbrio ðlaitai
gali bûti laikomi paèiomis turtingiausiomis Nepticulidae faunos vietovëmis, taèiau
treèdaliui Kopetdago faunos atstovauja euraziniø arealø rûðys, o apie 93% iðaiðkintos
Gisaro kalnagûbrio faunos sudaro Centrinës Azijos endemikai. Nustatytas
bendras Centrinës Azijos Nepticulidae endemizmas siekia 71%.
44

8. Autoriaus tyrimai parodë, kad beveik pusë (53%) Centrinës Azijos neptikulidø
rûðiø, kuriø trofiniai ryðiai iðaiðkinti, yra susijusios su Rosaceae ðeima (minuoja
ant Aflatunia, Cerasus, Crataegus, Malus, Prunus, Pyrus, Rosa, Rubus, Spiraea
genèiø augalø). 11% trofiðkai iðtirtø rûðiø minuoja Salicaceae augalus (Populus,
Salix), po 8% minuoja Ulmaceae (Celtis, Ulmus), Aceraceae (Acer), Rhamnaceae
(Paliurus, Rhamnus, Ziziphus), 6% – Betulaceae (Betula). Po vienà rûðá nustatyta
ant Hypericaceae, Moraceae ir Fagaceae augalø.
Pirmieji duomenys apie Omano, Indijos ir Nepalo Nepticulidae
9. Iðtyrus Omane ir Indostane surinktà kolekcinæ medþiagà, ið viso atogràþø
regione – nuo Arabijos pusiasalio iki Himalajø priekalniø – nustatytos 32 Nepticulidae
rûðys. Dvi borealinio arealo rûðys iðaiðkintos Himalajø aukðtikalnëse. Ið viso
apþvelgtame regione nustatytos 34 rûðys, ið kuriø 12 yra naujos.
10. Stigmella hoplometalla ir Acalyptris melanospila radimas Himalajø atogràþiniuose
miðkuose þymiai prapleèia duomenis apie ðiø rûðiø, iki ðiol þinomø tik
Bombëjaus regione, paplitimà ir leidþia teigti apie platø rûðiø arealà Indostano
subkontinente.
11. Himalajø atogràþiniuose miðkuose aptiktos trys labai artimos, bet lengvai
diagnozuojamos Acalyptris genties rûðys (A. melanospila, A. auratilis ir A. nigripexus),
kuriø buveiniø ir skraidymo laiko sutapimas leidþia teigti apie simpatrinæ ðiø
rûðiø kilmæ. Tuo tarpu aukðtuosiuose Himalajuose gyvenanti Stigmella tenebrica
morfologiðkai þymiai artimesnë europinei S. sorbi negu tos paèios grupës rûðims,
gyvenanèioms geografiðkai netolimoje Centrinëje Azijoje.
Revizuota ir papildyta Mandþiûrijos Nepticulidae fauna
12. Ið viso Rytø Azijoje nustatytos 105 Nepticulidae rûðys, ið kuriø 63 rastos
rytinëje Mandþiûrijoje. Vienà naujà rûðá – Stigmella auricularia – autoriaus apraðë
kartu su bendraautoriais.
13. Registruota 15 rûðiø, bendrø Europos ir Mandþiûrijos bei kaimyniniø regionø
faunai: Stigmella betulicola, S. luteella, S. sakhalinella, S. aurora, S. anomalella, S.
assimilella, S. salicis, S. obliquella, S. continuella, S. lediella, Bohemannia quadrimaculella,
Ectoedemia intimella, E. preisseckeri, E. occultella ir Fomoria weaveri. Dauguma jø
trofiðkai susijusios su Salicaceae (reèiau Rosaceae ir Ericaceae) ðeimø augalais.
14. Anksèiau þinomos kaip lokaliai europinio paplitimo, su Ulmus genties augalais
susijusios Ectoedemia preisseckeri radimas Rytø Azijoje patvirtina koncepcijà apie
dviejø geografiðkai nutolusiø – Mandþiûrijos ir Europos – faunø giminiðkumà.
Centrinës ir Pietø Amerikos Nepticulidae fauna
15. Ið viso tirtame regione nustatytos 74 Nepticulidae rûðys, ið kuriø 16 naujø,
t. y. Enteucha acuta, E. guajavae, Stigmella austroamericana, S. montanotropica, S.
nubimontana, S. rubeta, Fomoria repanda, F. tabulosa, Acalyptris ecuadoriana, A.
onorei, A. basihastatus, A. pseudohastatus, A. articulosus, A. rotundus, A. amazonius,
A. insolentis, disertantas apraðë kartu su bendraautoriais.
16. Tyrimais nustatytas neotropinës faunos endemizmas ir atskleistas Acalyptris
genties vyravimo fenomenas: tarp Belize iðaiðkintø neptikulidø rûðiø Acalyptris
genèiai priklauso 48%, Amazonës baseino vakarinëje dalyje – apie 50% rûðiø.
45

Nauji duomenys apie Pietø Afrikos Tischeriidae
17. Ðiuo metu Pietø Afrikoje nustatytos 10 Tischeriidae rûðiø, tai yra lygiai
tiek, kiek Europoje; 9 naujas Afrikos tiðeridø rûðis (Tischeria sparmanniae, T.
martinkrugeri, T. antilope, Coptotriche zimbabwiensis, C. africana, C. basipectinella ir
kt.) autorius apraðë su R. Puplesiu.
18. Iðaiðkinti tirto regiono tiðeridø trofiniai ryðiai, kurie ðiuo metu susijæ su 4
ðeimø ir 6 genèiø augalais: Rhamnaceae (Scutia), Tiliaceae (Grewia, Sparmannia,
Triumfeta), Anacardiaceae (Rhus) ir Combretaceae (Terminalia).
Pasaulio Nepticuloidea ir Tischerioidea taksonominë apþvalga
19. Ðiuo metu Þemëje þinomos 1027 Nepticuloidea ir Tischerioidea rûðys (ið jø
Nepticulidae – 780 rûðiø, Opostegidae – 130 rûðiø ir Tischeriidae – 117 rûðiø).
20. Apraðant Nepticuloidea ir Tischerioidea rûðis, ið viso dalyvavo 123 autoriai
(be bendraautoriø arba su bendraautoriais, tai yra ávairiø autoriniø kombinacijø).
Apraðant rûðis, svarbiausi aktyvumo pakilimai buvo apie 1911–1915, 1976–1980 ir
ypaè 1984–1985 bei 2000 metus. Disertacijos autorius ið viso atrado ir apraðë
(vienas arba su bendraautoriais) 44 naujas Nepticulidae ir 36 Tischeriidae rûðis.
21. Daugiausia Nepticuloidea ir Tischerioidea rûðiø (445 rûðys arba apie 44%)
registruota Palearkties regione. Ið Nepticulidae ðeimos tik Stigmella ir Fomoria yra
kosmopolitinio paplitimo, o Acalyptris, skiringai negu buvo galvojama anksèiau,
taip pat aptinkamas beveik visuose biogeografiniuose regionuose. Pabrëþtinas
Stigmella dominavimas visø Þemës regionø faunose, o Acalyptris – Neotropiniame
regione (30–35% iðaiðkintø regiono rûðiø, nors kai kuriose vietovëse Acalyptris
rûðys sudaro 50% neptikulidø faunos).
22. Remiantis Nepticulidae genèiø paplitimu, Palearkties ir Afrotropinio regiono
endemizmas siekia apie 20%, Neotropionio regiono – apie 17%, o Australinio
apie 50%.
23. Amerikos þemyno – didþiausio Tischeriidae ávairovës ir galbût kilmës centro
– iðskirtinumà pabrëþia taksonominë èia aptinkamos faunos struktûra: be
Coptotriche ir Tischeria genèiø, kurios þinomos kiekviename regione, iðskyrus Australiná,
Neotropiniame regione vyrauja Astrotischeria rûðys (74% tiðeridø faunos).
24. Sprendþiant pagal tas rûðis, kuriø trofiniai ryðiai iðaiðkinti, pasaulio Nepticuloidea
ir Tischerioidea minuoja ant 66 ðeimø augalø: daugiausiai ant Rosaceae
(123 rûðys) ir Fagaceae (93 rûðys), kiek maþiau ant Rhamnaceae (32) ir Asteraceae
(31), Fabaceae (31), Salicaceae (28) ir Betulaceae (27).
25. Labiausiai apibendrintame poklasiø lygyje iðaiðkëja, kad Nepticuloidea ir
Tischerioidea evoliucija vyko adaptuojantis minavimui ant 7-iø ið 12-os augalø
poklasiø; keturi ið ðiø poklasiø (Hamamelididae, Dilleniidae, Rosidae ir Lamiidae)
yra bendri ir Nepticulidae bei Opostegidae, ir Tischeriidae ðeimoms.
Nauja Tischeriidae sistema
26. Remiantis parengta nauja ðeimos charakteristika, iðskirti 6 morfologiniai
poþymiai, padedantys identifikuoti ðeimà ir parodantys taksonà, kaip monofiletinës
kilmës sistematiná vienetà: priekinis galvos kuokðtas, ilgos ir siûliðkos antenø
sensilos (sensilla chaetica), ðukiðki priedai virð akies (pecten), labai susiaurëjæs aede-
46

agus, sklerotizuotos pateliø kiauðdëèiø iðaugos, 4–5 apofiziø poros patelës genitalijose.
27. Atnaujintas Coptotriche genties taksonominis statusas, o anksèiau apraðytos
Emmetia Leraut (1993) genties pavadinimas nurodytas kaip jaunesnysis Coptotriche
genties sinonimas.
28. Remiantis 34 apomorfiniais poþymiais, iðvardytais disertacijoje, ir originalia
kladograma, atskleistos trys Tischeriidae ðeimõs evoliucinio vystymosi kryptys
arba filogenetinës ðakos: 1) rûðys, kuriø patino genitalijose iðsivysto juxta, o patelës
– antrum; 2) rûðys, kurioms bûdinga iðsivysèiusi valva nugarinë (dorsalinë)
skiautë ir kurios perëjo minuoti iðimtinai ant Malvaceae ir ypaè Asteraceae augalø;
3) rûðys, kuriø patino genitalijose iðsivystë transtilla, diafragma pasidengë spygliukais,
aedeagus ágavo „tulpës“ formà, o membraniðkoji patelës genitalijø ductus
spermathecae dalis labai pailgëjo.
29. Remiantis kladistinës analizës duomenimis, apraðyta nauja Astrotischeria
gentis, ðiuo metu vienijanti 30 amerikinio paplitimo rûðiø ir identifikuojama pagal
valva nugarinæ skiautæ, didelá vinculum, trumpà uncus ir penkias apophyses poras
patelës genitalijose bei mitybinius Malvaceae ir Asteraceae ðeimø augalus.
47

LIST OF PUBLICATIONS CONTAINING MATERIALS OF THE THESIS
Papers in scientific journals
1. Puplesis, R., Diðkus, A. 1995. Acalyptris argyraspis sp. n., a remarkable species
from Tadzhikistan (Lepidoptera: Nepticulidae). Phegea, 23 (1): 51–54.
2. Puplesis, R., Diðkus, A. 1996. First record of the genus Etainia Beirne from
Central Asia with descriptions of two new species and some provisional notes on
the world fauna (Lepidoptera: Nepticulidae). Phegea, 24 (1): 41–48.
3. Puplesis, R., Diðkus, A. 1996. A review of the Stigmella sorbi species-group
with descriptions of two new species from Turkmenistan and Tadzhikistan (Lepidoptera:
Nepticulidae). Phegea, 24 (4): 171–182.
4. Puplesis, R., Diðkus, A. 1996. Five new mining Lepidoptera (Nepticulidae,
Bucculatricidae) from Central Asia. Tijdschrift voor Entomologie, 139 (2): 181–190.
5. Puplesis, R., Sruoga, V., Diðkus, A., Puplesienë, J. 1996. Minuojanèiø drugiø
(Lepidoptera) autochtoninës evoliucijos centrai Vakarø Azijoje. In: Jonaitis, V.
(ed.). Lietuvos entomologø darbai (Lietuvos entomologø draugijos 30-meèiui). 48–51.
Vilnius.
6. Diðkus, A., Puplesis, R. 1996. Palearktikos Borealinio regiono Nepticulidae
(Lepidoptera) fauna: taksonominë sudëtis ir zoogeografiniai ryðiai. In: Jonaitis,
V. (ed.). Lietuvos entomologø darbai (Lietuvos entomologø draugijos 30-meèiui). 52–
55. Vilnius.
7. Diðkus, A. 1996. “Europinës” drugiø minuotojø (Lepidoptera: Nepticulidae,
Opostegidae, Tischeriidae, Bucculatricidae) rûðys Centrinës Azijos faunoje.
In: Jonaitis, V. (ed.). Lietuvos entomologø darbai (Lietuvos entomologø draugijos 30meèiui).
56–59. Vilnius.
8. Puplesis, R., Diðkus, A., Nieukerken, E. J. van. 1997. Stigmella divina sp. n.,
a remarkable species from Turkmenistan and Turkey (Lepidoptera, Nepticulidae).
Tijdschrift voor Entomologie, 140 (1): 55–58.
9. Diðkus, A. 1998. Review of the Tischeriidae (Lepidoptera) of Central Asia.
Acta Zoologica Lituanica, 8 (3): 23–33.
10. Sruoga, V., Diðkus, A. 2001. Stephensia brunnichella (Lepidoptera: Elachistidae)
new species for Lithuania. Acta Zoologica Lituanica, 11 (1): 73–77.
11. Puplesis, R., Diðkus, A., Robinson, G. S. 2002. New Neotropical Nepticulidae
(Lepidoptera) from the western Amazonian rainforest and the Andes of
Ecuador. Bulletin of the Natural History Museum, London (Entomology), 71 (1):
19–58.
12. Puplesis, R., Diðkus, A., Robinson, G. S., Onore, G. 2002. A review and
checklist of the Neotropical Nepticulidae (Lepidoptera). Bulletin of the Natural
History Museum, London (Entomology), 71 (1): 59–76.
Monographs or chapters in monographs
13. Puplesis, R., Diðkus, A. 1997. Sem. Nepticulidae – neptikulidy (in Russian).
In: Ler, P. A. (ed.). Key to the insects of Russian Far East. Trichoptera and
Lepidoptera, 5 (1): 302–319. Dal’nauka Publishers, Vladivostok.
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14. Puplesis, R., Diðkus, A. 2003. Nepticuloidea ir Tischerioidea (Lepidoptera)
pasaulio ir Lietuvos faunoje. The Nepticuloidea & Tischerioidea (Lepidoptera) – a
global review, with strategic regional revisions. 512 pp. „Lututë“, Kaunas.
Diðkus, A., Puplesis, R. 2003. Tyrimø metodai ir medþiaga. In: Puplesis, R.,
Diðkus, A. Nepticuloidea ir Tischerioidea (Lepidoptera) pasaulio ir Lietuvos
faunoje. 22–31. „Lututë“, Kaunas.
Diðkus, A., Puplesis, R. 2003. Nepticuloidea ir Tischerioidea pasaulio faunoje.
In: Puplesis, R., Diðkus, A. Nepticuloidea ir Tischerioidea (Lepidoptera)
pasaulio ir Lietuvos faunoje. 38–175. „Lututë“, Kaunas.
Puplesis, R., Diðkus, A. 2003. Strateginiø regionø taksonominës revizijos
ir naujø rûðiø apraðai. In: Puplesis, R., Diðkus, A. Nepticuloidea ir Tischerioidea
(Lepidoptera) pasaulio ir Lietuvos faunoje. 176–289. „Lututë“, Kaunas.
Diðkus, A. 2003. Revizuota Lietuvos Nepticulidae fauna. In: Puplesis, R.,
Diðkus, A. Nepticuloidea ir Tischerioidea (Lepidoptera) pasaulio ir Lietuvos
faunoje. 290–317. „Lututë“, Kaunas.
Diðkus, A., Puplesis, R. 2003. Catalogue of the world Nepticuloidea &
Tischerioidea. In: Puplesis, R., Diðkus, A. Nepticuloidea ir Tischerioidea (Lepidoptera)
pasaulio ir Lietuvos faunoje. 318–436. „Lututë“, Kaunas.
15. Puplesis, R., Diðkus, A., Mey, W. 2004. Tischeriidae. In: Mey, W. (ed.). The
Lepidoptera of the Brandberg Massif in Namibia. Esperiana Memoir, 1: 39–51. Berlin.
Other publications
16. Diðkus, A., Puplesis, R., Sruoga, V., Puplesienë, J. 1994. The results of leafmining
Lepidoptera collections in the Persian province (Western Kopet Dag, 1993).
IX European Congress of Lepidopterology. 35–37. Konvoj Publishers, Lednice.
17. Diðkus, A. 1997. Taksonominiai Nepticulidae (Lepidoptera) ávairovës aspektai:
artimø rûðiø grupiø spektras Stigmella gentyje. Lietuvos bioávairovë (bûklë,
struktûra, apsauga). 58–59. Vilnius.
18. Diðkus, A., Juchneviè, V. 2001. Nepticulidae (Lepidoptera) minavimo laikas
Lietuvoje. VPU Gamtos mokslø fakulteto bakalaurø ir magistrantø mokslinës
konferencijos praneðimø medþiaga. 125–128. Vilnius.
19. Diðkus, A. 2003. Revizuota Lietuvos Nepticulidae (Lepidoptera) fauna.
Lietuvos biologinë ávairovë (bûklë, struktûra, apsauga). 23–24. Vilnius.
20. Diðkus, A., Puplesis, R. 2003. Nepticuloidea ir Tischerioidea rûðiø apraðymas,
trofiniø ryðiø bei geografinio paplitimo ypatumai. Lietuvos biologinë ávairovë
(bûklë, struktûra, apsauga). 24–25. Vilnius.
21. Puplesis, R., Diðkus, A. 2004. Ar Lietuvoje gyvena Maþieji gaubtagalviai?
Þurnalas apie gamtà, 6: 26–29.
49

50
DUOMENYS APIE AUTORIØ
Vardas, pavardë Arûnas Diðkus
Gimimo data ir vieta 1971 04 26, Marijampolë
El. paðtas a.diskus@vpu.lt
Iðsilavinimas ir profesija
1989–1994 m. Aukðtasis iðsilavinimas, Vilniaus pedagoginis universitetas, toliau
vadinamas VPU, Gamtos ir geografijos fakultetas; ágyta biologijos mokytojo
specialybë.
Darbo patirtis:
1990 m. – 1996 m. VPU, Gamtos mokslø fakultetas, toliau vadinamas GMF,
Biosistematikos laboratorijos laborantas.
1996 m. – iki ðiol VPU, GMF, Biologinës ávairovës ir technologijø laboratorijos
jaunesnysis mokslo darbuotojas.
2001 m. – iki ðiol VPU Zoologijos katedros asistentas. Vadovauja biologijos
specialybës studentø pratyboms, lauko praktikoms ir studentø kursiniams darbams.
Mokslinës staþuotës:
1994 m. rugsëjis Vienos Gamtos muziejus (Austrija).
1995 m. lapkritis–1996 m. vasarisKopenhagos universitetas, Entomologijos
laboratorija (Danija).
1997 m. gruodis–1998 m. vasaris Britø Gamtos muziejaus Entomologijos
laboratorija, Londonas (Didþioji Britanija).
2001 m. gruodis–2002 m. sausis Britø Gamtos muziejaus Entomologijos
laboratorija, Londonas (Didþioji Britanija).
Mokslinës ekspedicijos:
1990 m. spalis–lapkritis Ekologiniai tyrimai Picundos-Miusero rezervate
(Abchazija).
1991 m. birþelis–liepa Entomologinë ekspedicija Varzobe, Tadþikistano MA
botaninës stoties stacionare (Tadþikistanas).
1991 m. lapkritis–gruodisEntomologinës medþiagos rinkimas ir tyrimai
Aðchabado ir Duðanbës apylinkëse (Turkmënistanas ir Tadþikistanas).
1993 m. vasaris–kovas Entomologinës medþiagos rinkimas ir tyrimai Aðchabado
apylinkëse (Turkmënistanas).
1993 m. balandis–rugpjûtis (5 mën.) Entomologinë ekspedicija vakarø
Kopetdago kalnagûbryje (Choðdemiras ir Siunt Chasardago rezervatas),
bendradarbiaujant su Turkmënistano MA kolegomis (Turkmënistanas).
2005 m. sausis–vasaris Entomologinë ekspedicija Andø kalnuose (Pietø
Amerika, Ekvadoras).

52
Arûnas Diðkus
NEPTICULOIDEA IR TISCHERIOIDEA:
TAKSONOMINËS STRATEGINIØ REGIONØ REVIZIJOS IR PASAULIO
FAUNOS APÞVALGA
(INSECTA: LEPIDOPTERA)
Daktaro disertacijos santrauka
Biomedicinos mokslai, zoologija (05B)
Redagavo autorius
Maketavo Laura Barisienë
SL 605. 3,25 Sp. l. Tir. 100 egz. Uþsak. Nr. 05-025
Iðleido Vilniaus pedagoginis universitetas, Studentø g. 39, LT–08106 Vilnius
Spausdino VPU leidykla, T. Ðevèenkos g. 31, LT–03111 Vilnius
Kaina sutartinë