Pygmy moths (Nepticulidae, Lepidoptera) from Eocene Baltic amber are described with five new specimens, representing the only specimens beyond one described before. Some of the findings differ from Stigmellites baltica Kozlov in species and sex. They are formally described as new species Bohemannia butzmanni and B. aschaueri. Phylogenetic and paleoecological aspects of Nepticulidae from Baltic amber are discussed.Fünf neue Arten von Zwergminiermotten (Nepticulidae, Lepidoptera) werden aus Baltischem Bernstein (Eozän) beschrieben; zusammen mit einer bereits früher beschriebenen Form (Stigmellites baltica Kozlov) stellen sie die einzig bekannten Fossilien dieser Tiergruppe dar. Da einige der hier beschriebenen Fossilien sich von S. baltica deutlich unterscheiden, werden sie als neue Arten, Bohemannia butzmanni und B. aschaueri, beschrieben. Phylogenetische und paläoökologische Aspekte der Nepticulidae im Baltischen Bernstein werden diskutiert

Fischer, Thilo C.

Universität München: Elektronischen Publikationen

An International Journal  
of Palaeontology and Geobiology
Series A /Reihe A
Mitteilungen der Bayerischen Staatssammlung 
für Paläontologie und Geologie
München 2013
Zitteliana
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Zitteliana A 53 (2013) 85
Pygmy moths (Lepidoptera, Nepticulidae)  
from Baltic Amber (Eocene)
Thilo C. Fischer
Ludwig-Maximilians-University Munich, Department I, Botany, Großhadernerstr. 2–4, 
D-82150 Planegg-Martinsried,Germany
E-mail: thilo.fischer@biologie.uni-muenchen.de
 Zitteliana A 53, 85 – 92
 München, 31.12.2013
 Manuscript received 
30.09.2013; revision 
accepted 27.11.2013
 ISSN 1612 - 412X
Abstract
Pygmy moths (Nepticulidae, Lepidoptera) from Eocene Baltic amber are described with five new specimens, representing the only 
specimens beyond one described before. Some of the findings differ from Stigmellites baltica Kozlov in species and sex. They are formal-
ly described as new species Bohemannia butzmanni and B. aschaueri. Phylogenetic and paleoecological aspects of Nepticulidae from 
Baltic amber are discussed.
Key words: Baltic amber, Bohemannia, Eocene, leaf mining, Nepticulidae, pygmy moths
Zusammenfassung
Fünf neue Arten von Zwergminiermotten (Nepticulidae, Lepidoptera) werden aus Baltischem Bernstein (Eozän) beschrieben; zusam-
men mit einer bereits früher beschriebenen Form (Stigmellites baltica Kozlov) stellen sie die einzig bekannten Fossilien dieser Tiergruppe 
dar. Da einige der hier beschriebenen Fossilien sich von S. baltica deutlich unterscheiden, werden sie als neue Arten,  Bohemannia butz-
manni und B. aschaueri, beschrieben. Phylogenetische und paläoökologische Aspekte der Nepticulidae im Baltischen Bernstein werden 
diskutiert.  
Schlüsselwörter: Baltischer Bernstein, Bohemannia, Eozän, Blatt-minierer, Nepticulidae, Zwergminiermotten
1. Introduction
Nepticulidae are a monophyletic (natural) family 
of glossatan, monotrysian and heteroneurous Le-
pidoptera (Scoble 2002). Together with the close-
ly related extant family Opostegidae they form the 
superfamily Nepticuloidea. The family Nepticulidae 
comprises the smallest imagos of all Lepidoptera; 
the larvae mine leaves or other living plant surface 
tissues (petioles, bark, samaras) (Scoble 2002; Gri-
maldi and Engel 2005). With its leaf mining habit, the 
family is of great ecological importance and also has 
economic impact on a number of crop plants.
As a basal group of Lepidoptera phylogeny of 
Nepticulidae is of concern, but members of the fa-
mily are extremely rare as fossils (Sohn et al. 2012). 
Furthermore, most reports refer only to the typical 
traces of mining activity found in fossil leaves (most-
ly described as Stigmellites; see Sohn et al. 2012), 
the oldest of which come from the Mid-Cretaceous 
Dakota formation and are suggested to have been 
produced by Stigmella and Ectoedemia (Nepticuli-
dae) (Labandeira et al. 1994). Lepidopterous mines 
in leaves of the seed fern Pachypteris from the Aus-
tralian Upper Jurassic/Lower Cretaceous might be 
an even older evidence for Nepticulidae (Rozefelds 
1988). Undescribed putative nepticulid imagos are 
known from Cretaceous Siberian amber (mentioned 
in Grimaldi and Engel 2005) and from Canadian Late 
Cretaceous amber (mentioned in Kristensen and 
Skalski 1999). The evolution of Nepticulidae and 
other lepidopterous leaf miners is supposed to be 
closely linked to angiosperm radiation and hence 
host plant evolution (Labandeira et al. 1994; Grimaldi 
and Engel 2005). A first fossil nepticulid imago was 
mentioned by Skalski (1976, 1990) from Eocene Bal-
tic amber and affiliated to Ectoedemia sp. without 
description or even information on its deposition. 
Kozlov (1988) described a female imago as Stigmel-
lites baltica. Some of the nepticulid imagos studied 
Zitteliana A 53 (2013) 86
tip, three-segmented downwards-pointing maxilla-
ry palps, very short and blunt proboscis, abdomen 
slightly flat with tips of valves visible, no epiphysis, 
coxa and femur with transversal rows of scales, spur 
formula 0-2-4, middle hind spurs at half the length of 
hind tibia, five tarsomers, thorax with triangular tegu-
las, cilia at forewings and hindwings, twice as long at 
the hindwings, wing tips acute, venation consisting 
of sc, r1, r2-3, r4, r5, m1, m2, cu, and a; m1, m2 and 
cu distally diverging from r5.
Differential diagnosis: In comparison to Stigmel-
lites baltica Kozlov 1988 this specimen differs in the 
following characters: wing venation, (sex,) length of 
antennas, tibial spurs, hindwings broader, coxa and 
femur with transversal rows of scales. Affiliation to 
the extant genus Bohemannia is mainly supported 
by the structure of the venation and the presence of 
a collar of scales.
Description: Total length 2,2 mm, head 400 μm 
broad, vertex smooth, frontal tuft, epicranial notch 
visible, collar with piliform scales, eyes 130 μm in 
diameter, bases of antennas forming eye caps, fili-
form antennas with a length of about ¾ of forewing 
length with 30–35 segments (partly invisible), labial 
palps 150 μm with two segments, last segment with 
a thick middle part and a fine tip, pointed upwards, 
maxillary palps 250 μm with three visible segments, 
folded downwards, proboscis very short (100 μm) 
and blunt, mesonotum 400 μm broad, abdomen 870 
μm long, slightly flat and with irregularly preserved 
scales, male genitalia (tips of valves) visible, no epi-
physis at front legs, coxae and femurs with transver-
sal rows of scales, spur formula 0-2-4, middle hind 
spurs positioned at half the length of the hind tibia, 
five tarsomers, thorax with triangular tegulas, forew-
ings 1,8 mm long, cilia at forewings and hindwings, 
twice as long at the hindwings (up to 500 μm), scale 
covering only preserved at the proximal parts of the 
wings, wing tips acute, venation consisting of sc, r1, 
r2+3, r4+5, m1, m2, cu, and a; m1, m2 and cu di-
verging from r4+5 in the very distal part of the latter. 
There are no syninclusions preserved in this piece 
of amber.
Specimen 5217
Imago of pygmy moth (Nepticulidae) 
Fig. 2
Description: Total length 2,2 mm, head 400 μm 
broad, frons and vertex smooth, collar without sca-
les, diameter of eyes ca. 150 μm, eye caps at bases 
of antennas, filiform antennas with ca. 20 segments 
and a length half of that of forewings, maxillary (or 
labial?) palps bent and with 2 visible segments, other 
palps not visible, proboscis very small, abdomen 
length ca. 800 μm, mesonotum ca. 450 μm broad, 
spur formular 0-2-4, forewing length 1,9 mm, cilia at 
wings.
here clearly differ from Stigmellites baltica, and also 
represent the first male specimens discovered from 
Baltic amber.
2. Material and Methods
The amber pieces were purchased from trusted 
amber dealers. The inclusions were photographed 
with a binocular microscope (Zeiss Stemi 2000-C) 
to which is attached a digital camera system (Lei-
ca DFC 320, software LAS 4.1) (specimens 5058, 
5166); other images were obtained with the inclusi-
ons (specimens 5198, 5199, 5217). The specimens 
were preserved by treatment with an acryl varnish 
and are kept in the author’s collection at constant 
temperature in plastic clip bags within metal boxes 
excluding oxygen and light. However, transfer to a 
public collection (Bayerische Staatsammlung für Pa-
läontologie und Geologie, München, Deutschland) is 
envisaged.
3. Systematic paleontology
A total of five new nepticulid imagos have been 
discovered in the Eocene Baltic amber pieces.
Lepidoptera Linnaeus, 1758
Superfamily Nepticulidea Stainton, 1854
Family Nepticulidae Stainton, 1854
Genus Bohemannia Stainton, 1859
Bohemannia butzmanni nov. spec.
Fig. 1
Holotypus: Specimen ex coll. Fischer no. 5058, 
Fig. 1
Locus typicus: Amber mine of Yantarni, GUS
Stratum typicum: “Blaue Erde” (Upper Eocene – 
Lower Oligocene)
Etymology: The name is given to acknowledge 
the numerous paleontological activities of Mr. Rainer 
Butzmann (Munich, Germany)
Repository: Bayerische Staatssammlung für Palä-
ontologie und Geologie, Munich, Germany; accessi-
on number SNSB-BSPG 2013 I 93
Diagnosis: Very small moth with broad head, ver-
tex smooth but with frontal tuft, epicranial notch 
visible, piliform scales at collar, bases of antennas 
form eye caps, filiform antenna, length about ¾ of 
forewing length with 30–35 segments (partly invisi-
ble), two-segmented upwards-pointed labial palps, 
last segment with a thick middle part and a fine 
Zitteliana A 53 (2013) 87
Figure 1: Male pygmy moth (Nepticulidae), Bohemannia butzmanni nov. spec., from Eocene Baltic amber (SNSB-BSPG 2013 I 93); 
(a) Dorsal view; (b) Ventral view; total length of imago: 2,2 mm; (c) Corresponding line drawings with organ identification. Scale bars = 1 
mm.
Zitteliana A 53 (2013) 88
rewing length 1,95 mm, heteroneurous, wing tips 
pointed, hindwings not broad, cilia at front and hind 
wings, antennas with at least 26 segments (but an-
tennas broken), labial palps three-segmented, ma-
xillary palps with two- (or three-) visible segments, 
valves.
Differential diagnosis: Venation same as 5058 
(hence different to Stigmellites baltica), but frons with 
erect scales and galeae of proboscis slightly separa-
ted.  Preparation of genitals as performed in studies 
with extant Nepticulidae is not possible. Affiliation to 
the extant genus Bohemannia is due to the similarity 
of the venation.
Specimen 5198
Imago of pygmy moth (Nepticulidae) 
Fig. 4
Total length with legs 2,8 mm, head 400 μm broad, 
vertex with erect scales, collar with bundles of pili-
form scales behind antennas, eyes 180 μm in dia-
meter, eye caps, filiform antennas short (2/5 of front 
wing length) with 19 segments and pointed at the tip, 
labial palps two-segmented and downwards-poin-
ting, maxillary palps not visible, proboscis very short 
and with slightly separated galeae, mesonotum 400 
μm broad, abdomen 1100 μm long, no epiphysis, 
spur formula 0-2-4, tibias with scales, middle hind 
spurs positioned at 2/5 of length from basis of femur, 
five tarsomers, forewing length 1,8 mm, wings with 
scales preserved, cilia and longitudinal scales up to 
200 μm long at wing edges, wing tips acute.
This specimen may be conspecific with specimen 
5058, showing the same fore wing venation and also 
a smooth vertex.
Syninclusions: Cecidomyiidae (no. 5218), Germa-
raphis sp. (no. 5219).
Genus Bohemannia Stainton, 1859
Bohemannia aschaueri nov. spec.
Holotypus: Specimen ex coll. Fischer no. 5199, 
Fig. 3
Locus typicus: Amber mine of Yantarni, GUS
Stratum typicum “Blaue Erde” (Upper Eocene – 
Lower Oligocene)
Etymology: The name is given to acknowledge 
the numerous paleontological activities of Birgitt and 
Karl Aschauer (Waidhofen an der Ybbs, Austria).
Repository: Bayerische Staatssammlung für Palä-
ontologie und Geologie, Munich, Germany; accessi-
on number SNSB-BSPG 2013 I 94
Syninclusions: Trichomes, Arthropleones.
Diagnosis: Eye caps with scales, trochanter with 
rows of scales, proboscis very small, galeae sepa-
rated, spur formula 0-2-4, frons with erect scales, 
vertex smooth (with scales?), tegulas triangular, fo-
Figure 2: Pygmy moth (Nepticulidae) from Eocene Baltic amber (coll. T Fischer no. 5217); (a) Dorsal view; total length of imago: 2,2 mm; 
(b) Corresponding line drawing. Scale bars = 1 mm.
Zitteliana A 53 (2013) 89
Figure 3: Male pygmy moth (Nepticulidae), Bohemannia aschaueri nov. spec., from Eocene Baltic amber (SNSB-BSPG 2013 I 94); 
(a) Ventral view; total length of imago: 3 mm, body 2 mm; (b) Dorsal view; (c & d) Corresponding line drawings. Note that in dorsal view 
the right visible wing is the hindwing with a different venation than the forewing. Scale bars = 1 mm
Zitteliana A 53 (2013) 90
Figure 4: Female (?) pygmy moth (Nepticulidae) from Eocene Baltic amber (coll. T Fischer no. 5098); (a) Dorsal view; (b) Ventral view; total 
length of imago: 3 mm, body 2 mm; (c & d) Corresponding line drawings. Scale bars = 1 mm.
Zitteliana A 53 (2013) 91
clusion is positioned deep within the amber piece, 
the ventral side is hidden behind a large air bubble.
Description: Total length 2,2 mm, head 400 μm 
broad, epicranial notch visible, collar with piliform 
scales, eyes 150 μm in diameter, antennas filiform, 
length about half of the forewing, bases of antennas 
form eye caps, antenna length about 2/3 of forewing 
length with ca. 22 segments, mesonotum 300 μm 
broad, abdomen 1 mm long, end of the abdomen 
pointed as ovipositor, spur formula ?-?-4, five tar-
somers, thorax with triangular tegulas, forewings 1,9 
mm long, cilia at forewings and hindwings, wing tips 
acute, venation not visible.
Syninklusions: Ensifera (no. 5162), Dolichoderinae 
(no. 5163), Chironomidae (no. 5164) with two Leptus 
sp. (Acari) (no. 5165), Trichoptera, trichomes.
4. Discussion
Morphology and size clearly indicate that the fos-
sils described in the paper belong to the superfamily 
Nepticuloidea. Eye caps are also characteristic, even 
if similar caps are also found in other families. Among 
the Nepticuloidea, the family Opostegidae can be 
excluded since members in this family are charac-
terized by a reduced wing venation, with all the veins 
separate and relatively straight (Hoare 2000; Davis 
1999). Conversely, Nepticulidae possess branched 
and stalked veins (Davis 1999). Moreover, Oposte-
gidae are somewhat larger than extant Nepticuli-
dae (Parenti 2000) and the fossils considered here. 
Based on fore wing venation (Van Nieukerken 1986) 
the specimens with visible venation are affiliated to 
the extant genus Bohemannia Stainton, 1859. Extant 
species of Bohemannia occur in central, western 
and northern Europe, as well as in eastern Russia 
and Japan (Van Nieukerken 1986).
Skalski (1976, 1990) mentions the occurrence of 
the closely related genus Ectoedemia in Baltic am-
ber. Since this record does not contain a detailed de-
scription of the fossil and no repository information, 
the new fossils described in this paper could not be 
compared with this specimen.
The syninclusions of the Nepticulidae imagos 
represent systematic groups that are quite common 
as inclusions in Baltic amber, including Nematocera 
(Cecidomyiidae, Chironomidae), Collembola (Arthro-
pleones), Acari (Leptus sp.), Trichoptera, Formicidae 
(Dolichoderinae), Aphidae (Germaraphis sp.), and 
oak trichomes (Quercus sp.). Only Ensifera are not 
abundant fossils among these inclusions. As a result, 
taphonomy of Nepticulidae imagos in Baltic amber 
does not seem to have involved special life commu-
nities or specific processes in fossilization. It seems 
quite likely that the rarity of Nepticulidae inclusions 
in Baltic amber reflects their rarity in the amber for-
est habitat. Most extant Nepticulidae live on an-
Due to the lack of observable wing venation iden-
tification is only possible at the family level (Nepti-
culidae).
Specimen 5166
Imago of pygmy moth (Nepticulidae) 
Fig. 5
The specimen can positively be identified as be-
longing to the Nepticulidae because of the small 
body length of 2 mm and the presence of eye caps. 
However, identification is only possible at the fami-
ly level (Nepticulidae) due to the lack of observable 
wing venation Also sex is not determinable. The in-
Figure 5: Pygmy moth (Nepticulidae) from Eocene Baltic amber 
(coll. T Fischer no. 5166); (a) Dorsal view; (b) Corresponding line 
drawing. Scale bars = 1 mm.
Zitteliana A 53 (2013) 92
6. References
Conwentz H. 1886. Die Angiospermen des Bernsteins. In: HR 
Göppert, A Menge: Die Flora des Bernsteins und ihre Bezie-
hungen zur Flora der Tertiärformation und der Gegenwart, Bd. 
2, Leipzig, Engelmann.
Czeczott H. 1961. Sklad i wick flory burztynow baltyckich. Prace 
Muzeum Ziemi 4, 119–145.
Davis DR. 1999. The Monotrysian Heteroneura. In: NP Kristensen 
(Ed.), Lepidoptera, Moths and Butterflies, Volume 1: Evolution, 
Systematics and Biogeography, Handbook of Zoology, Vol. 
IV Arthropoda, Insecta, Part 35. Berlin, New York, Walter de 
Gruyter, 65–90.
Grimaldi D, Engel MS. 2005. Evolution of the Insects. Hong Kong, 
Cambridge University Press.
Hoare RJB. 2000. A new genus of primitive Nepticulidae (Lepi-
doptera) from Eastern Australia, with a revised diagnosis 
of nepticulid subfamilies. Zoological Journal of the Linnean 
Society 128, 289–317.
Kozlov MV. 1988. Paleontology of the Lepidoptera and problems of 
the phylogeny of the order Papilionoidea. In: AG Ponomarenko 
(Ed.), The Mesozoic-Cenozoic Crisis in the Evolution of Insects. 
Moscow, Academy of Sciences, 16–69.
Kristensen NP, Skalski AW. 1999. Phylogeny and Palaeontology. 
In: NP Kristensen (Ed.), Lepidoptera, Moths and Butterflies, 
Volume 1: Evolution, Systematics and Biogeography, Handbook 
of Zoology, Vol. IV Arthropoda, Insecta, Part 35. Berlin, New 
York, Walter de Gruyter, 7–26.
Labandeira CC, Dilcher DL, Davis DR, Wagner DL. 1994. Ninety-
seven million years of angiosperm-insect association: Paleo-
biological insights into the meaning of coevolution. Proceedings 
of the National Academy of Sciences USA 91, 12278–12282.
Parenti U. 2000. A guide to Microlepidoptera of Europe – Guide I. 
Turin, Museo Regionale di Scienze Naturale Torino.
Rozefelds AC. 1988. Lepidoptera mines in Pachypteris leaves 
(Corystospermaceae; Pteridospermophyta) from the Upper 
Jurassic/Lower Cretaceous Battle Camp Formation, North 
Queensland. Proceedings of the Royal Society of Queensland 
99, 77–81.
Scoble MJ. 2002. The Lepidoptera – Form, Function and Diversity. 
Oxofrd, The Natural History Museum / Oxford University Press.
Skalski AW. 1976. Les lépidoptères fossiles de l’ambre. Etat actuel 
de nos connaissances. Linneana Belgica, Pars VI, 7, 154–233.
Skalski AW. 1990. The families Nepticulidae and Thyrididae in Baltic 
amber (Lepidoptera). Proceedings of the VII. Congress of Eu-
ropean Lepidopterology, Lunz 3.–8.9. Nota Lepidopterologica, 
Supplement 1992 nr. 4, 144–145.
Sohn, JC, Labandeira CC, Davis D, Mitter C. 2012. An annotated 
catalog of fossil and subfossil Lepidoptera (Insecta : Holom-
etabola) of the world. Zootaxa 3286, 1–132.
Van Nieukerken EJ 1986. Systematics and phylogeny of holarctic 
genera of Nepticulidae (Lepidoptera, Heteroneura: Monotrysia). 
Zoologische Verhandlingen 236, 1–93.
giosperms, whereas the amber forest is assumed to 
have been dominated by an amber-producing gym-
nosperm and also oak trees. It is therefore reason-
able to assume that the Nepticulidae lived on oak or 
some of the rare, diverse angiosperm taxa which are 
known from the fossil record of Baltic amber (Con-
wentz 1886; Czeczott 1961). Woody plants of the 
orders Hamamelidae, Rosidae, and Dilleniidae are 
important host plants of extant Nepticulidae (Laban-
deira et al. 1994). For some extant species of Bohe-
mannia biology is known; these animals live on de-
cidous trees, including Malus sp. (Rosaceae), Alnus 
glutinosa (Betulaceae) and Quercus spp. (Fagaceae) 
(Van Nieukerken 1986).
5. Conclusions
The fossil Nepticulidae from Baltic amber are re-
markably similar in morphology and size to extant 
taxa. Two new fossil species are described and af-
filiated to the extant genus Bohemannia. Along with 
another three fossil imagos they provide a more 
complete insight into the evolution of Nepticulidae, 
allowing also the definition of a minimal age of the 
genus and calibration of phylogenetic trees.
Acknowledgements 
I gratefully acknowledge Tamara Kruegel (Max-
Planck-Institute for Chemical Ecology, Jena) for 
providing a translation of the Russian text of Kozlov 
(1988). Dirk Brandis and Lina Rosotta (both Kiel) are 
acknowledged for allowing and organizing the use of 
the Microlepidoptera collection housed in the Zoolo-
gical Museum Kiel. Erik van Nieukerken (Utrecht) is 
acknowledged for support with literature, and Wolf-
ram Mey (Berlin) for reviewing the manuscript.


English

Pygmy moths (Lepidoptera, Nepticulidae) from Baltic Amber (Eocene)

Open Access LMU

An International Journal  of Palaeontology and GeobiologySeries A /Reihe AMitteilungen der Bayerischen Staatssammlung für Paläontologie und GeologieMünchen 2013ZittelianaZitteliana An International Journal of Palaeontology and Geobiology Series A/Reihe A 5353Zitteliana A 53 (2013) 85Pygmy moths (Lepidoptera, Nepticulidae)  from Baltic Amber (Eocene)Thilo C. FischerLudwig-Maximilians-University Munich, Department I, Botany, Großhadernerstr. 2–4, D-82150 Planegg-Martinsried,GermanyE-mail: thilo.fischer@biologie.uni-muenchen.de Zitteliana A 53, 85 – 92 München, 31.12.2013 Manuscript received 30.09.2013; revision accepted 27.11.2013 ISSN 1612 - 412XAbstractPygmy moths (Nepticulidae, Lepidoptera) from Eocene Baltic amber are described with five new specimens, representing the only specimens beyond one described before. Some of the findings differ from Stigmellites baltica Kozlov in species and sex. They are formal-ly described as new species Bohemannia butzmanni and B. aschaueri. Phylogenetic and paleoecological aspects of Nepticulidae from Baltic amber are discussed.Key words: Baltic amber, Bohemannia, Eocene, leaf mining, Nepticulidae, pygmy mothsZusammenfassungFünf neue Arten von Zwergminiermotten (Nepticulidae, Lepidoptera) werden aus Baltischem Bernstein (Eozän) beschrieben; zusam-men mit einer bereits früher beschriebenen Form (Stigmellites baltica Kozlov) stellen sie die einzig bekannten Fossilien dieser Tiergruppe dar. Da einige der hier beschriebenen Fossilien sich von S. baltica deutlich unterscheiden, werden sie als neue Arten,  Bohemannia butz-manni und B. aschaueri, beschrieben. Phylogenetische und paläoökologische Aspekte der Nepticulidae im Baltischen Bernstein werden diskutiert.  Schlüsselwörter: Baltischer Bernstein, Bohemannia, Eozän, Blatt-minierer, Nepticulidae, Zwergminiermotten1. IntroductionNepticulidae are a monophyletic (natural) family of glossatan, monotrysian and heteroneurous Le-pidoptera (Scoble 2002). Together with the close-ly related extant family Opostegidae they form the superfamily Nepticuloidea. The family Nepticulidae comprises the smallest imagos of all Lepidoptera; the larvae mine leaves or other living plant surface tissues (petioles, bark, samaras) (Scoble 2002; Gri-maldi and Engel 2005). With its leaf mining habit, the family is of great ecological importance and also has economic impact on a number of crop plants.As a basal group of Lepidoptera phylogeny of Nepticulidae is of concern, but members of the fa-mily are extremely rare as fossils (Sohn et al. 2012). Furthermore, most reports refer only to the typical traces of mining activity found in fossil leaves (most-ly described as Stigmellites; see Sohn et al. 2012), the oldest of which come from the Mid-Cretaceous Dakota formation and are suggested to have been produced by Stigmella and Ectoedemia (Nepticuli-dae) (Labandeira et al. 1994). Lepidopterous mines in leaves of the seed fern Pachypteris from the Aus-tralian Upper Jurassic/Lower Cretaceous might be an even older evidence for Nepticulidae (Rozefelds 1988). Undescribed putative nepticulid imagos are known from Cretaceous Siberian amber (mentioned in Grimaldi and Engel 2005) and from Canadian Late Cretaceous amber (mentioned in Kristensen and Skalski 1999). The evolution of Nepticulidae and other lepidopterous leaf miners is supposed to be closely linked to angiosperm radiation and hence host plant evolution (Labandeira et al. 1994; Grimaldi and Engel 2005). A first fossil nepticulid imago was mentioned by Skalski (1976, 1990) from Eocene Bal-tic amber and affiliated to Ectoedemia sp. without description or even information on its deposition. Kozlov (1988) described a female imago as Stigmel-lites baltica. Some of the nepticulid imagos studied Zitteliana A 53 (2013) 86tip, three-segmented downwards-pointing maxilla-ry palps, very short and blunt proboscis, abdomen slightly flat with tips of valves visible, no epiphysis, coxa and femur with transversal rows of scales, spur formula 0-2-4, middle hind spurs at half the length of hind tibia, five tarsomers, thorax with triangular tegu-las, cilia at forewings and hindwings, twice as long at the hindwings, wing tips acute, venation consisting of sc, r1, r2-3, r4, r5, m1, m2, cu, and a; m1, m2 and cu distally diverging from r5.Differential diagnosis: In comparison to Stigmel-lites baltica Kozlov 1988 this specimen differs in the following characters: wing venation, (sex,) length of antennas, tibial spurs, hindwings broader, coxa and femur with transversal rows of scales. Affiliation to the extant genus Bohemannia is mainly supported by the structure of the venation and the presence of a collar of scales.Description: Total length 2,2 mm, head 400 μm broad, vertex smooth, frontal tuft, epicranial notch visible, collar with piliform scales, eyes 130 μm in diameter, bases of antennas forming eye caps, fili-form antennas with a length of about ¾ of forewing length with 30–35 segments (partly invisible), labial palps 150 μm with two segments, last segment with a thick middle part and a fine tip, pointed upwards, maxillary palps 250 μm with three visible segments, folded downwards, proboscis very short (100 μm) and blunt, mesonotum 400 μm broad, abdomen 870 μm long, slightly flat and with irregularly preserved scales, male genitalia (tips of valves) visible, no epi-physis at front legs, coxae and femurs with transver-sal rows of scales, spur formula 0-2-4, middle hind spurs positioned at half the length of the hind tibia, five tarsomers, thorax with triangular tegulas, forew-ings 1,8 mm long, cilia at forewings and hindwings, twice as long at the hindwings (up to 500 μm), scale covering only preserved at the proximal parts of the wings, wing tips acute, venation consisting of sc, r1, r2+3, r4+5, m1, m2, cu, and a; m1, m2 and cu di-verging from r4+5 in the very distal part of the latter. There are no syninclusions preserved in this piece of amber.Specimen 5217Imago of pygmy moth (Nepticulidae) Fig. 2Description: Total length 2,2 mm, head 400 μm broad, frons and vertex smooth, collar without sca-les, diameter of eyes ca. 150 μm, eye caps at bases of antennas, filiform antennas with ca. 20 segments and a length half of that of forewings, maxillary (or labial?) palps bent and with 2 visible segments, other palps not visible, proboscis very small, abdomen length ca. 800 μm, mesonotum ca. 450 μm broad, spur formular 0-2-4, forewing length 1,9 mm, cilia at wings.here clearly differ from Stigmellites baltica, and also represent the first male specimens discovered from Baltic amber.2. Material and MethodsThe amber pieces were purchased from trusted amber dealers. The inclusions were photographed with a binocular microscope (Zeiss Stemi 2000-C) to which is attached a digital camera system (Lei-ca DFC 320, software LAS 4.1) (specimens 5058, 5166); other images were obtained with the inclusi-ons (specimens 5198, 5199, 5217). The specimens were preserved by treatment with an acryl varnish and are kept in the author’s collection at constant temperature in plastic clip bags within metal boxes excluding oxygen and light. However, transfer to a public collection (Bayerische Staatsammlung für Pa-läontologie und Geologie, München, Deutschland) is envisaged.3. Systematic paleontologyA total of five new nepticulid imagos have been discovered in the Eocene Baltic amber pieces.Lepidoptera Linnaeus, 1758Superfamily Nepticulidea Stainton, 1854Family Nepticulidae Stainton, 1854Genus Bohemannia Stainton, 1859Bohemannia butzmanni nov. spec.Fig. 1Holotypus: Specimen ex coll. Fischer no. 5058, Fig. 1Locus typicus: Amber mine of Yantarni, GUSStratum typicum: “Blaue Erde” (Upper Eocene – Lower Oligocene)Etymology: The name is given to acknowledge the numerous paleontological activities of Mr. Rainer Butzmann (Munich, Germany)Repository: Bayerische Staatssammlung für Palä-ontologie und Geologie, Munich, Germany; accessi-on number SNSB-BSPG 2013 I 93Diagnosis: Very small moth with broad head, ver-tex smooth but with frontal tuft, epicranial notch visible, piliform scales at collar, bases of antennas form eye caps, filiform antenna, length about ¾ of forewing length with 30–35 segments (partly invisi-ble), two-segmented upwards-pointed labial palps, last segment with a thick middle part and a fine Zitteliana A 53 (2013) 87Figure 1: Male pygmy moth (Nepticulidae), Bohemannia butzmanni nov. spec., from Eocene Baltic amber (SNSB-BSPG 2013 I 93); (a) Dorsal view; (b) Ventral view; total length of imago: 2,2 mm; (c) Corresponding line drawings with organ identification. Scale bars = 1 mm.Zitteliana A 53 (2013) 88rewing length 1,95 mm, heteroneurous, wing tips pointed, hindwings not broad, cilia at front and hind wings, antennas with at least 26 segments (but an-tennas broken), labial palps three-segmented, ma-xillary palps with two- (or three-) visible segments, valves.Differential diagnosis: Venation same as 5058 (hence different to Stigmellites baltica), but frons with erect scales and galeae of proboscis slightly separa-ted.  Preparation of genitals as performed in studies with extant Nepticulidae is not possible. Affiliation to the extant genus Bohemannia is due to the similarity of the venation.Specimen 5198Imago of pygmy moth (Nepticulidae) Fig. 4Total length with legs 2,8 mm, head 400 μm broad, vertex with erect scales, collar with bundles of pili-form scales behind antennas, eyes 180 μm in dia-meter, eye caps, filiform antennas short (2/5 of front wing length) with 19 segments and pointed at the tip, labial palps two-segmented and downwards-poin-ting, maxillary palps not visible, proboscis very short and with slightly separated galeae, mesonotum 400 μm broad, abdomen 1100 μm long, no epiphysis, spur formula 0-2-4, tibias with scales, middle hind spurs positioned at 2/5 of length from basis of femur, five tarsomers, forewing length 1,8 mm, wings with scales preserved, cilia and longitudinal scales up to 200 μm long at wing edges, wing tips acute.This specimen may be conspecific with specimen 5058, showing the same fore wing venation and also a smooth vertex.Syninclusions: Cecidomyiidae (no. 5218), Germa-raphis sp. (no. 5219).Genus Bohemannia Stainton, 1859Bohemannia aschaueri nov. spec.Holotypus: Specimen ex coll. Fischer no. 5199, Fig. 3Locus typicus: Amber mine of Yantarni, GUSStratum typicum “Blaue Erde” (Upper Eocene – Lower Oligocene)Etymology: The name is given to acknowledge the numerous paleontological activities of Birgitt and Karl Aschauer (Waidhofen an der Ybbs, Austria).Repository: Bayerische Staatssammlung für Palä-ontologie und Geologie, Munich, Germany; accessi-on number SNSB-BSPG 2013 I 94Syninclusions: Trichomes, Arthropleones.Diagnosis: Eye caps with scales, trochanter with rows of scales, proboscis very small, galeae sepa-rated, spur formula 0-2-4, frons with erect scales, vertex smooth (with scales?), tegulas triangular, fo-Figure 2: Pygmy moth (Nepticulidae) from Eocene Baltic amber (coll. T Fischer no. 5217); (a) Dorsal view; total length of imago: 2,2 mm; (b) Corresponding line drawing. Scale bars = 1 mm.Zitteliana A 53 (2013) 89Figure 3: Male pygmy moth (Nepticulidae), Bohemannia aschaueri nov. spec., from Eocene Baltic amber (SNSB-BSPG 2013 I 94); (a) Ventral view; total length of imago: 3 mm, body 2 mm; (b) Dorsal view; (c & d) Corresponding line drawings. Note that in dorsal view the right visible wing is the hindwing with a different venation than the forewing. Scale bars = 1 mmZitteliana A 53 (2013) 90Figure 4: Female (?) pygmy moth (Nepticulidae) from Eocene Baltic amber (coll. T Fischer no. 5098); (a) Dorsal view; (b) Ventral view; total length of imago: 3 mm, body 2 mm; (c & d) Corresponding line drawings. Scale bars = 1 mm.Zitteliana A 53 (2013) 91clusion is positioned deep within the amber piece, the ventral side is hidden behind a large air bubble.Description: Total length 2,2 mm, head 400 μm broad, epicranial notch visible, collar with piliform scales, eyes 150 μm in diameter, antennas filiform, length about half of the forewing, bases of antennas form eye caps, antenna length about 2/3 of forewing length with ca. 22 segments, mesonotum 300 μm broad, abdomen 1 mm long, end of the abdomen pointed as ovipositor, spur formula ?-?-4, five tar-somers, thorax with triangular tegulas, forewings 1,9 mm long, cilia at forewings and hindwings, wing tips acute, venation not visible.Syninklusions: Ensifera (no. 5162), Dolichoderinae (no. 5163), Chironomidae (no. 5164) with two Leptus sp. (Acari) (no. 5165), Trichoptera, trichomes.4. DiscussionMorphology and size clearly indicate that the fos-sils described in the paper belong to the superfamily Nepticuloidea. Eye caps are also characteristic, even if similar caps are also found in other families. Among the Nepticuloidea, the family Opostegidae can be excluded since members in this family are charac-terized by a reduced wing venation, with all the veins separate and relatively straight (Hoare 2000; Davis 1999). Conversely, Nepticulidae possess branched and stalked veins (Davis 1999). Moreover, Oposte-gidae are somewhat larger than extant Nepticuli-dae (Parenti 2000) and the fossils considered here. Based on fore wing venation (Van Nieukerken 1986) the specimens with visible venation are affiliated to the extant genus Bohemannia Stainton, 1859. Extant species of Bohemannia occur in central, western and northern Europe, as well as in eastern Russia and Japan (Van Nieukerken 1986).Skalski (1976, 1990) mentions the occurrence of the closely related genus Ectoedemia in Baltic am-ber. Since this record does not contain a detailed de-scription of the fossil and no repository information, the new fossils described in this paper could not be compared with this specimen.The syninclusions of the Nepticulidae imagos represent systematic groups that are quite common as inclusions in Baltic amber, including Nematocera (Cecidomyiidae, Chironomidae), Collembola (Arthro-pleones), Acari (Leptus sp.), Trichoptera, Formicidae (Dolichoderinae), Aphidae (Germaraphis sp.), and oak trichomes (Quercus sp.). Only Ensifera are not abundant fossils among these inclusions. As a result, taphonomy of Nepticulidae imagos in Baltic amber does not seem to have involved special life commu-nities or specific processes in fossilization. It seems quite likely that the rarity of Nepticulidae inclusions in Baltic amber reflects their rarity in the amber for-est habitat. Most extant Nepticulidae live on an-Due to the lack of observable wing venation iden-tification is only possible at the family level (Nepti-culidae).Specimen 5166Imago of pygmy moth (Nepticulidae) Fig. 5The specimen can positively be identified as be-longing to the Nepticulidae because of the small body length of 2 mm and the presence of eye caps. However, identification is only possible at the fami-ly level (Nepticulidae) due to the lack of observable wing venation Also sex is not determinable. The in-Figure 5: Pygmy moth (Nepticulidae) from Eocene Baltic amber (coll. T Fischer no. 5166); (a) Dorsal view; (b) Corresponding line drawing. Scale bars = 1 mm.Zitteliana A 53 (2013) 926. ReferencesConwentz H. 1886. Die Angiospermen des Bernsteins. In: HR Göppert, A Menge: Die Flora des Bernsteins und ihre Bezie-hungen zur Flora der Tertiärformation und der Gegenwart, Bd. 2, Leipzig, Engelmann.Czeczott H. 1961. Sklad i wick flory burztynow baltyckich. Prace Muzeum Ziemi 4, 119–145.Davis DR. 1999. The Monotrysian Heteroneura. In: NP Kristensen (Ed.), Lepidoptera, Moths and Butterflies, Volume 1: Evolution, Systematics and Biogeography, Handbook of Zoology, Vol. IV Arthropoda, Insecta, Part 35. Berlin, New York, Walter de Gruyter, 65–90.Grimaldi D, Engel MS. 2005. Evolution of the Insects. Hong Kong, Cambridge University Press.Hoare RJB. 2000. A new genus of primitive Nepticulidae (Lepi-doptera) from Eastern Australia, with a revised diagnosis of nepticulid subfamilies. Zoological Journal of the Linnean Society 128, 289–317.Kozlov MV. 1988. Paleontology of the Lepidoptera and problems of the phylogeny of the order Papilionoidea. In: AG Ponomarenko (Ed.), The Mesozoic-Cenozoic Crisis in the Evolution of Insects. Moscow, Academy of Sciences, 16–69.Kristensen NP, Skalski AW. 1999. Phylogeny and Palaeontology. In: NP Kristensen (Ed.), Lepidoptera, Moths and Butterflies, Volume 1: Evolution, Systematics and Biogeography, Handbook of Zoology, Vol. IV Arthropoda, Insecta, Part 35. Berlin, New York, Walter de Gruyter, 7–26.Labandeira CC, Dilcher DL, Davis DR, Wagner DL. 1994. Ninety-seven million years of angiosperm-insect association: Paleo-biological insights into the meaning of coevolution. Proceedings of the National Academy of Sciences USA 91, 12278–12282.Parenti U. 2000. A guide to Microlepidoptera of Europe – Guide I. Turin, Museo Regionale di Scienze Naturale Torino.Rozefelds AC. 1988. Lepidoptera mines in Pachypteris leaves (Corystospermaceae; Pteridospermophyta) from the Upper Jurassic/Lower Cretaceous Battle Camp Formation, North Queensland. Proceedings of the Royal Society of Queensland 99, 77–81.Scoble MJ. 2002. The Lepidoptera – Form, Function and Diversity. Oxofrd, The Natural History Museum / Oxford University Press.Skalski AW. 1976. Les lépidoptères fossiles de l’ambre. Etat actuel de nos connaissances. Linneana Belgica, Pars VI, 7, 154–233.Skalski AW. 1990. The families Nepticulidae and Thyrididae in Baltic amber (Lepidoptera). Proceedings of the VII. Congress of Eu-ropean Lepidopterology, Lunz 3.–8.9. Nota Lepidopterologica, Supplement 1992 nr. 4, 144–145.Sohn, JC, Labandeira CC, Davis D, Mitter C. 2012. An annotated catalog of fossil and subfossil Lepidoptera (Insecta : Holom-etabola) of the world. Zootaxa 3286, 1–132.Van Nieukerken EJ 1986. Systematics and phylogeny of holarctic genera of Nepticulidae (Lepidoptera, Heteroneura: Monotrysia). Zoologische Verhandlingen 236, 1–93.giosperms, whereas the amber forest is assumed to have been dominated by an amber-producing gym-nosperm and also oak trees. It is therefore reason-able to assume that the Nepticulidae lived on oak or some of the rare, diverse angiosperm taxa which are known from the fossil record of Baltic amber (Con-wentz 1886; Czeczott 1961). Woody plants of the orders Hamamelidae, Rosidae, and Dilleniidae are important host plants of extant Nepticulidae (Laban-deira et al. 1994). For some extant species of Bohe-mannia biology is known; these animals live on de-cidous trees, including Malus sp. (Rosaceae), Alnus glutinosa (Betulaceae) and Quercus spp. (Fagaceae) (Van Nieukerken 1986).5. ConclusionsThe fossil Nepticulidae from Baltic amber are re-markably similar in morphology and size to extant taxa. Two new fossil species are described and af-filiated to the extant genus Bohemannia. Along with another three fossil imagos they provide a more complete insight into the evolution of Nepticulidae, allowing also the definition of a minimal age of the genus and calibration of phylogenetic trees.Acknowledgements I gratefully acknowledge Tamara Kruegel (Max-Planck-Institute for Chemical Ecology, Jena) for providing a translation of the Russian text of Kozlov (1988). Dirk Brandis and Lina Rosotta (both Kiel) are acknowledged for allowing and organizing the use of the Microlepidoptera collection housed in the Zoolo-gical Museum Kiel. Erik van Nieukerken (Utrecht) is acknowledged for support with literature, and Wolf-ram Mey (Berlin) for reviewing the manuscript.

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Pygmy moths (Lepidoptera, Nepticulidae) from Baltic Amber (Eocene)

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