Publication date: December 2021
113 pages, over 200 illustrations
- New graphic illustrations
- Colour, large-format photographs
- Detailed microscopic illustrations, with
- A book with a similar concept has ne-
ver been published in Europe on the spe-
Book price: 49 Euro + postage
The address of the order is:
This book, based on the author's own research and following a crical analysis of the lit-
erature, presents an account of the Pterophoridae of the Balkan Peninsula. The geo-
graphical and Balkan distribuon of the 94 species is described. The biology of the spe-
cies is analysed, including the host plants, the preferred and potenal habitats. Based on
data from collecons and literature, the ﬂight mes and number of generaons of the
species are given. Several species are diﬃcult to name with certainty, and generalized
drawings and diagrams are provided to assist idenﬁcaon.
No similar book has ever been produced in Europe. The habitus and genitalia illustraons
are detailed and of appropriate size. The author has tried to present details and speciﬁc
features accurately. Therefore, the book will not only be useful for the idenﬁcaon of
Balkan Pterophoridae, but also for the study of Central European and European species
in general. The more than 200 illustraons have been produced by the author using a
unique digital technique.
This paper, based on the author's own research and following a critical analysis of the
literature, presents an account of the Pterophoridae of the Balkan Peninsula. The
geographical and Balkan distribution of the species is described. The biology of the
species is analysed, including the host plants, the preferred and potential habitats. Based
on data from collections and literature, the flight times and number of generations of the
species are given. Several species are difficult to name with certainty, and generalised
drawings and diagrams are provided to assist identification. Maps of the distribution
patterns of some rare species are included. The work presented here is a summary of the
author's earlier work in the Balkans, which will be followed by the processing of new
collections not yet examined.
Worldwide, nearly 1500 species of Pterophoridae have been described, in more than 90
genera. This study summarises for the first time the species that inhabit the Balkan
Knowledge of biodiversity among Pterophoridae in the Balkan Peninsula is still far
from complete. Compared to the neighbouring Pannonian ecoregions, the Pterophoridae
fauna in Balkan is very poorly known. It is not easy to give a definition of the Balkan
region, or South-Eastern Europe as it is more popularly known today, either
geographically or in a geopolitical sense. Geographically, it refers roughly to the region
bounded by the rivers Sava and Danube, the Black Sea, and the Mediterranean Sea.
In the broadest geopolitical sense, the Balkan Peninsula comprises Albania, Bulgaria,
Greece, Romania, Turkey, and five other countries, parts of the former Yugoslavia: Bosnia
and Herzegovina, Croatia, Macedonia, Serbia and Montenegro and Slovenia.
During the last Ice Age, the Balkan Peninsula was a refuge area for many species. They
have survived there, due to the presence of suitable niches (habitats) in the great variety
The northern border of the Balkan Peninsula is generally considered to be the Sava and
Danube rivers. For practical reasons, the border of the river Sava is extended here to the
river Drava, the Hungarian border.
Thus, the territory of Croatia is more homogeneous for the purpose of the study. The
Peninsula is bordered to the west by the Adriatic Sea, to the southwest by the Ionian Sea,
to the south by the Aegean Sea and the Mediterranean Sea, to the east by the Black Sea,
and to the southeast by the Dardanelles and Bosporus Straits, which separate the Peninsula
from the Anatolian part of Turkey. In this paper, Romania and Slovenia are not included
among the Balkan countries.
Institute. The identiﬁcation of these is in progress. Several Hungarian collectors, including
Imre Balogh † H-Budapest, Levente Ábrahám and H-Kaposvár have visited the Balkan
Peninsula. For several decades, Ernst Arenberger († 2020, Vienna) was consulted
intensively by the author, and gave him a lot of very important information, for which he is
According to Eastwood (2004), the vegetation of southeast Europe and the Balkans is
strongly inﬂuenced by region, and may be divided into two distinct communities:
Mediterranean vegetation community in the south, comprising elements of warm
deciduous forest, and a Central European (Continental) plant community in the north, with
elements of cold deciduous forest; the line on map broadly shows the boundary between
these two major vegetation communities. The Mediterranean vegetation community
consists of evergreen forest, Pinus halepensis, evergreen Quercus spp. and Juniperus, with
Mediterranean deciduous forest, deciduous Quercus spp., Carpinus orientalis, Fraxinus
ornus, Ostrya carpinifolia, Pistacia terebinthus and Acer spp. in inland and cooler areas.
This unit is found from sea level to~700 m a.s.l., while from 700-1700 m a.s.l. coniferous
forests of Abies and Juniperus are found, with sub-alpine and alpine vegetation from 1700-
3000 m a.s.l. Central European (Continental) plant community comprising mixed oak-
hornbeam forest of Quercus and Carpinus with Fagus, Ulmus, Tilia, Fraxinus and Acer is
found from 0-700 m a.s.l. inﬂuenced by Climate, altitude, soils, and human disturbance.
Higher, with montane forest comprising mostly Fagus and above these coniferous forests
of Abies, Picea and Pinus from 700-1700 m a.s.l. Subalpine and alpine vegetation together
Fig 3. The northern and eastern boundaries of the study area
(red line), with the names of the states.
Material and methods
This paper is based on unpublished data from our own research and published data gleaned
from the literature. The presence of an expected, potentially occurring species, [Stenoptilia
annadactyla Sutter, 1988] is predicted, placed here in square brackets. It is thought to be
important to include potential species in the catalogue because it draws attention to the
possibility their discovery and a more thorough analysis of problem species groups. For
several species faunistic and bionomic observations of "plume moth" species from the
Carpathian Basin (mainly Hungary), are reported, but which are indirectly related to the
Balkans. This provides a better overview of the faunal genetics and faunal history of the
two neighbouring areas.
Genitalia dissections were made in accordance with Robinson (1976). Some of the
genitalia were mounted in Euparal on slides; others are preserved in micro-vials ﬁlled with
glycerol. Genital analysis of worn, damaged specimens of Stenoptilia was performed using
the simple and rapid method of Wanke and Rajaei (2018).
Fig. 7. Daytime collection and observation, in Serbia, along the Danube River (a), setting
up a light tower (160 W HLMI) for night collection (b), examination of moths and genitalia
with a stereo microscope (c), evening and night collection with 125 W mercury vapour
lamp (d). The pictures show the author (Photo: Irma F. Visnyei).
Alexander K. Drenowski
Fig. 6. Researchers who have contributed to the knowledge of the Balkan Pterophoridae
fauna (schematic overview). See their work in the literature.
Jaroslaw Buszko Cees Gielis Imre Fazekas
15. Platyptilia nemoralis Zeller, 1841
Distribution. Platyptilia nemoralis has spread from Japan through Russia to the British
Isles. It is very local and rare in the Mediterranean Region. Recently also detected in Iran
(Alipanah & Gielis, 2010).
Occurrence in the Balkan Peninsula. Bulgaria, North Macedonia.
Biology. Flight period early July to end of August in a single generation. Larvae
narrowly oligophagous on Senecio fuchsii, S. ﬂuviatilis, S. sarracenicus, S. sulphureus, and
S. nemorensis. Shoot and stem borer, several larvae may be present on one shoot. Stems
enlarge in response to feeding. Frass is extruded through a hole in stem. The species lives in
damp places in forests and swamps, and along streams.
Remarks. P. nemoralis has only been found singly in the Balkans.
16. Platyptilia tesseradactyla (Linnaeus, 1761)
Distribution. Holarctic species with numerous fragmented populations, for example in
the USA, Canada, and the central regions of Eurasia.
Occurrence in the Balkan Peninsula. Bulgaria, Greece, North Macedonia.
Biology. Flight period in many regions starts as early as the end of May, with a peak
period in June and July, the moth is univoltine ﬂies only in June and July. It is probably two
generations, but this needs to be investigated further the last specimens collected in early
August. Larvae oligophagous on and young leaves of Helichrysum arenarium and
Antennaria dioica. This species has a wide ecological plasticity, which extends from north
of the Arctic Circle to the warmth of the Mediterrenean climate in the south. It favours birch
woodland, hairgrass meadows, hilly and mountain meadows, and can also be found in
rocky and sandy meadows.
Remarks. In continental Europe, P. tesseradactyla prefers sandy landscapes and
clearings and edges of pine forests. In mountain grasslands it reaches altitudes of 2200-
2500 m, where of its preferred host plants is abundant.
Figs. 19-20. Comparison of wing patterns, highlighting the hindwing (scale-tooth);
19 Platyptilia nemoralis, 20 P. tesseradactyla
23. Stenoptilia atlanticola Zerny, 1936
Distribution. Known only from Bulgaria (Fazekas, 1991) and Morocco.
Occurrence in the Balkan Peninsula. Bulgaria (Osogovo). See Fazekas (1991, Abb.
1.), gen. prep. Fazekas. Nr. 2196.
Biology. Specimens were collected in June and July, but the ﬂight period needs further
investigation The hostplant was stated by Arenberger (1995) to be unknown, but according
to Bigot et al. (2009) it is Nepeta stachyoides. It probably feeds also on other Neptea
species. In the Atlas Mountains, it reaches an altitude of 2700 m.
Remarks. The original description of the taxon atlanticola, as Stenoptilia
stigmatodactyla f. atlanticola Zerny, 1936. Mem. Soc. Sci. nat. phys. Maroc. 42:130.
Arenberger (1984) considered it to be a valid species in his revision. The genital structure
indicates that atlanticola belongs to the Stenoptilia pterodactyla complex. In external
features, it diﬀers from S. pterodactyla chieﬂy in the presence of the large crevice that sits
directly on the crevice. In the male genitalia, the formation of the cucullus is decisive for the
distinction. Based on Moroccan specimens, Arenberger (1984, Figs. 1-2) illustrated the
male and female genitalia as shown in Figs. 27).
Fig. 27. Stenoptilia atlanticola; 27a male genitalia, Bulgaria (Fazekas 1991), 27b male
genitalia, Morocco (Arenberger 1984), 27c female genitalia Morocco (Arenberger
27a 27b 27c
Fig. 28. Male (28a) and female (28b)
genitalia of Steoptilia pterodactyla
28b ♀ Fig. 29. Forewing pattern of Stenoptilia
atlanticola (29a) and S. pterodactyla
(29b). The basic colour of the wings is
30. Stenoptilia lutescens (Herrich-Schäﬀer,1855)
Distribution. Reported from France, Spain, Switzerland, Italy, Sardinia, Macedonia,
Serbia, and Albania (Gielis,2003), but otherwise little is known.
Occurrence in the Balkan Peninsula. Albania (“Krrabë” South-east of Tirana),
Bosnia-Herzegovina (Vučjabaja), North Macedonia, Serbia.
Biology. Flight period in mountainous regions between700 and 2500 m from May to
early November, probably in two generations. Larva monophagous on Gentiana lutea, G.
clusii. Reported on shoots, leaves, and stems.
Remarks. The pattern of the forewing makes it easily to mistake for S. coprodacyta,
and it is always necessary to examine the genitalia. According to Arenberger (1988), the
type specimen of Stenoptilia lutescens has been searched for in all museums and
collections without success. Some of the type specimens of species described by Herrich-
Schäﬀer were lost when his Collection was dispersed among private collections. According
to the original description there is no doubt that S. lutescens is a diﬀerent species from S.
coprodacytla, from which it diﬀers by the larger size and the darkening of the forewing. The
genitalia are also diﬀerent. The wing pattern and genitalia of the sister species
lutescens/coprodactyla are shown in Figures 36-37.
Fig. 36. Wing pattern of Stenoptilia coprodactya (36a) and S. lutescens (36b)
Fig. 37. Male genitalia of Stenoptilia coprodactya (37a). Male genitalia of S. lutescens
(37b); sacculus extension small, uncus stout and short, anellus slender and ﬁne tipped,
1/2 lenght of tegumen.
37a ♂ 37b ♂
woods, hedge banks, grassland, rock outcrops, upland screes, road verges, railway banks
and waste ground, found on most soil types except the most impoverished. S. pterodactyla
is also widespread in the Balkans and in many parts of the Palaearctic.
Remarks. Personal research has shown this species to be one of the most widespread
plume moths in the Balkan Peninsula. Fresh specimens can be distinguished by their white
costal cilia and ochreous wing colour from the similar Stenoptilia bipunctidactyla, which
has dark costal cilia and grey-brown wings. These diﬀerences may be less clear in worn
specimens. Examination of the genitalia is always important, and this species has been
misidentiﬁed in many collections.
Fig. 41. Diagnostic characters (indicated) of
species. 41a Stenoptilia pelidnodactyla, 41b
S. pneumonanthes, 41c S. pterodactyla
Fig. 42. The genitalic characters playing a decisive role in much Pterophorid's taxonomy
and identiﬁcation. 42a ♂ genitalia, Stenoptilia pterodactyla, 42b ♂ S. pelidnodactyla,
42c ♂, S. pneumonanthes. Females, antrum; 42d S. pterodactyla, 42e S. pelidnodactyla,
42f S. pneumonanthes
Buszkoiana Koçak, 1981
42. Buszkoiana capnodactyla (Zeller, 1841)
Distribution. Known from Denmark, Poland, southern Germany, the southern tip of
the Netherlands, Belgium, Italy, Austria, the Czech Republic, Slovakia, Hungary, Romania
(Locus typicus: “Bánát”), Bulgaria, North Macedonia, Albania, Ukraine, and southern
Occurrence in the Balkan Peninsula. Albania, Bulgaria, North Macedonia,
Biology. Flight period in June and July. Larvae monophagous on Petasites hybridus.
According to the literature, the habitat requirements coincide with those of Hydraecia
petasites (Doubleday, 1847) (Noctuidae), and larvae of both species can be found together
in hostplant (Finke et al., 1998).
Remarks. The preferred habitats are alder swales and the more humid highland
vegetation that accompanies streams. These habitats have been severely depleted by
climate change and watercourse regulation. The species is gradually disappearing from
highly fragmented habitats.
Figs 46-48. Wings characters of the species (indicated): 46 , Paraplatyptilia metzneri
third lobe of hindwing with distally placed scale-tooth, costal triangle well-developed,
47 , 48 , colour brown-grey, and Amblyptilia acanthadactyla Amblyptilia punctidactyla
specked with wihite scales.
Fig. 49. Buszkoiana capnodactyla,
scale-tooth on third lobe of
hindwing at apex . The costal
triangle is blurred, preceded by a
bright spot (indicated).
costal triangle, variable
Fig. 55. Forewing pattern of Capperia species, showing main morphological (indicated).
fringe with scale-tooth
of 1st lobe
of 2nd lobe
Figs 56-62. Morphological characters of Balkanian Capperia species (forewings,
indicated): 56 C. celeusi, colour from brown to yellow-brown; 57 C. fusca, the white
cross-stripes reduced, almost absent; 58 C. hellenica, the lobes of the forewings are
slender, the white crossbars are broad but variable; 59 C. maratonica, the white cross-
stripes and white apical border are vigorous, the fringe with scale-tooth are dark brown; 60
C. polonica, colour of the forewing dark brown, on the anal ange of 1st lobe are teh fringe
very dark. 61 C. trichodactyla, similar to the previous species, but the fringes are lighter,
the white cross-stripes are narrower; 62 C. washbourni, forewing as in C. celeusi, but the
white cross-stripes of the forewing broader, but the base colour is usually a much darker