15 - Olah - U 2017

FOLIA ENTOMOLOGICA HUNGARICA
R OVARTA N I KÖZ L E M É N Y E K
Volume 78 2017 pp. 111–255
Trichoptera endemic in the Carpathian Basin and the adjacent areas
János Oláh
H-4032 Debrecen, Tarján u. 28, Hungary. E-mail: [email protected]
Abstract – Original text of taxonomic diagnosis and data of type specimens are collected and cited
for each Trichoptera species endemic or subendemic in and around the Carpathian Basin, together
with taxonomic history of relevant changes in nomeclature. The total number of the described cad-
disfly species endemic in or around the Carpathian Basin is 263. By describing 78 new incipient
phylogenetic sibling species a surprisingly high ratio of unknown caddisfly diversity were detected
during only a few years and with only moderate collecting effort. This clearly indicates the highly
depressed state of the western contemporary taxonomy. The recently diverged taxa are products
of Pleistocene speciation process during isolated integration in crenon/epicrenon habitats on sky
islands of high mountain ranges. Rhyacophila (32 species) and Drusus (62 species) genera and the
Chaetopterygini (56 species) and Stenophylacini (42 species) tribes are the groups richest in en-
demic species diverged in spring and spring stream habitats of high elevations.
Key words – Balkan Peninsula, caddisfl ies, Carpathian Basin, taxonomic history
INTRODUCTION
During the last few years we have conducted studies on speciation proc-
esses acquiring intuitive insight into the contemporary entity divergences simply
by empirical detecting and describing over 120 young incipient new caddisfly
species in the European Trichoptera fauna (Oláh et al. 2012, 2013a, b, c, 2014,
2015a, b, 2017). This amount of new taxa, discovered easily with moderate effort
in the so-called “intensively studied and well known” faunal region, clearly indi-
cates that our knowledge is still far from complete.
Our method is based on the adaptive speciation traits, integrated in sexual
processes, and enabled us to delimit and delineate recently diverged or even con-
temporary diverging phylogenetic species in sibling species complexes (Oláh et al.
2015a, 2017). Most of these young species are the product of the Pleistocene spe-
ciation. This integrative process has been realised in isolated crenon/hypocrenon
habitats of sky islands of high elevations and highly influenced by the complex-
ity of European high mountain systems (Schmitt 2009). Just in and around the
DOI: 10.17112/FoliaEntHung.2017.78.111 Folia ent. hung. 78, 2017

112 J. Oláh
Carpathian Basin sensu lato we have collected and described 78 new endemic incip-
ient phylogenetic species from the total of 263 endemic or subendemic Carpathian
species. They populate the spring habitats in sky islands of high mountain ranges.
In order to distinguish these young sibling species applying the discovered
sensitive speciation traits we had to examine old historical specimens of several
unsettled taxa deposited and curated in various European and North American
museums. The study on the taxonomic history of these old taxa has been resulted
in this revision of the endemic caddisfl ies of the Carpathian Basin. The terri-
tory under revision is the Carpathian Basin sensu lato. Endemic species under
the present taxonomic revision are not restricted to the Carpathian caddisfl ies
sensu stricto, i.e. to species inhabiting sorroundings of the Carpathian Mountains.
The taxonomy of endemic or subendemic Trichoptera species having distribu-
tion centre in Carpathian Basin or described from the Carpathian Basin and the
adjoining Balkan and Alps territories are revised. In geographic terms the area
of the Carpathian Basin sensu lato covers the Danube drainage basin from River
Morva nearby the Dévény Gate where Danube River enters the Carpathian Basin
to River Morava nearby the Iron Gate where Danube River leaves the Carpathian
Basin. In ecological terms, taking into account the dispersion habitats and dis-
tribution capacities of the most active caddisfly species, the environmental ter-
ritory of the Carpathian Basin under revision covers the entire drainage basin
and extended to running waters flowing to adjoining drainage basins possibly
hundreds of kilometers over the watershed.
In this revision of the endemic Trichoptera of the Carpathian Basin sensu
lato I have collected data of the taxonomic history of listed species.
Diagnosis (if available) together with the locality data of type specimens
are given by citing the relevant information from the original publications in the
original language of the descriptions. Species diagnoses, frequently hardly acces-
sible, give a brief summary of specific character states or character combinations
for both specialist and non-specialist readers, together with the close relatives or
siblings of the species groups or species complexes. This essence of delineation
or delimitation of the endemic species in the Carpathian Basin, accumulated in a
single paper, may help scientists and conservationists directly by giving practical
guidance in nature conservation practices.
The taxonomic position of some Trichoptera subspecies is not studied yet.
Reviewing the present taxonomic status of the Carpathian Trichoptera species we
have accepted the subspecies status of the biological species concept. However,
we believe that similarly to all of our previous systematic investigations on the
fine structures of speciation traits, the search of initial split criteria in these sub-
species reveals in most cases the natural status of incipient sibling species elabo-
rated in the phylogenetic species concept.
Folia ent. hung. 78, 2017

Trichoptera endemic in the Carpathian Basin and the adjacent areas 113
MATERIAL
Thinking of population concept in the new taxonomy requires more elabo-

rated field collecting strategies. To collect many specimens from many popula-
tions is the prime target of any research project aimed to find the first signs of
reproductive isolation, to search species boundaries, to delimit closely related
incipient taxa, and to recognise the young phylogenetic species. Biodiversity re-
search and conservation are badly limited by the lack of population field sam-
pling, which is expensive. Staggering in the deprived discipline of taxonomy
and suffering the lack of adequate collecting we have been forced to outline the
principles and practice of cooperation to put together what we have (Oláh et
al. 2013c). There are historical materials scattered in museums, universities and
private collections. We have laboured an idea of cooperation how to realise com-
prehensive studies when funding is removed from taxonomy to more modern
regarded disciplines of genetics, ecology and conservartion and no resource re-
mained available even for adequate population sampling (Oláh et al. 2015b).
The examined freshly collected materials and the old historical type and
non-type specimens are deposited and curated in the following natural history
collections:
BDOL = Biologiezentrum des Oberösterreichischen Landesmuseums, Linz, Austria;

CCPC = Ciubuc Private Collection, Sinaia, Romania;
CMZL = Cantonal Museum of Zoology Laussane, Switzerland;
CNHM = Croatian Natural History Museum, Zagreb, Croatia;
DBFMNSUP = Department of Biology, Faculty of Mathematics and Natural Sciences,
University of Prishtina, Prishtina, Kosovo;
DEI = Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany;
HNHM = Hungarian Natural History Museum, Budapest;
MCSNBG = Museo Civico di Scienze Naturali “E. Caffi”, Bergamo, Italy;
MPC = Malicky Private Collection Lunz-am-See, Austria;
MFN = Museum für Naturkunde der Humboldt Universität zu Berlin, Germany;
MM = Mátra Museum of Hungarian Natural History Museum, Gyöngyös, Hungary;
MP ISEZ = NHM-ISEA;
NMPC = The National Museum (Natural History Department) in Prague, Czech
Republic;
NHMB = HNHM;
NHM-ISEA = Natural History Museum of the Institute of Systematics and Evolution of
Animals, Polish Academy of Sciences, Kraków, Poland;
NMNHBAS – National Museum of Natural History, Bulgarian Academy of Sciences,
Sofia;
NMP = The National Museum (Natural History Department) in Prague, Czech Republic;
NMPC = NMP;

114 J. Oláh
NMW = Naturhistorisches Museum Wien, Austria;

OPC = Oláh Private Collection, under national protection by the Hungarian Natural
History Museum, Debrecen, Hungary;
PMS = Slovenian Museum of Natural History, Ljubljana, Slovenia;
SMNHL = National Academy of Sciences of Ukraine, State Museum of Natural History,
Lviv, Ukraine;
WM = NMW
The original texts are cited verbatim, but the original character formatting
is not followed.
TAXONOMY
PHILOPOTAMIDAE
Wormaldia McLahlan, 1865
Wormaldia albanica Oláh, 2010
Wormaldia albanica Oláh, 2010: 68–69: “Holotype, male, HNHM. Albania:

Tepelenë county, Tepelenë, Uji i Ftohtë (Cold Water Spring), 165 m, N40°15.011’
E20°03.548’, 13.III.2008, leg. Sz. Czigány & D. Murányi.”
Diagnosis – Oláh (2010: 68–69): “This new species belongs to the spe-
cies complex described from the Balkan Mountains with enlarged endothecal
microspine cluster and tapering harpagones: W. kimminsi Botoşăneanu , 1960;
W. khourmai Schmid, 1959; W. balcanica Kumanski, 1979; W. bulgarica Novák,
1971. Most close to W. bulgarica described from Bulgaria but differs by having
(1) conspicuous basolateral flange of sclerites present on Xth segment and well
visible both in lateral and dorsal view; (2) in lateral view Xth segment has no
dorsal excision and no any dorsal subapical hook, dent or elevation, both present
and very conspicuous on W. bulgarica; (3) cerci slightly S-forming tapering in
dorsal view, not straight and clavate; (4) harpagones longer than coxopodites,
not shorter; (5) harpagones slender, tapering and downcurving apically; (6) en-
dotheca with a large spine and a group of four smaller spines besides the long
microspine cluster, not only with a single large spine.”
Etymology – “It is named for the country in which the type was collected.”
Wormaldia balcanica Kumanski, 1979

Wormaldia khourmai balcanica Kumanski, 1979a: 63–65: “Bulgaria, Stran-
dza Mts., hygropetric biotop near Katun-dere stream, not far from the bridge on
the road Malko Tarnovo-Zvezdetz (about 100 m a.s.l.), 11.VII.1976, 16²² and
12³³ (leg. Kumanski); Greece, Isl. Rhodos, 4²², Isl. Lesbos, 8²², and Isl. Chios,

12 ²² (all. leg. Malicky); Turkey, Prov. Samsun, 10 km NW Havza (500 m.a.s.l.),

41°30’N/35°33’, 15.V.1975, 1² (leg. H. & U. Aspoeck, H. & U. Rausch and P.
Ressl). Holotype ² (among the Bulgarian specimens), 22²² and 8³³. Paratypes
(19 from Bulgaria, 2 from Rhodos, 4 from Lesbos and 5 from Chios) in the au-
thor’s collection, 18 ²² and 4 ³³ Paratypes (4²² and 4³³ from Bulgaria, 1² from
Turkey and others from Greece) in coll. Malicky, Lunz am See, Austria.”
Wormaldia balcanica Kumanski, 1979: Kumanski (1985a: 167): raised to
species status.
Wormaldia bosniaca Botoşăneanu, 1960

Wormaldia occipitalis bosniaca Botoşăneanu, 1960a: 262, 263, 274: “Tre-
bevic, près de Sarajevo, 15. VII. 1955, entre 1100 et 1300 m. Petits ruisseaux as-
sez agités, coulant sous des sapins bas. Wormaldia (W.) occipitalis bosniaca n. ssp.
très commun.” “Trescavica, pas loin de Sarajevo, 18–20. VII. 1955. Rivière de
taille moyenne, en-dessous de Kosia Luka, verss 1200 m, dans la forêt; chassé
en-dessous de la source. Wormaldia (W.) occipitalis bosniaca n. ssp. 4².” Trnovo,
localité située au pied de Trescavica, vers 1000 m environ, 23. VII. 1955. Rivière
de 8–10 m de large, assez agitée. Wormaldia (W.) occipitalis bosniaca n. ssp. assez
commun.” “Je pense qu’il s’agit d’un nouvelle sous-espèce, que je vais dénommer
bosniaca n. ssp. 1² et 1³ de Trebevic ont été désignés comme holotype et allotype
(FS)[Fernand Schmid]; 1² parat.: DEI [Deutsches Entomologisches Institut,
Berlin]; 1² parat.: LB [Lazar Botoşăneanu ].”
Wormaldia bosniaca Botosaneanu, 1960: Malicky (2005b: 549): unclear
taxonomic position.
Wormaldia bosniaca Botosaneanu, 1960: Neu (2015: 107): raised to species
status.
Wormaldia bulgarica Novák, 1971

Wormaldia khourmai bulgarica Novák, 1971: 105–106: “Holotypus: Fun-
dorte: Bach bei Rilski Monastir, 14.VII.1962, 1², in der Sammlung des Autors.
Paratypen: Fundorte: Rila-Gebirge; Zufluss des Malovice Flusses, Rilski Monas-
tir, 14.VII.1962, 5², 3³; Bach bei Rilski Monastir, 14.VII.1962, 1²; Fluss Malovice,
Rilski Monastir, 11², 2³. (in der Sammlung von L. Botosaneanu).”
Wormaldia bulgarica Novák, 1971: Kumanski (1979a: 61–62): raised to
species status.
Wormaldia daga Oláh, 2015

Wormaldia khourmai Kumanski, 1979a: 62–63 (ssp.?): misidentification.

116 J. Oláh
Wormaldia daga Oláh, 2015 in Oláh & Kovács (2015: 99–100): “Holotype:
Bulgaria: Bosna Mts. Dudenovo, Dudenska Reka, between Vizitza and Novo
Panicharevo, 249m, N42°10’25” E27°34’07”, 26.VII.2012 at light leg. S. Beshkov
& M. Beshkova (1², NMNHBAS).”
Diagnosis – Oláh & Kovács (2015: 99–100): “We have collected W. khour-
mai Schmid 1959 on the Thales slopes near to the locus typicus: Irán, Thales Mts.
Masula River, 12.VIII.1990, leg. J. Oláh (3 males, OPC). The most important
genital structures involved in sexual selection, the endothecal spine pattern and
the the head of segment X are clearly different. This species with charactersitic
endothecal spine clusters and narrowing harpagones belongs to the Wormaldia
khourmai, W. bulgarica, W. balcanica, W. mahiri and W. erzincanica group of spe-
cies and most close to W. khourmai, but differs by having more swollen apex of
segment X, rounded, not truncate apex of cerci and only a single endothecal
spine, not three spines.”
Etymology – “Daga, from “dagadt”, swollen in Hungarian, refers to the apical
shape of the segment X in lateral view.”
Wormaldia hellenica Jacquemart, 1962

Wormaldia hellenica Jacquemart, 1962: 3–4: “Récoltés par la mission E.
Janssens-R Tollet, Mont Pélion W. Drakia (Khani Zisi) (1.200 m), 28/30-VII-
1953, 5 preparations microscopiques, Mont Olympe E Stavros a Prioni, 1.000 m,
21/23-VII-1953, 1 preparation microscopique.”
Wormaldia hellenica Jacquemart, 1962: Malicky (2005: 549): unclear taxo-
nomic position.
Wormaldia hellenica Jacquemart, 1962: Neu (2015: 107): raised to species
status.
Wormaldia homora Oláh, 2015

Wormaldia triangulifera asterusia Kumanski, 1975: 59: misidentification.
Wormaldia triangulifera asterusia Kumanski, 1975: Kumanski & Malicky
(1976: 103): misidentification.
Wormaldia triangulifera McLachlan, 1878: Kumanski (1985: 165–166):
misidentification.
Wormaldia homora Oláh, 2015 in Oláh & Kovács (2015: 102–103): “Holo-
type. Bulgaria, Eastern Rodopi, near Strazhetz, above the crossroad Gugutka-
Krumovgrad, 575m, N41°21’11” E25°50’35”, 24.VII.2012, at light, leg. S. Beshkov
& M. Beshkova (1², NMNHBAS).”
Diagnosis – Oláh & Kovács (2015: 102–103): “The species under the name
of Wormaldia triangulifera asterusia Malicky were collected from several regions in

Bulgaria: Stara Planina, Pirin Mts., Strandscha Mts. (Kumanski & Malicky, 1976).
Specimens from various regions exhibit rather stable genital structures: especially
the head of segment X and the endothecal spine pattern are conservative. We have
a single specimen from the Eastern Rodopi Mts. and compared its fine structure
with specimens of W. asterusia collected from Greece (Crete) W. homora sp. n. is
most close to W. asterusia, but differs by having apex of segment X high, not low,
apical portion of cerci truncate, not rounded in lateral view and the subapical mesal
projection triangular, not rounded lobe in dorsal view; apices of harpago narrowing
and downward curving, not broad; endothecal spine structure different.”
Etymology – “Homora, from “homorú”, concave in Hungarian, refers to the
apical shape of the segment X in lateral view. The apical portion beyond the sub-
apical dorsal point is concave in lateral view.”
Wormaldia juliani Kumanski, 1979

Wormaldia juliani Kumanski, 1979a: 58–60: “Bulgaria, Maleshevska Mt.,
streamlet falling into Struma River, nearly 2 km over railway station Kresna
(about 400 m a.s.l.), 9.VI.1975, Holotype ² and 4 Paratypes (3²² and 1³). Stran-
dzha Mts., spring Aidere near the bridge on the road Malko Tarnovo-Zvezdetz,
11.VII.1975, 1² Paratype. Holotype and 4 Paratypes deposed in the author’s col-
lection in the National Natural History Museum, Sofia; 1² Paratype in coll. H.
Malicky, Lunz am See, Austria. I am very obliged to Dr. Malicky for reexamining
and confirming the new species.”
Wormaldia occipitalis vaillantorum Botosaneanu, 1980a: 168–169: “Ho-
lotype ² de l’ île grecque de Kérkira (= Corfou): Or. (= Mt.) Pantokrator, 900 m.
alt., Mai (“16/30”) 1971, coll. B. van Aartsen. J’ai trouvé cet exemplaire dans les
collections entomologiques de notre Musée; il y est conservé à sec, abdomen dans
un petit tube à glycérine.”
Wormaldia juliani vaillantorum Botosaneanu, 1980: Botosaneanu (2004:
162): the subspecies Wormaldia occipitalis vaillantorum Botosaneanu, 1980 trans-
ferred to W. juliani Kumanski, 1979.
Wormaldia juliani Kumanski, 1979, Wormaldia vaillantorum Botosaneanu,
1980: Malicky (2005: 549): unclear taxonomic position.
Wormaldia juliani Kumanski, 1979: Neu (2015: 107): Wormaldia occipitalis
vaillantorum Botosaneanu, 1980, and Wormaldia juliani vaillantorum Botosanea-
nu, 1980 synonymised with Wormaldia juliani Kumanski, 1979.
Wormaldia kimminsi Botoşăneanu, 1960

Wormaldia triangulifera kimminsi Botoşăneanu, 1960a: 270–271: “4² et
5³ de Perister 12–16.VIII.1955, que j’ai designé comme holotype ², allotype ³ et

118 J. Oláh
paratypes ² et ³; holot. ² + allot. ³: F. Schmid; 1 ² parat.+ 1³ parat.: Deutsches

Entomologisches Institut, Berlin; 2²+2³ parat.: L. Botoşăneanu.”
Wormaldia kimminsi Botosaneanu, 1960: Malicky (1977: 68): raised to
species status.
Wormaldia subterranea Radovanović, 1932

Wormaldia subterranea Radovanović, 1932: 103–107: “Die Höhle Pom-
peska jama liegt 40 km südöstlich von Ljubljana (26 km Luftlinie), ungefähr 2 km
von der Station Dobre-polje an der Lokalbahn Ljubljana-Kocevje, und steht mit
dem gleich danebenliegenden Pokrito brezno (etwa: “zugedeckter Abgrund”)
in unterirdischer Verbindung. Dieser letztere ist nur eine unterirdische Schlucht
ohne direkte verbindung mit der Aussenwelt und wurde erst im Jahre 1914 durch
Sprengarbeiten zufälligerweise aufgeschlossen; der Eingang wurde aber sofort
wieder zugedeckt. Seitdem wurde diese Höhle nur zweimal noch besucht (am 12.
Juni 1927 und am 19. Juni dieses Jahres) und gleichzeitig zoologisch gründlich
erforscht. Am Grunde dieses unterirdischen Ganges, der sich bei seinem engen
Eingang gleich vertical 6 m in die Tiefe senkt und zirka 30 m lang ist, befindet
sich ein Bach, der jedenfalls aus weiter Ferne herkommt (man weiss nicht woher);
er verschwindet bal din einer engen Höhlung, erscheint dann wieder 45 m weiter
abwärts als Bach “k” (Kenk und Seliskar) in der Podpeska jama (84 m von dem
Eingang der Höhle entfernt und 9.5 m tiefer als dieser gelegen), um dann sofort
wieder unterirdisch zu verschwinden.
Die Larven und Nymphen der neuen Wormaldia-Form wurden in grosser Zahl
in dem eben erwahnten Bäche angetroffen, und die Imagines waren gleich daneben
an den feuchten Wanden haftend oder langsam kriechend zu finden. Es sind seit
Oktober 1928 bis 1. Juni dieses Jahres in der gleichzeitig als Höhlenlaboratorium
(Kenk-Seliskar) dienenden Podpeska jama insgesamt 85 Stück Imagines beo-
bachtwet worden. Sie sind zwar in allen Monaten des Jahres an dieser Stelle aufge-
funden worden, jedoch konnte eine auffällige Periodizität in ihrem Vorkommen
festgestellt werden (darüber spater). Die Tiere leben aber im unvergleichlich
grösser Menge in Pokrito brezno, und dieses muss als eigentliche Heimat der neu-
en Wormaldia-Form angesehen werden. Sie machen hier gleichzeitig bei weitem
die überwiegende Mehrzahl der gesamten Tierbevölkerung aus.”
Wormaldia occipitalis subterranea Radovanović, 1932: Kimmins (1953: 51):
treated as a subspecies.
Wormaldia occipitalis occipitalis Pictet, 1934: Botosaneanu (1989: 166): W.
subterranea and W. occipitalis subterranea are synonyms of W. occipitalis occipitalis.
Wormaldia subterranea Radovanović, 1932: Neu (2015: 107): species rank
resurrected.

PSYCHOMYIIDAE
Psychomyia Latreille, 1829
Psychomyia klapaleki Malicky, 1995
Psychomyia klapaleki Malicky, 1995: 443–444: “Holotypus ² und Para-
typen: Slowenien, Fluss Kolpa bei Gsparci, 23.VII.1994 (25², 1³); Paratypen,
Kocevie, 25.VI.1994, leg Wimmer (9², 18³); in meiner Sammlung. Weitere
Paratypen: Bosnien, Rogatica, 2.VIII.1897 (7²) und Gorazda, 9.VIII.1897 (1²)
leg. Klapálek (diese sind ein Teil der bei Klapálek (1898) erwahnten Ausbeute);
Slowenien, Sanntaler Alpen, Leutsch, 3–10.VII.1942 (14²) und 21-27.VII.1942
(3²), leg. Zerny; alle im Naturhistorischen Museum Wien.”
Tinodes Curtis, 1834

Tinodes andrasi Oláh, 2010
Tinodes andrasi Oláh, 2010: 74–76: “Holotype, male, HNHM Croatia:
Konavli Mts, Ljuta (near Gruda), Ljuta stream at the Konaviski dvori watermill,
60 m, N4° 32.076 E18° 22.610’, 7.X.2008, leg. L. Dányi, Z. Fehér, J. Kontschán &
D. Murányi.”
Diagnosis – Oláh (2010: 74–76): “This new species is close to the widely
distributed Tinodes rostocki MacLachlan, 1878, but differs by having (1) IXth
sternite short and tall, not long and low; (2) phallicata with S-forming thin api-
cal end, not C-forming; (3) paraproctal processes armed with 4–5 megasetae on
the middle, besides the apical set; (4) apical margin of coxopodite has altogether
only 3 spinelike processes, not 5 processes; apicomesal short and blunt processes
lacking; (5) dorsal pair of processes on basal plate low arching, not hook-form-
ing; (6) anterior apodeme of basal plate very enlarged and clavate.”
Etymology – “The name of this tiny Tinodes species was dedicated to András,
the newly-born son of the collector, Dávid Murányi.”
Tinodes polifurculatus Botoşăneanu, 1956

Tinodes polifurculatus Botoşăneanu, 1956: 382–386: “La prise No 30 con-
tenait un riche materiel (46 ²² et ³³): Bulgarie: Région de la ville Varna, Ceaika,
dans la region des ruisseaux, près du rivage de la mer. IX.1955, leg. A. Valkanov
& B. Rusev.”
Tinodes popovi Kumanski, 1975

Tinodes popovi Kumanski, 1975 in Kumanski & Malicky (1975: 25–
27): “Das gesamte Material stammt aus Bulgarien und umfasst acht Exemplare:
Holotypus ² und 4³ Paratypen: Ost-Stara Planina-Gebirge, Fluss Kotlenska ober-

120 J. Oláh
halb Kotel (ca. 800 m), 6.VII.1970. 1 ² Paratypus vom selben Fundort, 7.VI.1968,
leg. Kumanski. 1 ² Paratypus: Sliwen (dasselbe Gebiet), 1913 leg. Tchhorbadjieff.
1² Paratypus West-Stara Planina Planina-Gebirge, Zufluss des Flusses Zaselja,
Bov, 1.VII.1962, leg. Novak. Ein paratypus (vom Fluss Kotlenska) in coll Malicky,
der ²-Paratypus aus der Westlichen Stara Planina in coll. Novak, Praha, die
übrigen Typen in coll. Kumanski im Zoologischen Institute und Museum, Sofia.”
Tinodes raina Botoşăneanu, 1960

Tinodes raina Botoşăneanu, 1960b: 113–114: “Holotype ²: ruisseau
Trestenic, affluent droit de la Cerna, un peu en aval du ruisseau Presacina, Banat,
Roumanie, 4.X.1956. Conservé dans les collections du British Museum (Nat.
Hist.), Department of Entomology.”
Tinodes unidentatus Klapálek, 1894

Tinodes unidentatus Klapálek, 1894: 491–492: “Frequents the rapid streams
on the slopes of the Vitosa Mountain near Sophia, Bojana, 20th July, 1893”
Tinodes pseudorostocki Botoşăneanu, 1960a: 276–278: “J’ai eu à ma dis-
position 6² et 4³ de Perister, 12–16. VIII. 1955, dont j’ai designé le holotype ²,
l’allotype ³ ainsi que des paratypes ² et ³. Holot. ² + allot. ³: FS.; 1² parat.: DEI;
3² + 2³ parat.: LB.”
Tinodes pseudorostocki Botoşăneanu, 1960: Botoşăneanu & Sykora
(1963: 122): synonymised with T. unidentatus. “Espèce decrite par Klapálek
(1894, 1894) de ruisseaux rapides du massif Vitocha, et retrouvée à Kostenetz
(Mts. Ryla), le 16.VI.1959. Elle est connue aussi de Roumanie (Mts. Bihar).”
Tinodes urdhva Oláh, 2010

Tinodes urdhva Oláh, 2010: 77–78: “Holotype male, HNHM. Albania:
Dibër district, Korab Mts, Radomirë, torrent E of the village, 1460m, N41°49.131’,
E20°30.160’, 26.VI.2007, leg. D. Murányi.”
Diagnosis – Oláh (2010: 77–78): “This new species belongs to the Tinodes
kimminsi Sykora, 1962; T. maculicornis (Pictet, 1934); T. sarisa Malicky, 1975,
T. unicolor (Pictet, 1934) species complex having simple monolobed harpagones
and tall paraproct. Most close to T. sarissa Malicky described from Greece, but
differs by having (1) number and pattern of megasetae on paraproctal processes
different; (2) phallicata is simple rod without any structures; (3) harpagones up-
ward directed, not downward; (4) basal plate of gonopods with different struc-
tural units both in lateral and ventral view.”
Etymology – “Name was given with reference to the upward directed harp-
agones, upward “urdhva” in Sanscrit.”
POLYCENTROPODIDAE
Plectrocnemia Stephens, 1836
Plectrocnemia kisbelai Botosaneanu, 1967
Plectrocnemia kisbelai Botosaneanu, 1967a: 171–172: “Holotype ² (con-
servé en alcool, dans ma collection): 20.VI.1964, Mts. Apuşeni (ou de Bihor),
Scârişoara-Belioara, leg. Kis Béla. J’ai le plaisir de dédier cette espèce à son dé-
couvreur, entomologue distingué de Cluj, auquel je sui redevable pour bien des
captures interessantes de Trichoptères.”
Plectrocnemia kisbelai Botosaneanu, 1967: Botosaneanu (1995: 67):
“Only the holotype of this rare, endemic species was known. I was very pleased
to find more specimens in the recently studied collections. Some of them are, like
the holotype, from the Bihor mountains: 3 ²², 1³ taken on 13, 15, and 17. VII.
1971 along small tributaries of Valea Disghitului between Garda and the hamlet
Iarba Rea (catchment of Arieşul Mare), L. Botosaneanu leg. But 1 ² was caught
on 9 VI. 1966 in Valea Ciuluii, tributary of Valea Nadaşului (catchment of the
Mureş), not far from Cabana Debela Gora, Zarand mountains, L. Botosaneanu
leg. This last record means a slight extension of the known range.”
Plectrocnemia minima Klapálek, 1899

Plectrocnemia minima Klapálek, 1899a: 436–437: “Ein ² von Korniareva
im Krassó-Szörényer Comitat.”
Plectrocnemia minima Klapálek, 1899: Botosaneanu (1967a: 171): “Plect-
rocnemia minima Klap. est un endémite carpatique, et son aréal comprend unique-
ment les Mts. du Banat. Capturée d’abord à Cornereva (type ² de Klapálek); j’ai
capturé entre 1954 et 1961 à peu près 25 ²² (la ³ reste inconnue, ce qui est assez
curieux!) dans plusieurs sources, ruisselets de source et petits ruisseaux des envi-
rons des localités Băile Herculane (Herkulesbad), Mehadia et Cornereva (bassins
donc de la Cerna et de la Belareca). Or. Les Mts. du Banat étant bien explorées
au point de vue trichoptérologique, nous serions tentés de conclure à l’existence
d’un aréal extrémement restreint dans le cadre de ces montagnes; cependant, 1 ²
à été pris dans la “source” de la Beuşniţa (bassin de la Nera). Les zones des réseaux
lotiques que P. minima habite, sont: eukrenal, hypokrenal, epirhithral et, beau-
coup moins fréquemment, metarhithral. Les dates des captures sont comprises
entre le 1.V. et le 11.VII.”
Plectrocnemia mojkovacensis Malicky, 1982

Plectrocnemia mojkovacensis Malicky, 1982: 161: “Holotypus ² und Para-
typus ²: Crna Gora, Bistrica, Mojkovac, 19. VII. 1981, leg Sivec, coll. Malicky.
Paratypus ² mit gleichen Daten in coll Prirodoslovni Muzej Slovenije, Ljubljana.”
122 J. Oláh
Plectrocnemia smiljae Marinković-Gospodnetić, 1966

Plectrocnemia smiljae Marinković-Gospodnetić, 1966a: 112: “Holotype
², the source of the stream Zunovnica, Hadzici near Sarajevo, 28. V. 1964. Leg.
M. Dokić.”
Polycentropus Curtis, 1835

Polycentropus devetaki Krušnik et Malicky, 1992
Polycentropus devetaki Krušnik & Malicky, 1992: 56: “Holotype ²: Crna
Gora, Mojkovac, 10.VII.1985, leg D. Devetak, coll. Krušnik.”
Polycentropus ierapetra septentrionalis Kumanski, 1986

Polycentropus ierapetra septentrionalis Kumanski, 1986: 185–186: “Material
and localities: Struma valley, railway station Stara Kresna, 10.VI.1975, 1², and
24.VI.1981, 1² (leg. A. Slivov); Sestrino village, the foothills of Ograzhden Mt.,
19.IX.1984, 2²² (leg. J. Ganev); Rhodopes Mts., Lukovitza River, above Asenovgrad,
8.VIII.1983, 2²², and 15.VIII.1983, 2²² (leg. J. Ganev); Strandzha Mt., Ropotamo
River, 2 km above Krushevetz village, 4.VIII.1981, 2²²; Eastern part of Stara Planina
MT., streamlet, left tributary of the Kamtschia River, ca. 2 km from the “Kamtschia”
barrage, 15.VI.1984, 2²². All localities are below 350 m a.s.l. All samples were made
at artificial light. Holotype chosen among the specimens from the Rhodopes.”
Polycentropus ierapetra slovenica Malicky, 1998

Polycentropus ierapetra slovenica Malicky, 1998: 326–328: “Holotypus:
Slowenien, Mini, Socerga, 29.VI.1990, leg. C. Krušnik, in meiner Sammlung.”
Polycentropus schmidi Novák et Botoşăneanu, 1965

Polycentropus schmidi Novák et Botoşăneanu, 1965: 139–140: “1 holotype
² provenant des collections du musée de Budapest, et mis a notre disposition par
le Dr. F. Schmid. L’étiquette porte i’inscription suivante: “Vratna, Dr. Pasziczky”.
Vratna est une localité située dans la montagne Malá Fatra en Slovaquie. Insecte
conservé à sec, ailes déployées, abdomen en preparation microscopique, coll. F.
Schmid. 1 paratype ² (leg. K. Novák): ruisseau près de Lopusna, Slovaquie du
NO, 450 m. alt., 18.VI.1963. Conservé en alcool, coll. K. Novák.”
HYDROPSYCHIDAE
Hydropsyche Pictet, 1834
Hydropsyche botosaneanui Marinković-Gospodnetić, 1966a

Hydropsyche botosaneanui Marinković-Gospodnetić, 1966a: 112.

“Holotype ², Bosnia, river Miljacka (Mokro), 2.VI.1964.”
Hydropsyche dinarica Marinković-Gospodnetić, 1979

Hydropsyche dinarica Marinković-Gospodnetić, 1979: 165. “Holotype:
Sutjeska (river system of Drina), Tjentiste, 24.V.1965. Allotype ³: Sutjeska,
Tjentiste, 24.V.1965. Paratypes: Sutjeska, Tjentiste, 6²² 8³³, 24.V.1965; 1² 1³,
16.V.1967. Tributaries of the river Sutjesca at Tjantiste 2²², 19.VI.1968; 1²,
29.VI.1968. Josanica, a tributary of Drina at Coca, 1², 19.VI.1968. The source of the
river Pliva (river system of Vrbas), 1², 22.VI.1976. The source of the river Zeta, 4²²
1³, 274.V.1973. Bregava (Stolac), 1², 289.IV.19758. Holotype in autor’s collection.”
Hydropsyche emarginata Navas, 1923

Hydropsyche emarginata Navas, 1923: 90, 159–160: “Museum Paris Rég.
D’Iven et ravins de la côte 1422 (SE de Monastir) D. Vergné 1917”; “Mai”;
“Hydropsyche emarginata Nav./P. Navás S.J. det”; “Type” (Botosaneanu 1980b:
191).
Hydropsyche tjederi Botosaneanu et Marinković-Gospodnetić,
1966: 514–516: “1 ² (holotype). 31.VIII.1956, riviere Belareca a Mehadia. Banat.
Roumanie (leg. I. Capuse). L’holotype, en alcool, est conservé dans la coll. L.
Botosaneanu.” Synonymised by Botosaneanu (1980b: 191).
Hydropsyche mostarensis Klapálek, 1898

Hydropsyche mostarensis Klapálek, 1898b: 127–128: “Mostar 18./VIII.,
sklepána s dubu při cestě ku pramenu Jasenice.”
Hydropsyche peristerica Botosaneanu et Marinković-Gospodnetić, 1966

Hydropsyche peristerica Botosaneanu et Marinković-Gospodnetić, 1966:
516: “2 ²; torrent de largeur, au fond d’une vallée principale descendant sur Bitolia,
captures entre 1 800 et 2 500 m. (Mt. Persister, Macédonie Yougoslave), 12–16.VIII.
1955, leg F. Schmid. L1holotype ² (à sec, ailes déployées, abdomen dans un petit
tube d’alcool glyceriné) et l’allotype ³ (à sec, ailes déployées) dans la coll. F. Schmid;
paratype ² dans la coll. L. Botosaneanu (à sec, abdomen en alcool glyceriné)”
Hydropsyche sarnas Oláh, 2015

Hydropsyche sarnas Oláh, 2015 in Oláh & Kovács (2015: 106–107):
“Holotype. Albania, Gjirokastër District, N of Humelicë, shore vegetation of

124 J. Oláh
river Drino, 170m, N40.17854° E20.07981°, 10.05.2014, leg. Z. Barina, D. Pifkó

& G. Puskás (1², OPC).”
Diagnosis – Oláh & Kovács (2015: 106–107): “Belongs to the Hydropsyche
angustipennis species group and to the Hydropsyche pellucidula species clus-
ter of Oláh & Johanson (2008). Close to Hydropsyche dinarica Marinković-
Gospodnetić but differs by the lateral profile of the median keel of segment X, by
the clearly twopartite apical profile of segment X and by the extremely enlarged
subapical lateral projection on the head of the phallic organ.”
Etymology – “Sarnas, from “szárnyas”, winged in Hungarian, refers to the
very much produced, wing-like angular, subapical, lateral projections before the
cleft apex of the phallotheca.”
Hydropsyche sinuata Botosaneanu et Marinković-Gospodnetić, 1966

Hydropsyche sinuata Botosaneanu et Marinković-Gospodnetić,
1966: 513–514: “1 ² et 3 ³, 23.VI.1960, ruisselet Silistea, petit affluent gauche
du Lapusnic, a 2 km environ en amont du village Lapusnica Mare (Mts de Banat,
Roumanie) La materiel est conservé en alcool. Holotype ², allotype ³ et 2 para-
types ³ dans la coll. L. Botosaneanu.”
Hydropsyche smiljae Marinković-Gospodnetić, 1979

Hydropsyche smiljae Marinković-Gospodnetić, 1979: 167–169:
“Holotype ² and allotype ³: Buna, the left tributary of the river Neretva, 27.V.1975.
Paratypes: 8²² 1³, 28.VI.1967; 4²² 4³³, 27.V.1975; 2²² 1³, 8.VII.1975. Trebizat,
the right tributary of the river Neretva: 1², 28.IV.1974. (Vitina) 2²² 2³³,
28.IV.1974. Trebisnjica, 4²² 1³, 30.VI.1967. Holotype in the author’s collection.”
Hydropsyche fischeri Botosaneanu, 1980a: 169: Synonymised by Botosa-
neanu (2004: 166).
Hydropsyche tabacarui Botoşăneanu, 1960

Hydropsyche tabacarui Botoşăneanu, 1960b: 114–115: “29.X.1959, Vallée
de Iezerul, affluent de l’Ampoi, peu en aval du lac Ighiel, pas loin de Alba Iulia,
Transsylvanie, Roumanie. Leg. I. Tabacaru. Holotype ², allotype ³, 1 paratype ²
et 6 paratypes ³, dans les collection de l’auteur; 1 paratype ², 1 allotype ³ dans les
collections du British Museum (Nat. Hist.), Department of Entomology.”
RHYACOPHILIDAE
Rhyacophila Pictet, 1834
Rhyacophila akutila Oláh, 2010

Rhyacophila akutila Oláh, 2010: 82–83: “Holotype male, HNHM. Bulgaria:

Sofia province, Rila Mts, Borovec, Prava Marica Stream at Zavračica mountain
hut, 2190 m, N42°10.075’, E23°38.504’, 8.IX.2005, leg. M. Földvári & D. Murányi.
Paratype, female, HNHM. Bulgaria: Sofia province, Rila Mts, Borovec, Prava
Marica Stream at Zavračica mountain hut, 2190 m, N42°10.075’, E23°38.504’,
8.IX.2005, leg. M. Földvári & D. Murányi.”
Diagnosis – Oláh (2010: 82–83): “This new species belongs to the R. tris-
tis species group. Most close to R. pseudotristis Kumanski, 1987, but differs by
having (1) abdominal tergites and sternites with very pronounced light coloured
alveolar haloes, R. pseudotristis has no light ring around alveoli at all; (2) in dor-
sal view the apical excision on Xth segment almost pentangular, not triangular;
(3) in dorsoapical view the pair of apicoventral flaps on the dorsal branch of the
Xth segment laterad directed, not parallel or mesad oriented; (4) the vertically
oriented ventral branch of the Xth segment short and straight, not long and arch-
ing; (5) apices of the vertical branche of the Xth segment deeply bifid, not shal-
low; (6) mesal lobe of the apices pointed, not blunt; (7) membranous, almost in-
discernible blunt parameres much shorter than the paired less sclerotized dorsal
processes of the phallobase.”
Etymology – “This species is named with reference to the peculiar straight
ventral branch of the Xth segment; “akutila” is “straight” in Sanscrit.”
Rhyacophila balcanica Radovanović, 1953

Rhyacophila balcanica Radovanović, 1953: 20–22, 32, 38–39: “Fundort:
ein Männchen auf dem Bjelassitza-Gebirge in Montenegro (am 12. VIII. 1948).”
Rhyacophila biegelmeieri Malicky, 1984

Rhyacophila biegelmeieri Malicky, 1984: 297: “Holotypus ²: Griechenland,
Grammos-Gebirge bei Eptachorion, 1000 m, 10. VII. 1976, leg. H. Biegelmeier,
in meiner Sammlung.”
Rhyacophila biegelmeieri Malicky, 1984: Malicky (2005a: 48): recorded
from six localities in Greece.
Rhyacophila biegelmeieri Malicky, 1984: Oláh (2010: 84): recorded from
Albania (Tropojë district).
Rhyacophila biegelmeieri Malicky, 1984: Oláh & Kovács (2013: 110): re-
corded from Albania (Librazhd district).
Diagnosis – Oláh & Beshkov (2016: 95): “Apicodorsal process long hori-
zontal, broad based with triangular apex. Cerci enlarged and elongated quadratic
with paralell lateral margins. Dorsoapical process of segment X shorter than cer-
ci. Epiproct (anal sclerite) short. Apicodorsal excision on harpagones low. Easily

126 J. Oláh
distinguished from its sibling species (R. diakoftensis Malicky, 1983, R. neretva
sp. n., R. nyurga sp. n., R. pascoei McLachlan, 1879) by the enlarged parameres
and by the paramere dorsum densely packed with strong setae. R. neretva, R.
nyurga and R.pascoei have only a small cluster of setae on parameres subapically
ventromesad. R. diakoftensis has no any additional setae on parameres; only the
single terminal seta is present. The tripartite aedeagus has the paired dorsal arm
very short, long at all the other sibling species; Ventral arm of the aedeagus long
and spatulate in ventral view.”
Rhyacophila bosnica Schmid, 1970

Rhyacophila bosnica Schmid, 1970: 161. “Holotype ²: Yougoslavie, Bosnie,
Vucjaluka.”
Rhyacophila braaschi Malicky et Kumanski, 1976

Rhyacophila braaschi Malicky et Kumanski, 1976 in Kumanski & Malicky
(1976: 98–99): “Holotypus ² und 8 paratypen ²: Bulgarien, Stara Planina, Ne-
benbach der Strjama oberhalb Sopot, 1700 m, 11.IX.1971 (=BStpl 2). Weitere
Paratypen ²²: BStpl 8 (2²), BPir 4 (1²), BRo 3 (4²). Holotypus und Paratypen
von allen Platzen in coll. Malicky, Paratypen (ausser vom Pirin) ebenso in coll.
Kumanski.”
Rhyacophila brevifurcata Kumanski, 1986

Rhyacophila brevifurcata Kumanski, 1986b: 55–57: “Bulgaria, Western
Stara Planina, the river Barzija near the Petrohan-pass (ca. 1400 m a.s.l.), 10.VI.
1979, 1 ², and the stream Sini Vir at the tourist home Probojnica (ca. 1000 m
a.s.l.), 18–22.VIII.1980, 3². Holotype chosen among the males from the second
locality. One paratype in coll. L. Botosaneanu (Amsterdam); the other types in
the author’s collection at the National Natural History Museum (Sofia).”
Rhyacophila cibinensis Botosaneanu et Marinković, 1967

Rhyacophila cibinensis Botosaneanu et Marinković, 1967: 1145–1149:
“Le 19 mai 1963, 5² et 3³ (leg. B. Kis) furent capturés à Paltini, près de deux af-
fluent du Rîul Mare; cette localité est située dans les Monts de Cibin, Carpates
méridionales, 1400 m alt. environ, pas loin de Sibiu (Hermannstadt). 2² et 1³
furent capturés, le même jour et par le même collégue, près d’un ruisselet affluent
du Sadu, dans les mêmes Monts de Cibin, près du sommet Batrîna (1500–1600 m
alt.). Holotype ² et allotype ³ dans la collection L. Botosaneanu; les paratypes ²
et ³ sont gardés dans les collections L. Botosaneanu, M. Marinković et F. Schmid.

Le 23 mai 1966, un de nous (L.B.) a réussi à retrouver R. cibinensis près du ruis-

seau Izvorul Danesii (Monts de Cibin, bassin de Cibin, à quelques centaines de
mètres de Paltinis). 13² et 8³ furent capturés dans la végétation des bords du
ruisseau.”
Rhyacophila confinium Botoşăneanu, 1957

Rhyacophila confinium Botoşăneanu, 1957a: 62–63: “Bach Comanul,
Nebenfluss des Baches Wasser, 3 km von der Quelle abwärts (Marmarosh) 19.
VII. 1956. 1² in Alkohol (Holotypus SMF N3).”
Rhyacophila confinium Botoşăneanu, 1957: Szczęsny & Chvojka (2008:
160): “Ukraine, Czarnohora Massif, Arendarski stream, 16.VII.1995, leg. B.
Szczęsny, 4²², 1³.”
Rhyacophila denticulifera Kumanski, 1986

Rhyacophila denticulifera Kumanski, 1986b: 51–55: “Material and locali-
ties. Rila Mountain: Resorts “G. Dimitrov” and Dolna Banja, 13–14.IX.1967, 2²
and 1³; brooklets with hygropetric niches in the vicinity of Kartalska poljana
(1400–1600 m a.s.l.), 15–17.VIII.1972, 15² and 2³; The Rhodopes: the river
Sirikolaska, upper flow (1200–1500 m a.s.l.) 31.VIII.1968, 3², and 6.X.1976,
1²; brooklet in Caira-locality, 20.VII.1972, 1²; brooklet, right tributary of the
Cerna River above Smoljan, 17.VII.1971, 4². Strandza Mountain: the Ropotamo
River near Krusevec village (ca. 300 m a.s.l.), 25.IV.1970, 1² (leg. D. Braasch).
Holotype chosen among the males of the long series from Rila. One paratype ²
in coll. H. Malicky (Lunz) and another ² (indicated earlier as Rh. furcifera) in
coll. A. Dyakonoff (Leiden) (both specimens from the Cerna river). Holotype
and other paratypes in the author’s collection at the National Natural History
Museum (Sofia).”
Rhyacophila dohleri Botoşăneanu, 1957

Rhyacophila dohleri Botoşăneanu, 1957b: 180–182: “19.VII.1956, Torrent
du Comanul, affluent de la rivière Wasser, à environ 3 km en aval de la source
(Maramouresch, Nord des Carpathes orientales, environ 1300 m alt.). 1 ² en al-
cool (Holotype, dans les collections du Rijksmuseum van Natuurlijke Historie
de Leyde).”
Rhyacophila fagarashiensis Botosaneanu, 1964

Rhyacophila fagarashiensis Botosaneanu, 1964: 177–179: “Holotype ²:
7.VIII.1962, cours superieur (zone alpine) du torrent alpin Podragul, massif de

128 J. Oláh
Fagarash – Alpes de Transylvanie, Carpates Meridionales, ca. 2.100 m. alt. (dans

la collection de l’auteur). 2 paratypes ²: 8.VIII.1962, le torrent alpin Podragul dans
la zone superieure de la forêt d’épicea, massif de Fagarash-Alpes de Transylvanie,
Carpates Meridionales, 1700-1800 m. alt. (un exemplaire dans la collection F.
Schmid, l’autre dans la collection de l’auteur).”
Rhyacophila fischeri Botoşăneanu, 1957

Rhyacophila fischeri Botoşăneanu, 1957b: 186–188: “11.VI.1956. Le ruis-
seau de forêt Scocina, affluent droit de la rivière de Cerna à environ 1 km en
amont du point denommé “7 izvoare reci” (Banat, pas loin de Bâile Herculane-
Herkulesbad). Holotype ² (avec aedeagus en extension totale), dans les collec-
tions du Rijksmuseum van Natuurlijke Historie de Leyde). Paratype ² dans la
collection de l’auteur. Material en alcool.”
Rhyacophila flava Klapálek, 1898

Rhyacophila flava Klapálek, 1898a: 489–490: “Hungary: Aus Marmaros.
1²”; Klapálek (1899a: 437–438).
Rhyacophila furcifera Klapálek, 1904

Rhyacophila meyeri var. furcifera Klapálek, 1904: 729: “Goetzenberg
3/8. 2².”
Rhyacophila furcifera Klapálek, 1904: Botoşăneanu (1952a: 547–550):
raised to species rank. “In pâdure de conifere pe valea Galeşului, masivul Retezat,
9.VI.1951.”
Rhyacophila joosti Mey, 1979

Rhyacophila joosti Mey, 1979: 124–125: “Holotyp ² und Paratyp ²: 2.X.1976,
rechter Nebenbach der Strjama oberhalb Sopot, Balkangebirge, Bulgarien. Die
Typen werden im Museum der Natur, Gotha aufbewahrt.”
Rhyacophila kimminsiana Botosaneanu, 1958

Rhyacophila kimminsi Botoşăneanu, 1957: 179–180: “20.VIII.1956,
Torrent de Cocora, affluent de la rivière Ialomitza, près de la cabane de Pestera,
massif de Bucegi (Carpathes méridionales, environ 1600 m alt). 1 ² en alcool
(Holotype, dans les collections du Rijksmuseum van Natuurlijke Historie de
Leyde)”. Homonym: preoccupied by Ross (1956: 122).
Rhyacophila kimminsiana Botosaneanu, 1958: 139: replacement name.

Rhyacophila kownackiana Szczęsny, 1970

Rhyacophila kownackiana Szczęsny, 1970: 773–772: “Holotype ²: le tor-
rent Stakatsu? Dans Botev, 18.VIII.1969. Le spécimen dans la collection du
Musée de Zoologie à Sofia.”
Rhyacophila liutika Oláh, 2010

Rhyacophila liutika Oláh, 2010: 85–87: “Holotype male, HNHM. Macedo-
nia: Southeastern region, Belasica Mts. Kolesino, waterfall of the Kolesino stream
in platan-beech forest above the village, 500 m, N41° 23’, E22° 48’, 18.X.2006, leg.
L. Dányi & D. Murányi.”
Diagnosis – Oláh (2010: 85–87): “This new species belongs to the species
group of R. stigmatica and close to species complex described from the Car-
pathians: R. furcifera and from various parts of the Balkan: R. kownackiana
Szczęsny, R. morettina Botosaneanu, R. brevifurcata Kumanski, R. denticulifera
Kumanski. Most close to R. denticulifera described from Bulgaria but differs by
having (1) dorsomedial process of IXth tergite lost; (2) apical part of segment
X rounded balloon-shaped in lateral view, not rectangular; (3) two vertical setal
lines both located inside a single circular area in the caudal concavity of the Xth
segment, not in two separate vertically oval area; (4) the dorsal processes of the
phallobase differently shaped and their apical lobes smooth, not serrated.”
Etymology – “This species is named to remember Liütika (today Levente).
He was the first son of the Hungarian Prince Árpád and led the Turkic Kabar
tribes to balance the Turkic Bolgars on the Balkan during the great Hungarian
return to the Carpathians.”
Rhyacophila margaritae Kumanski, 1998

Rhyacophila margaritae Kumanski, 1998 60–61: “All the localities are
streams in the forest zone of the Central Stara Planina Mts. Stara Ribaritsa
riv., 1000–1200 m .s.l., 19 males (25.09.1997); same river, 950-1000 m, 1 male
(23.10.1996); Bolovandzhishka riv., 1050 m, 7 males and 1 female (26.09.1997);
Krayovitsa riv., above “Yavorova luka”-hut, 740–1000 m, 3 males and 1 female
(22–24.09.1997); Beli Osum riv., above “Haidushka pesen”-hut, 1000–1100 m,
15 males; left tributary to the Kositsa riv. above “Antshova batshyia”-hut, 100–
1200 m, 41 males (26.10.1997); Svinska (left tributary to Tsherni Vit river) ca.
600 m a.s.l. 1 male (03.11.1996), and 3 males (27.10.1997). Holotype male chosen
among the insects of the second locality; all other specimens designated as para-
types. The whole type series collected by the author and deposited in alcohol in
the National Museum of Natural History, Bulgarian Academy of Sciences, Sofia.”

130 J. Oláh
Rhyacophila mocsaryi Klapálek, 1898

Rhyacophila mocsaryi Klapálek, 1898: 489: “Görgény im Com. Maros-
Torda. 1², 14³.” Klapálek (1899a: 437).
Rhyacophila morettina Botosaneanu, 1980

Rhyacophila morettina Botosaneanu, 1980a: 165–166: “Holotype ² de
Yougoslavie, Bosnie: Jajce, 6.VI.1963, coll. F.C.J. Fischer. Conservé à sec, abdo-
men dans un petit tube à glycérine. Cet exemplaire était dans une collection de
trichoptères de Yougoslavie que m’avait léguée F.C.J. Fischer.”
Rhyacophila motasi Botoşăneanu, 1957

Rhyacophila motasi Botoşăneanu, 1957a: 61–62: “Kleiner Nebenfluss des
Dragan-Tales bei Zerna (Bihar-Gebirge); 26.VI.1953. 1² in Alkohol (Holotypus
SMF N2).”
Rhyacophila neretva Oláh, 2016

Rhyacophila neretva Oláh, 2016 in Oláh & Beshkov (2016: 97): “Holotype:
Bosnia & Herzegovina, Neretva River before Mostar, 13.IX.1989, light, leg. J.
Oláh (male, OPC). Paratypes: same as holotype (44 males OPC).”
Diagnosis – Oláh & Beshkov (2016: 97): “This new species forms a spe-
cies complex with the closely related sibling species of R. biegelmeieri Malicky,
1984, R. diakoftensis Malicky, 1983, R. nurga sp. n. and R. pascoei McLachlan,
1879. Distinguished from all the other siblings by the strongly downward curv-
ing clavate dorsal process of segment IX, by the length ratio of cerci and dorsal
process of segment X, by the configuration of the tripartite aedeagus.”
Etymology – “Named after the Neretva River where the specimens were col-
lected.”
Rhyacophila nurga Oláh, 2016

Rhyacophila nurga Oláh, 2016 in Oláh & Beshkov (2016: 98): “Holotype:
Montenegro, Pivska Planina, Crkvicko Polje, near Rudine Village, 1117 m,
N43°19’48”; E018°53’41”, 7.VIII.2015, leg. S. Beshkov & A. Nahirnic (male,
OPC).”
Diagnosis – Oláh & Beshkov (2016: 98): “A member of the Rhyacophila
pascoei new species complex. Differs from R. diakoftensis Malicky, 1983 by hav-
ing longer dorsal apical lobe on segment X; apical lobe narrow tapering, not
broad middle; cerci very long, not the shortest in the species complex, like at R.

diakoftensis; epiproct long, not short; configuration of the tripartite aedeagus

different.”
Etymology – “Nurga, from “nyurga”, slender or elongate in Hungarian with
reference to the elongate apicodorsal process of segment IX, cerci, paraproct and
epiproct.”
Rhyacophila obtusa Klapálek, 1894

Rhyacophila obtusa Klapálek, 1894: 492–493: “Three ² at a brook in
Dragalevci flowing from the Vitosa Mountains near Sophia, 22nd July (1893).”
Rhyacophila olahorum Oláh, 2016

Rhyacophila orghidani Botoşăneanu, 1952: Schmid (1970: 66): the new
drawing was prepared not from the holotype of R. orghidani, but from the here
distinguished and described closely related sibling, R. olahorum sp. n. as detect-
able from the lateral view of the harpago. Misidentification.
Rhyacophila orghidani Botosaneanu, 1952: Malicky (1983: 6, 2004: 7): re-
producing the erroneous drawing of Schmid. Misidentification.
Rhyacophila olahorum Oláh, 2016 in Oláh & Beshkov (2016: 91):
“Holotype: Romania, Munţii Apuseni, Munţii Gilăului, Staţiunea Muntele
Băişorii, spring stream of Vadului, N46°31.954’, E23°16.852’, 1552 m, 26.V.2013,
singled, leg. J. Oláh, E. Bajka, Cs. Balogh & G. Borics (male, OPC). Allotype: same
as holotype (female, OPC). Paratypes: Romania, Munţii Apuseni, Someşul Cald
Gorge, 1143 m, 5–15.VI.1999, leg. L. Újvárosi (8 males, 8 females, OPC); Munţii
Apuseni, Cetatea Radusei, 14.VI.1999, leg. L. Újvárosi (1 male, OPC); Munţii
Apuseni, Someşul Cald Gorge, 1143 m, 10.VI.2007, leg. M. Bálint (1 male, OPC).
Munţii Apuseni, Munţii Gilăului, Staţiunea Muntele Băișorii, three-branched
stream, N46°30.701’, E23°16.279’, 1620 m, 19.VI.2013, singled, leg. J. Oláh, Cs.
Balogh & S. Fekete (1 male, OPC); Munţii Apuseni, Munţii Gilăului, Muntele
Mare, spring stream area of Valea Mare, 1826 m, 19.VI.2013, singled, leg. J. Oláh,
Cs. Balogh & S. Fekete (1 male, OPC). Munţii Apuseni, Masivul Vlădeasa, Stâna
de Vale, Pastravariei stream, N46°41.676’, E22°38.027’, 1277 m, 6.VI.2015, leg.
M. Kiss, J. Oláh & L. Szél (12 males, 4 females, OPC); Munţii Apuseni, Masivul
Vlădeasa, Stâna de Vale, small stream, crossing road to Culmea Baia Popii,
N46°40.452’, E22°38.045’, 1335 m, 6.VI.2015, leg. M. Kiss, J. Oláh & L. Szél (3
males, OPC); Munţii Apuseni, Masivul Vlădeasa, Stâna de Vale, upper section of
Jád stream, N46°41.5’, E22°36.725’, 1135 m, 5.VI.2015, leg. M. Kiss, J. Oláh & L.
Szél (3 males, OPC); Munţii Apuseni, Masivul Vlădeasa, Stâna de Vale, upper sec-
tion of Jád stream, N46°41.867’, E22°36.666’, 1075 m, 5.VI.2015, leg. M. Kiss, J.
Oláh & L. Szél (5 males, OPC); Munţii Apuseni, Masivul Vlădeasa, Stâna de Vale,

132 J. Oláh
Galbenele stream, N46°40.809’, E22°37.147’, 1180 m, 7.VI.2015, leg. M. Kiss, J.

Oláh & L. Szél (12 males, 8 females, OPC); Munţii Apuseni, Masivul Vlădeasa,
Stâna de Vale, upper section of Ciripa stream, N46°40.546’, E22°38.515’, 1360 m,
6.VI.2015, leg. M. Kiss, J. Oláh & L. Szél (36 males, 13 females, OPC); Munţii
Apuseni, Cheile Someşului Cald, spring area, N46°37’59.77”, E 22°42’39.64”,
1247 m, 20.V.2015, leg. Cs. Balogh (26 males, 1 female, OPC); Munţii Apuseni,
V. Cuciulata, spring stream near Piatra Grăitoare, N46°38’38.41”, E 22°41’40.62”,
1521 m, 20.V.2015, leg. Cs. Balogh (9 males, OPC); Munţii Apuseni, Munţii
Gilăului, Staţiunea Muntele Băişorii, Muntele Mare (Öreghavas), Iara (Jára)
spring area, N46°28.914’, E23°13.294’, 1750 m, 19.VI.2015, singled, leg. J. Oláh,
Cs. Balogh & P. Juhász (1 male, 2 females, OPC); Munţii Apuseni, Munţii
Gilăului, Staţiunea Muntele Băişorii, Muntele Mare (Öreghavas), Creţoaia spring
area (Posaga tributary), N46°28.862’, E23°13.921’, 1750 m, 19.VI.2015, singled,
leg. J. Oláh, Cs. Balogh & P. Juhász (1 male, OPC).”
Diagnosis – Oláh & Beshkov (2016: 91): “In dorsal profile of segment
X the apical excision is deep and wide, not shallow and narrow. Ventral lobe
of harpago parallel-sided with obliquely truncate apex, not gradually tapering.
Apex of the dorsal process of the phallotheca quadrifid, not trifid. Apical margin
of the posterior process of the vaginal sclerite complex truncate, not bifid.”
Etymology – “This beautiful species is named for all Vlach peoples of Illyr-
ian-Thracian-Dacian origin scattered today as Aromans in Albania, Bulgaria,
Bosnia-Herzegovina, Croatia, Greece, Republic of Macedonia, Serbia and as well
as Romanians in Romania. In Hungarian language Vlach is pronounced as Oláh
to lessen consonant dominance. Consequently named also for all Hungarian
nominate Oláh families populated the Carpathian Basin, especially the Munţii
Apuseni, the habitats of Rhyacophila olahorum sp. n. and its surroundings from
the Middle Ages, including the ancestors of the first author.”
Rhyacophila orghidani Botoşăneanu, 1952

Rhyacophila orghidani Botoşăneanu, 1952b: 721–723: “ Rhyacophila
orghidani n. sp. a fost descoperită de noi în cursul verii anului 1951; 6 ² şi 2 ³ au
fost capturate în imediata apropiere a unei cascade eu apă puternic incrustantă
(afluent al Arieşului mic) în limitele satului Vidra de Jos, Munţii Apuseni.”
Notes – Oláh & Beshkov (2016: 94): “Dorsal profile of apical margin of
segment X with small mesal excision delimited by small lateral triangular lobes.
Ventral lobe of harpago gradually tapering. Dorsoventral profile of heavily scle-
rotized head of dorsal process of phallotheca trifid; mesal arm long and pointed.
Apical margin of posterior process of vaginal sclerite complex bifid, not trun-
cate.”

Rhyacophila pirinica Kumanski, 1980

Rhyacophila pirinica Kumanski, 1980: 197–198, 203: “Holotype ²: Pirin
Mts., hostel “Pirin” (1600 m a.s.l.), 27.VI.1979 (leg. A. Popov), in the National
Natural History Museum, Sofia.”
Rhyacophila pseudotristis Kumanski, 1987

Rhyacophila pseudotristis Kumanski, 1987a: 17–21: “Pirin Mts.: Circus Ban-
derski, the outlet of the Greater Banderski-lake (ca. 2250 m a.s.l.), 22.IX.1967, Holo-
type ² and 3² and 2³ paratypes (2 pairs in copula), leg. K. Kumanski; brooklet be-
low the Muratovi-lakes (ca. 2300 m a.s.l.); 22.IX.1967, 6² and 3³ paratypes (of them
2² and 2³ in collection of Dr. O. Nybom, Imatra, Finnland, labelled as Rh. tristis
by me); river Banderiska above the tourist home “Vihren” (2063 m a.s.l.), 21.IX.
1967, 7² and 5³ paratypes, leg. K. Kumanski, at light; brooklet below the peak
Vihren, 27.VIII.1979, 6² and 2³ paratypes (of them 2² in collection of Dr. S. Smith,
Washington, USA, labelled as Rh. tristis by me), leg. P. Beron; Circus Demianiski, riv-
er Demianiska (2000–2200 m a.s.l.), 8.X.1976, 6² and 2³ paratypes, leg. K. Kumans-
ki; small lake below the Popovo-lake (2200 m a.s.l.), 28.VII.1978, 2² paratypes, leg.
K. Kumanski. Rila Mts.: “The Seven Lakes”, Circus the outlet of the 4th lake (ca.
2250 m. a.s.l.), 9.VIII.1969, 1² paratype, leg. A. Ianev; Elenski-lake (2470 m. a.s.l.),
21.VIII.1972, 5² paratypes, leg. P. Beron; small brooklet with hygropetric niches,
right tributary of Cena-river above Smoljan (ca. 1100 m a.s.l.), 17.VII.1971, 1² para-
type, leg.K. Kumanski. Stara Planina Mts.: right tributary of Lesnica-river, ca. 10 km
aboveIasenova-village (ca.950 m a.s.l.), 20.VI.1984, 2² paratypes, leg. K. Kumanski
(together with a large series of the proper Rh. tristis). If not otherwise stated, all the
types are preserved in the collection of the National Natural History Museum, Sofia.”
Rhyacophila schmidinarica Urbanič, Krušnik et Malicky, 2000

Rhyacophila schmidinarica Urbanič, Krušnik et Malicky, 2000: 17–18:
“Holotype ²: Slovenia, Podplana, 19.V.1983, leg. I. Kos, coll. Krušnik. Paratypes:
Podplana, 22.V.1983, 1³; Zlebic, 11.V.1983, 1²; 29.V.1983, 1³; 5.VI.1983, 1²; all
leg. I. Kos, coll. Krušnik. Paratype: Crna Gora, Durmitor, Tara, 13.VII.1981, 1²,
leg. I. Sivec, coll. Malicky.”
Rhyacophila trescavicensis Botoşăneanu, 1960

Rhyacophila trescavicensis Botoşăneanu, 1960a: 265–266: “8² et 3³ de
Trescavica 18–20.VII.1955, que j’ai designé comme holotype ², allotype ³ et para-
types ² et ³; holot. ² + allot. ³: F. Schmid; 1 ² parat.: Deutsches Entomologisches
Institut, Berlin; 5²+2³ parat.: L. Botoşăneanu.”

134 J. Oláh
Rhyacophila vranitzensis Botosaneanu et Marinković-Gospodnetić, 1967

Rhyacophila vranitzensis Botosaneanu et Marinković-Gospodnetić,
1967: 1149–1151: “Nous avons eu a notre disposition 6 ² et 4 ³ en provenance
des localités suivantes de Bosnie: Mont Vranitza, ruisselet, 12 août 1956; Mont
Bjelachnitza, Chavnitzi, ruisselet, 23 juin 1957; rivière Miljacka a Mokro, 24
juillet 1964; le holotype ² et allotype ³ ont eté choisis parmi les exemplaires de
cette dernière localité, ils sons conservés dans la collection M. Marinković; un
paratype ² de Miljacka dans la coll F. Schmid et un autre dans la coll. L. Botosa-
neanu.”
GLOSSOSOMATIDAE
Glossosoma Curtis, 1834
Glossosoma bunae Marinković-Gospodnetić, 1988
Glossosoma bunae Marinković-Gospodnetić, 1988: 42–45: “Za sada
jedino poznato nalazište Glossosoma bunae je izvorište reke Bune u blizini
Mostara. Svi primerci Glossosoma discophorum, koji su nalaženi u Buni, u radu
Marinković-Gospodnetić (1978) pripadaju vrsti Glossosoma bunae n. sp. Ova vr-
sta je i kasnije tu nalažena, kako u proleónim mesecima (naročito u maju i junu),
tako i u jesen (u septembru, oktobru i novembru). Holotip mužjaka, alotip ženke
i paratipovi mužjaka i ženke nalaze se u autorovoj kolekciji.”
Glossosoma discophorum Klapálek, 1902

Glossosoma discophorum Klapálek, 1902: 165–166: “Stolac (Winneguth)
2² und 2³.”
Glossosoma discophorum Klapálek, 1902: Botosaneanu (1995: 62): “The
only Romanian records for this species being from the Apuşeni Mountains and
from the Eastern Carpathians, it is not uninteresting to mention its presence also
in the Southern Carpathians (20.VIII.1973, Retezat mountains: Gura Zlata, 1²,
leg. Eleonora Erhan-Dinca) as well as – with a slight doubt – in Oltenia (30–31.
VII.1971, Runc-Gorj, various springs, 2³³, leg. L. Botosaneanu).”
Glossosoma neretvae Marinković-Gospodnetić, 1988

Glossosoma neretvae Marinković-Gospodnetić, 1988: 45–47: “Glossoso-
ma neretvae n. sp. je nalažena samo u reci Neretvi i to na deonici od Vrapčića
ispred Mostara do Doljana (kod Metkovica). Odrasli oblici nalaženi su dva puta
godišnje, u proleće (april, maj, juni i početak jula) i u jesen (septembar, okto-
bar i novembar). U radu Marinković-Gospodnetić (1978) oznacena je imenom

Glossosoma neretvanus sp. n. (ali nije opisana ni ilustrovana). Holotip mužjaka,

alotip ženke i paratipovi mužjaka i ženke nalaze se u autorovoj kolekciji.”
Agapetus Curtis, 1834

Agapetus belareca Botoşăneanu, 1957
Agapetus belareca Botoşăneanu, 1957b: 188–189: “Première moitie de
juin 1956. Material pris à la lumière d’une lampe Petromax, à Mehadia (Banat); il
provient probablement de la rivière de Belareca. Holotype ² dans les collections
du Rijksmuseum van Natuurlijke Historie de Leyde. Paratype ² dans la collec-
tion de l’auteur. Material en alcool.”
Agapetus kampos Oláh, 2013

Agapetus kampos Oláh, 2013 in Oláh & Kovács (2013: 111–112): “Holo-
type. Montenegro, Bar municipality, Rumija Mts, Sutorman, Basa spring, N42°
09’25.6”, E19°06’06.3”, 770 m, 26.05.2013, leg. P. Juhász, T. Kovács, G. Magos, G.
Puskás (1 ², OPC). Allotype. Same as holotype (1 ³, OPC). Paratype. Same as
holotype (1 ², OPC).”
Diagnosis – Oláh & Kovács (2013: 111–112): “Its characteristic bifid
hook-headed structure of the phyllic organ differentiates this species from all the
known European species.”
Etymology – “Kampos from “kampós” (= hooked) in Hungarian, refers to
the hopoked head of the phallic organ.”
Agapetus montanus Kumanski, 1985

Synagapetus montanus Kumanski, 1985b: 76–79: “Pirin Mt., Kamenischki
Circus, small torrent following in the Great (3rd) Kamenischko-Lake (ca. 2200
m a.s.l.), 21.VII.1968, 6 mature pupae (3²²), one immature pupa, 2 praepupae
and 3 larvae; Rila Mt., tributary of Malyovishka River, 11.VII.1962, 1² and 1³
(leg. Novak). Holotype male chosen among the mature pupae from Pirin. The
other mature pupae and the adults from Rila designated as paratypes. All speci-
mens kept in the collection of the National Natural History Museum, Sofia.”
Agapetus montanus Kumanski, 1985: the genus Synagapetus was lowered to
subgeneric rank by Ross (1956: 144–149, 158–163).
Agapetus rectigonopoda Botoşăneanu, 1957

Agapetus rectigonopoda Botoşăneanu, 1957a: 63–64: “Dragan-Tal (Bihar-
Gebirge); 23.VI.1953. Iada-Tal (Bihar-Gebirge); 29.VII.1953. Holotypus ² (SMF
N4), Paratypoide: ³ (SMF N5) und ²³ (Sammlung des Verfassers), in Alkohol.”

136 J. Oláh
Agapetus slavorum Botoşăneanu, 1960

Agapetus slavorum Botoşăneanu, 1960a: 268–269: “J’ai eu à ma disposi-
tion 2² et 2³ de Trnovo, 23.VII.1955, qui ont été designés comme holotype ²,
allotype ³, paratype ² et paratype ³. Holot. ² allot. ³; F. Schmid, 1²+1³ parat.;
L. Botoşăneanu”.
HYDROPTILIDAE
Allotrichia McLahlan, 1880
Allotrichia marinkovicae Malicky, 1977
Allotrichia marinkovicae Malicky, 1977: 66–67: “Holotypus ² und zwei
Paratypen ²²: Hercegovina, Mostar, 26.VII.1916. Ferner (nicht als typen festge-
legt, aber vermutlich dazugehörig) von ebendort: 1³ vom 12.VII.1916 und 2³³
vom 27.VII.1916. Alle in coll. Naturhistorisches Museum Wien.”
Ithytrichia Eaton, 1873

Ithytrichia bosniaca Murgoci, Botnariuc et Botoşăneanu, 1948
Ithytrichia bosniaca Murgoci, Botnariuc et Botoşăneanu, 1948:
2–22: “La nouvelle espèce d’Ithytrichia à été trouvée en grand nombre dans les
eaux courantes presque patout où l’on a récolté ce matériel: dans la Bosna (30/
VII et 17/VIII 1947) à quelques kilomètres en aval de la gare de Kakany en face
du village du même nom, sur le gravier qui couvre le lit du fleuve et, surtout sur le
Potamogeton fluitans qui se développe en abondance dans certains endroits tran-
quilles du fleuve: 2) dans le ruisseau Rybnica (3/VIII.1947; T°de l’eau 17°) afflu-
ent de la Bosna pres du village Kakany; 3) dans la Lashva (12/VIII.1947; T°de
l’eau 18°) affluent gauche de la Bosna et 4) dans la Kusica (21/VIII.1947; T°de
l’eau 18°) affluent droit de la Lashva.”
Hydroptila Dalman, 1819

Hydroptila angulifera Kumanski, 1974
Hydroptila angulifera Kumanski, 1974: 71–74: “Rhodope, la rivière
Trigradska, juste avant sa confluence avec la rivière Tchairska (environ 800 metres
d’altitude), 2.VIII.1970, 5 paratypes (4² et 1³, nymphes) et 16 fourreaux vides;
20.VII.1971, 1² (Holotype) et 33 paratypes (17², 1³ et 15 nymphes mures dont 8²
et 7³). Holotype, 2² (1 adulte et 1 nymphe in sec) et 1³ paratypes (in sec) fixés sur
trois préparations microscopiques et les autres exemplaires – en alcool, sont gardés
dans la collection de l’Institut Zoologique et Musée près l’Académie Bulgare des
Sciences, à l’exception d’un ² paratype envoyé à la collection du D. H. Malicky,
Station biologique près l’Académie des Sciences de l’Autriche, Lunz am See.”

Hydroptila atalante Malicky, 1997

Hydroptila atalante Malicky, 1997: 147: “Holotypus ² und 5 Paratypen
²²: Bulgarien, Strandscha-Gebirge, 1 km S Kruschewez, 100 m, 18.VI.1980,
leg. Kumansky & Malicky, Sammlung Malicky, Lunz am See & Nationales
Naturwissenschaftliches Museum, Sofia.”
Hydroptila kalonichtis Malicky, 1972

Hydroptila kalonichtis Malicky, 1972: 30–31: “Holotypus ²: Kreta,
Kalonichtis, 24.IV.1971, leg. Malicky, Allotypoid ³ und Paratypoide ²², ³³ mit
den selben Daten. Zahlreiche weitere Paratypoide von drei weiteren Fundorten
in Kreta. Alle in meiner Sammlung.”
Hydroptila bureschi Kumanski, 1972: 1261–1263: “Balkangebirge, kleiner
Bach, Nebenfluss des Iskar beim Dorf Bov, 1.VII.1962, Holotypus ² (leg. Dr. K.
Novak); Rhodopen-Gebirge, Fluss Trigradska, 3 km unterhalb des Dorfes Trigrad,
2.VIII.1970, 1 ² Paratypes. Holotypus- und Paratypusmikroskoppräparate in
der vom Autor zusammengetragenen Sammlung des Zoologischen Instituts und
Museums der Bulgarischen Akademie der Wissenschaften.” Synonymised by
Botosaneanu & Malicky (1978b).
PHRYGANEIDAE
Phryganea Linnaeus, 1758
Phryganea ochrida Malicky, 1975
Phryganea ochrida Malicky, 1975: 85: “Holotypus ² und drei Paratypen
²²: Makedonien, Ochrid, 8.VII.1973, leg. R. Willmann, in meiner Sammlung.
Paratypus ²: Ochrid, “V 1.1835” (vermuthlich richtig: 1.V.1935), leg.
Wolfschlaeger, coll. Oberösterreichisches Landesmuseum Linz; Paratypus ²:
Ochrid, 24.VI.1954, leg. Thurner, coll. Hoelzel, Graz.”
BRACHYCENTRIDAE
Micrasema McLachlan, 1876
Micrasema sericeum Klapálek, 1902
Micrasema sericeum Klapálek, 1902: 164–165: “Pazaric, Krupthal 3 ²
(Winneguth), Stolac (Winneguth) 2³.”
LEPIDOSTOMATIDAE
Crunoecia McLachlan, 1876
Crunoecia bosniaca Marinković-Gospodnetić, 1971

138 J. Oláh
Crunoecia bosniaca Marinković-Gospodnetić, 1971a: 84: “Southeast

Bosnia, the small forest springs of tributaries of the Sutjeska, between Tjentiste
and Curevo”. Marinković-Gospodnetić (1971: 144): “Southeast Bosnia, ²²
³³ in small forest springs on the mountain Maglic.”
Crunoecia monospina Botoşăneanu, 1960

Crunoecia monospina Botoşăneanu, 1960a: 286–287: “J’ai eu à ma dis-
position 3² de Perister 12–16.VIII.1955, que j’ai designé comme holotype ² et
paratypes ²; holot. ²: F. Schmid; parat. ²: Deutsches Entomologisches Institut,
Berlin; parat.: L. Botoşăneanu.”
UENOIDAE
Thremma McLachlan, 1876
Thremma anomalum McLachlan, 1876
Thremma anomalum McLachlan, 1876: 266: “Greece (Parnassus, Krüper,
12th June); one pair in the Vienna Museum.”
APATANIIDAE
Apatania Kolenati, 1848
Apatania carpathica Schmid, 1954
Apatania carpathica Schmid, 1954: 11–12: “Cette espèce à été signalé des
Carpathes par Dziędzielewicz, sous le nom de meridiana. Elle est très voisine de
cette dernière forme et s’en distingue principalement par la forme du corps du
Xme segment. J’en ai vu 1² et 2³, que je designe comme holotype (²), allotype
(³) et paratype. Ils ont été capturés à Czarnohora et sont actuellement dans ma
collection.”
Apatania szczesnyorum Oláh, 2006

Apatania szczesnyorum Oláh, 2006: 11–12: “Holotype male: Poland, Polish
Tatras, Chocholowska Valley, 22. VIII. 1986, net leg. J. Oláh. Habitat. The adult
animal was resting on Carex canopy in a small stream-fed bog at the entrance of
a side valley in the middle section of Dolina Chocholowska. This small bog was
maintained by the tributary of the side valley. The stream water was spreading
and ending in this bog. This rare animal was accompanied with Apatania fimbri-
ata Pictet, which is an abundant species living in larger streams and small tribu-
taries of the whole Tatras. In this valley it was also just some minutes of sweeping
effort to collect 27 males.”

LIMNEPHILIDAE
Drusinae
Drusus Stephens, 1837
Drusus arbanios Oláh, 2010
Drusus arbanios Oláh, 2010: 98–100: “Holotype, male, HNHM. Albania:
Skrapar district, Ostrovicë Mts, Backë, stream beneath the pass between Mt.
Frengu and Mt. Faqekuq, 1913 m, N40°31.614’, E20°25.021’, 4.VII.2005, leg. Z.
Barina, D. Pifkó & D. Schmidt.”
Diagnosis – Oláh (2010: 98–100): “This dark species with almost ebony
black sclerites belongs to the species complex with large upward arching triangu-
lar gonopods and triangular or bilobed cerci inhabiting the Balkan Mountains.
Most close to Drusus illyricus sp. n., but differs by having (1) small animal with
forewing length of 8 mm, not large animal of 14 mm; (2) posterodorsal spinate
area on VIIIth tergite four-lobed in dorsal view, not trilobed; (3) the longitudinal
groove of IXth segment linear, not with ventral arm; (4) cerci with dorsal lobe
quadratic, not tapering in lateral view; (5) inner branch of paraproct forming
an almost regular quadratic plate in lateral view, not supplied with a dorsal pro-
nounced subapical wart-shaped process.”
Etymology – “The name was given to remind the old ancient city of Arbon
and his people the Arbanios, nearby the type locality.”
Drusus balcanicus Kumanski, 1973

Drusus discophorus Radovanović, 1942: Kumanski (1971: 102): “Stara
Planina, below “Tdza” (1500 m), 17. V. 1968, leg. P. Beron (3², 1³).” Misidentifi-
cation.
Drusus discophorus balcanicus Kumanski, 1973a: 113–114: “Die Fundorte
der neuen subspecies befinden in dem Zentral-Balkangebirge: Berghütte “Taza”
(1500 m Höhe), 17.V.1968, 3² und 1³; Bach bei der Berghütte “Raj”, 1³ und sub-
alpiner Bach (1900–2000 m Höhe), rechter Zufluss des Baches Taza, 10.VI.1971,
massenhaft anzutreffen (insgesamt 43² und 10³ gesammelt.”
Drusus discophorus balcanicus Kumanski, 1973: Kumanski & Malicky
(1976: 104): “Stara Planina, Teteven, Beli Vit, Ribariza und Nebenbache, 600
m, 15–18.V.1969, 1 male. Stara Planina, Trojan Pass, Einzugsgebiet des Tscherni
Osam, 1400–1600 m, 20.V.1969, 2 females.”
Drusus balcanicus Kumanski, 1973: Kumanski (1981: 142): raised to spe-
cies status.
Diagnosis – Kumanski (1973a: 113–114): “In allgemeinen Linien nach
dem Bau von D. discophorus discophorus konstruiert.”

140 J. Oláh
Drusus bosnicus Klapálek, 1899

Drusus bosnicus Klapálek, 1899b: 327–329: “Ilidže i Vrelo Bosne (Apfelb.),
Pazarič, Dolina Krupe, 3. novembra 1898. (Apfelb.).”
Drusus bosnicus Klapálek, 1899: Klapálek (1900: 674–675): “Ilidže und
Vrelo Bosna (Apfelb.), Pazarič, Krupathal, 3. November 1898 (Apfelb.).
Drusus bosnicus Klapálek, 1899: Schmid (1956: 31–32): “Cette espèce à été
décrite de Bosnie; M. Radovanović l’ à signalée de Sarajevo. J’ai étudié un assez
grand nombre d’exemplaires que j’ai capturés a mi-juilett en Bosnie au massif de
Treskavitza.”
Drusus bosnicus Klapálek, 1899: Oláh et al. (2017: 95): “Bosnia & Herze-
govina: Federation of Bosnia and Herzegovina, Sarajevo, Ilidža, Bosna springs,
43°49’08.4”, 18°16’09.4”, 645 m, 2.X.2015, leg. P. Juhász & T. Kovács (26 males,
14 females, OPC).”
Diagnosis – Klapálek (1900: 675): “Die Analanhänge dieser Art zeigen
eine grosse Aehnlichkeit mit den homologen Theilen des Dr. monticola, und ich
würde es nicht wagen sie als eine distincte Art zu beschreiben, ware nicht der
Unterschid in der Färbung und in der Form der Discoidalzelle so auffalend; auch
die Form der oberen Anhänge ist etwa verschieden.”
Schmid (1956: 32): “Dr. bosnicus est sans doute l’espèce la plus évoluée du
groupe; ceci est visible à ses appendices de forme très spécialisée: IXe segment
concave lateralement, appendices inférieurs très divergent, appendices interme-
diaires formant une assez grande surface propongeant la zone spinulifera.”
Kučinić et al. (2015): larval head shape relates D. bosnicus to D. ramae, al-
though the present efforts in larval research focus rather on morphological sepa-
rations than on phylogenetic relations. Surprisingly recent molecular study sug-
gested two different species from the Drusus bosnicus species group coexisting
in the spring area of Bosna River, but so far, there was no any records of adult of
last instar larvae of this possible new species. The fine structure analysis carried
out on the paraproct, paramere and vaginal sclerite complex of 26 males and 14
females, collected in the autumn of 2015 in the spring area of Bosna River, has
demonstrated the presence of only a single species: Drusus bosnicus.
Drusus botosaneanui Kumanski, 1968

Drusus botosaneanui Kumanski, 1968a: 214–216: “Fundort: Vitosa-Gebir-
ge, Bojanski-Bach bei der Berghütte “Bor” (1600 m Höhe), 3.IX.1967. Holotypus
(mit getrenntem Abdomen) und 2 Paratypen (alles in alcohol) in der Sammlung
des Autors.”
Diagnosis – Kumanski (1968a: 214–216): “Drusus botosaneanui gehört zur
Gruppe von annulatus (nach Schmid, 1956) und steht tenellus am nachsten.”

Drusus brunneus Klapálek, 1898

Drusus brunneus Klapálek, 1898a: 489: “Mehádia, Corniareva, Marmaros,
Boroszno. 6², 4³.”
Drusus brunneus Klapálek, 1898: Klapálek (1899a): 434–435, 489: Drusus
brunneus Klapálek, 1898: Schmid (1956: 48–49): “Cette espèce est localisée dans
les montagnes de l’Europe orientale: j’ai vu une dizaine d’individus, provenant de
Hongrie et de Roumanie.”
Diagnosis – Klapálek (1898: 489): “Habituell dem D. Muelleri sehr ähn-
lich.” Klapálek (1899: 434–435): “Obwohl diese Art durch ihr Aeusseres stark
an den Drusus Muelleri erinnert, so lässt sie sich durch die Genitalanhänge
sogleich von demselben unterscheiden.”
Schmid (1956: 49): “Dr. brunneus est caractérisé par ses deux paires d’ailes
brun-jaune foncé, par ses appendices intermédiaires armés de deux pointes très
petites, par le corps du Xe segment proéminent et par ses appendices inférieurs
assez petits et peu velus.”
Drusus bureschi Kumanski, 1973

Drusus bureschi Kumanski, 1973a: 114–117: ”Östliche Balkangebirge, a)
Eleno-Twardischki-Pass (1000 m Höhe), 25.V. 1969, 1² (leg. Al. Popov); b) Längs
eines kleinen Berbaches, 4 km östlich vom Pass und c) Bei einem ungestümen
Bergbach, am Beginn des Twardischka-Baches (eines Zuflusses der Tundza), un-
gefähr 10 km östlich von Pass, 12.VI.1971, ingesamt 3² und 4³; d) Balkangebirge
bei der Stadt Sliwen, Örtlichkeit Karandilja, 21.V.1969, 1² (leg. Al. Popov). Als
Typusexemplar bezeichnete ich ein ² von Fundort c). Dasselbe sowie auch die
übrigen Exemplare sind in Alkohol in der Sammlung des Zoologischen Instituts
mit Museum bei der Bulgarischen Akademie der Wissenschaften aufbewahrt.”
Drusus bureschi Kumanski, 1973: Kumanski (1975: 63): Bulgaria, Stara
Planina, Sliven, 4. VII. 1911, 1 male. Bulgaria, Below Raj, Basmandra, 1800 m,
22. VIII. 1970, 32 female.
Drusus bureschi Kumanski, 1973: Kumanski & Malicky (1976: 105): Bul-
garia, Stara Planina, Sliven, Oberlauf der Stara Reka, 500–800 m, 22. V. 1969, 1
male. Bulgaria, Stara Planina, Sliven, Tundscha-Einzugsgebiet mit Nebenbachen,
300–600 m, 23. V. 1969, 1 male.
Diagnosis – Kumanski (1973a: 114–117): “Nach vielen ihrer Merkmale
steht die neue Art D. discophorus ziemlich nahe; letztere galt als eine in dem
Rahmen der Gattung isolierte Art (Schmid, 1956). Die Affindung ihrer Unterart
balcanicus in der Balkangebirgen sowie auch das Antreffen von D. bureschi
erlaubt die Vereinigung der drei Taxa in eine neue Gruppe – die Gruppe von
discophorus. Die Zusammengehörigkeit dieser Gruppe beruht bei den ² auf

142 J. Oláh
dem Vertiefung der dorsalen Oberfläche des VIII. Tergiten, dem Gleichen
Aussehen der App. Inferiores und dem gemeinsamen Schema des X. Segments
mit App. Intermediales und bei den ³ auf der allgemeinen Ähnlichkeit der
Genitalstructuren. Die Eigenheiten, die D. bureschi individualisieren, sind fol-
gende: eine breite zona spinulata am VIII. Tergit; die eigenartige Form und die
weit entfernten Spitzen der App. Intermediales bei den ² und die zugespitzten
seitlichen Teile des X. Segments bei den ³.”
Drusus buscatensis Botoşăneanu, 1960

Drusus buscatensis Botoşăneanu, 1960: 369–370: “Holotypus ²: Băişoara
– Muntele Buscat (Rumanische Westliche Karpaten – Munţii Apuseni); Neben
einer Quelle, 21.V.1956 (leg. Kiss Béla, Sammlung L. Botoşăneanu). Paratypus ²:
selber Ort, 16.V.1956 (leg. L. Botoşăneanu, Samlung F. Schmid).”
Drusus buscatensis Botosaneanu, 1960: Botosaneanu (1975: 97–98):
“Drusus buscatensis Bots. ist einerseits vom Apuseni-Gebirge, anderseits vom
Cibin- oder Cindrel-Gebirge (Südkarpaten) bekannt. Frühlingsart (Mai); lebt in
Quellen und Bächlein in der Fichtenwaldzone (etwa 1400–1600 m).
Drusus buscatensis Botoşăneanu, 1960: L. Újvárosi (pers comm.) has collect-
ed 1² near the locus typicus (RO, Cluj county, Apuseni, Muntele Băişorii, Buscat,
springs 46.537505°N, 23.291260°E, 1529 m, 12.V.2012) and 1² near Paltinis,
Munţii Cindrel (RO, Sibiu county, Southern Carpathians, Complex Paltinis,
Cibin, Batrana springs 45.620050°N, 23.893600°E, 1720 m, 2.VI.2010).
Diagnosis – Botosaneanu (1975: 97–98): “Sicherlich nahe mit D. doehleri
verwandt, die ihre Schwesterart ist. Es ware interessant zu wissen, wann und un-
ter welchen Umstanden die Artaufsplitterung stattgefunden hat.”
Drusus carpathicus Dziędzielewicz, 1911

Drusus carpathicus Dziędzielewicz, 1911a: 206–209: “In Karpathibus
Orientalibus in regione alpina (Pini Mughi) apud fontes. Mons Chomiak 27.V. –
11.VI.1909 (supra 1300 m. s. m.). Czarnohora in montibus Dancerz et Howerla
30.V.1909. – 10.VI.1910.”
Drusus carpathicus Dziędzielewicz, 1911: Szczęsny (1980): specimen in
NHM-ISEA: 1²; this specimen is labelled with the following notes: “Czarnohora,
Dancerz 31.V.1911” and “Drusus carpathicus Dz. ² ver. C. Tomaszewski”, and
is registered in the inventory under number 8/25. Dziędzielewicz (1911)
described the species on the basis of specimens collected “near the summit of
Chomiak by springs of the stream Roskolski during the period 27.V.-11.VI.1909
and at Czarnohora near Howerla and Dancerz on the days 30.V.1909 and
10.VI.1910”. Dziędzielewicz also noted that all the specimens collected in this

period were deposited in the museum at Lvov. Therefore the male deposited at
Cracow does not belong to this series of specimens which enabled Dziędzielewicz
to describe a new species, nevertheless it does come from the “locus typicus”.
Drusus carpathicus Dziędzielewicz, 1911: Szczęsny & Godunko (2007):
“3²², 3³³; East Carpathians, Czarnohora Massif, Gorgany Massif (Chomiak);
1² and 1³ collected on slopes of Chomiak Mt, 3.VI.1909 and 11.VI.1909, respec-
tively (No E24.12.06.04/02 and 03), and 1³ caught 9.VI.1910 in the Czarnohora
massif (No E24.12.06.04/01) belong to the series of specimens on the basis of
which Dziędzielewicz described the species; the male from Chomiak is desig-
nated as a lectotype herein. To that series belong also several specimens stored in
NMP (P. Chvojka, pers. comm.).”
Drusus carpathicus Dziędzielewicz, 1911: Szczęsny & Chvojka (2008):
altogether 2²² and 4³³ paralectotype specimens were collected during 27.V.-11.
VI.1909 in the East Carpathians (Chomiak) and deposited in NMP.
Diagnosis – Dziędzielewicz (1911a: 206–209): “Habitu coloreque Druso
bosnico Klap. similis. Capite nigro, in fronte et occipite nigro piloso; protorace
rufo, nigro piloso; meso et metathorace nigris; abdomine griseo. Appendices
praeanales maris oblongae, lappaceae, pilosae; ungues praeanales perlongi, recti,
apice hamato; pedeg genitales bipartiti, in aspectu supero excavati, apice arcuato
exciso, in aspectu laterali conici, paullulum curvat.”
Drusus crenophylax Graf et Vitecek, 2015

Drusus crenophylax Graf et Vitecek, 2015 in Vitecek et al. (2015c: 88): “Ho-
lotype. 1 male: Bosnia and Herzegovina, Cvrka river; 44°32.932’N 17°23.562’E;
393 m a.s.l.; 1. X. 2014; leg. Dejan Dmitrovic, Goran Sukalo, specimen identifier:
f Dsp4501M. Paratypes: 2 females: Bosnia and Herzegovina, Spring of Cvcka
river, Vilenjska vrela; 44°33.003’N 17°23.580’E; 456 m a.s.l.; 12. IX. 20124;
leg. Dejan Dmitrovic, specimen identifier: f Dsp3401F. 4 males, 3 females, 19
larvae: Bosnia and Herzegovina, Cvrka river; 44°33.003’N 17°23.580’E; 456 m
a.s.l.; 12. IX. 2012; leg. Dejan Dmitrovic, Goran Sukalo, Goran Sukalo; speci-
men identifier for 3 larvae: f Dsp4502L, f Dsp4503L, f Dsp4504L. Holotype and
paratypes currently in coll. W. Graf, will deposited in the Biologiezentrum des
Oberösterreichischen Landesmuseums, Linz, Austria.”
Drusus crenophylax Graf et Vitecek, 2015: Oláh et al. (2017: 96): “Bosnia
& Herzegovina: Republika Srpska, Večići, Cvrcka river, 44°32’55.4”, 17°23’33.7”,
1.X.2015, leg. P. Juhász & T. Kovács (1 male, OPC).”
Diagnosis – Vitecek et al. (2015c: 88): “Males of the new species are most
similar to Drusus discophorus Radovanović and D. vernonensis Malicky, but ex-
hibit (1) subtriangular superior appendages I n lateral view, (2) subtriangular,

144 J. Oláh
low tip of the intermediate appendages in lateral view, and (3) simple, rounded
tips of intermediate appendages in caudal view. Drusus discophorus males have
suboval superior appendages and a high round tip of the intermediate appendage
in lateral view; D. vernonensis males have round superior appendages in lateral
view and trilobate tips of intermediate appendages in caudal view. Females of the
new species show the reduced median lobe of the vaginal sclerite and gigh base
of the lateral lobe of segment IX as typical for Balkan Drusinae and most similar
to Drusus vernonensis. Larvae of the new species are most similar to Drusus kla-
paleki Marinković-Gospodnetić and D. serbicus Marinković-Gospodnetić.”
Drusus croaticus Marinković-Gospodnetić, 1971

Drusus croaticus Marinković-Gospodnetić, 1971c: 105–107: “Sources
of the river Crna Reka, Plitvice, 30m, 1f, 13. V. 1971.”
Diagnosis – Marinković-Gospodnetić (1971c: 105–107): “Drusus croat-
icus is, probably, a member of the group discolor according to the form of inter-
mediate appendages, which are the most similar to those of D. macedonicus and
D. transylvanicus.”
Drusus dacothracus Oláh, 2010

Drusus dacothracus Oláh, 2010: 100–102: “Holotype, male, HNHM.
Albania: Dibër district, Dejë Mts, Varoshit Stream and its karst cave sidespring at
Shkanderbeu Cliff, W of Murrë Pass, 975 m, N41°38.792’, E20°11.390’, 11.X.2005,
leg. T. Deli & D. Murányi. Paratypes, HNHM. Albania: Dibër district, Dejë
Mts, Varoshit Stream and its karst cave sidespring at Shkanderbeu Cliff, W of
Murrë Pass, 975 m, N41°38.792’, E20°11.390’, 13.IV.2006, leg. Z. Erőss, Z. Fehér,
A. Hunyadi & D. Murányi (1 male). Dibër district, Dejë Mts, Varoshit Stream
and its karst cave sidespring at Shkanderbeu Cliff,Wof Murrë Pass, 975 m, N41°
38.792’, E20°11.390’, 11.X.2005, leg. T. Deli & D. Murányi (1 male, 3 associated
females). Dibër district, Lurë area, Cidhnë, aqueduct above Setë Stream in its
gorge, W of the village, 780 m, N41°45.149’, E20°14.732’, 10.X.2005, leg. Z. Erőss
& D. Murányi (1 male). Skrapar district, Ostrovicë Mts, Çeremica, brookWof the
village, 1820 m, N40°32.649’, E20°26.573’, 5.VII.2005, leg. Z. Barina, D. Pifkó &
D. Schmidt (1 male).”
Diagnosis – Oláh (2010: 100–102): “This dark species belongs to the spe-
cies complex with large upward arching triangular gonopods and triangular or
bilobed cerci inhabiting the Balkan Mountains. Most close to Drusus illyricus sp.
n., but differs by having (1) smaller size; (2) upper lobe of the trilobed spinate
area on tergite VIII narrow and clearly monolobed, not broad bilobed with
some mesal depressen; (3) sternal lateral suture of the fused IXth segment with-

out middle fork, not with well-developed middle ventral brach; (4) segment IX
not very long ventrally; (4) cerci with dorsal lobe short triangular, not long and
downward curving; (5) the ventral lobe of the cerci long triangular, not long
digitiform; (6) inner branch of paraproct differently shaped both in lateral, dor-
sal and caudal view.”
Etymology – “The name was given to remind Dacian-Thracian origin of the
Albanian people inhabiting the type locality. Three theories exist: the Illyrian,
Dacian-Thracian and Pelasgian origin of the Albanians.”
Drusus dardanicus Ibrahimi, Kučinić et Vitecek, 2015

Drusus dardanicus Ibrahimi, Kučinić et Vitecek, 2015 in Ibrahimi et al.
(2015: 558–561): “Type material. Holotype (1 male) and paratypes (2 males):
Kosovo: Podujeve Municipality, Shatorice Mountains, stream above Bollosice
Village, 1330 m a.s.l., 43.118169°N, 20.99330°E, 11.V.2014, leg Halil Ibrahimi.
Holotype deposited in the department of Biology, faculty of Mathematics and
Natural Scienxes, University of Prishtina “Hasan Prishtina”, Prishtine, Republic
of Kosovo. Paratype (3 males): Same collection and locality data, deposited in
the Croatian Natural History Museum, Zagreb (coll. Kučinić -Trichoptera),
Croatia. Paratypes (3 males): Same collection and locality data, deposited in the
Biologiezentrum des Oberösterreichischen Landesmuseum, Linz, Austria; speci-
men identifiers for 2 males: f Dsp4301M, f Dsp4302M.”
Diagnosis – Ibrahimi et al. (2015: 558–561): “Males of the new species are
most similar to Drusus discophorus, D. bureschi, and D. balcanicus.”
Drusus discophoroides Kumanski, 1979

Drusus discophoroides Kumanski, 1979: 67–70: “Belassitza Mt. (South-
Western Bulgaria), brooklet 1.5 km western from the mountain hostel “Belassitza”
(750 m a.s.l.), 10.VI.1975, 2²² (leg. Kumanski). Holotype ² and 1² Paratype in
the author’s collection in the National Natural History Museum Sofia.”
Diagnosis – Kumanski (1979: 67–70): “D. discophoroides n. sp. is near to D.
discophorus Rad. Both species have common plan of ² genitalia. This notwith-
standing they can easily be distinguished both after the darker general colour of
the new species and several details of the male genitalia: the upper appendages
of D. discophoroides n. sp. are so small that the intermediate appendages remain
wholly visible from the side; in D. discophorus the latter almost invisible; the
upper margin of the intermediate appendages rounded in the new species and
formed as an undulate horizontal line in discophorus; the perianal region oval
rounded (discophorpides n. sp.) or with nearly straight side borders (discophorus).”

146 J. Oláh
Drusus discophorus Radovanović, 1942

Drusus discophorus Radovanović, 1942: 184–190: “An den Quellen und
Bachen am 1. und 3. Gebirgssee auf dem Jablanitza-Gebirge.” “Dieses Gebirge
liegt etwa 16 km nordwestlich von Struga am Ochridsee und bildete damals das
Grenzgebiet zwischen Albanien und Jugoslawien. In einer Höhe von etwa 1960
m befinden sich vier Gebirgsseen von verschiedener Grösse. Diese liegen im
Bereiche der am Fusse des Gebirges gelegenen Dörfer Labuniste, Podgoratz und
Vevcane und werden auch nach diesen Dörfern als Labunisko (1. Und 2. See),
Podgoracko (3. See) und Vevcansko Jezero (4. See; jezero = See) genannt. Die
drei erstgenannten Seen konnte ich gelegentlich meines Besuches (13.–16. VII.
1939) ziemlich eingehend trichopterologisch erforschen.” ”Der 1. Gebirgsee auf
der Jablanitza (“1. Labunisko Jezero”) hat eine etwa trapezförmige Gestalt und
geht mit seinem nördlichen Winkel in einen gegen Labuniste abfl iessenden Bach
über der bald danach unterirdisch verschwindet. Westlich von See, unter dem
Gebirgsrücken, befinden sich einige kräftige Quellen, deren Wasser dem See zu-
fl iesst. Der 3. See (Podgoracko Jezero) liegt etwa 1300 m südlich von 1. See und
ist bedeutend grösser als die zwei vorher geschilderten.” “In dem Trichopteren-
Material, das mir nachträglich von Herrn Winneguth aus Sarajevo zum Bestimmen
nachgesandt wurde, befanden sich etliche Exemplare, die ebenfalls zu dieser Art
gehören. Von diesen waren zwei Stück am Limflusse bei Andrijewitza erbeutet,
und die zwei anderen (sämtlich ²) an der Wrujaquelle bei Gusinje.”
Diagnosis – Radovanović (1942: 184–190): Compared to Drusus bosnicus
and D. graecus among the known species in the Balkan Peninsula. “Von dieser
Art [Drusus graecus] sowie von Drusus bosnicus Klap. Unterscheiden sich die
Exemplare von der Jablanitza (Drusus discophorus) und von Labuniste (Drusus
plicatus) nach einigen augenscheinlichen charakteristischen Merkmalen, vorw-
iegend nach der Form der Genitalanhänge und nact der Gestalt des 8. Tergits.”
Drusus ekes Oláh, 2017

Drusus ekes Oláh, 2017 in Oláh et al. (2017: 150–151): “Holotype:
Romania, Apuseni Mts., Vlădeasa Mt., Stâna de Vale, upper section of Ciripa
stream, N46°40.546’ E22°38.515’, 1360 m, 6.VI.2015, leg. M. Kiss, J. Oláh & L. Szél
(1 male, OPC). Allotype: same as holotype (1 female, OPC). Paratypes: same as
holotype (12 males, OPC). Apuseni Mts., Vlădeasa Mt., Stâna de Vale, Galbenele
stream, N46°40.809’ E22°37.147’, 1180 m, 7.VI.2015, leg. M. Kiss, J. Oláh & L.
Szél (2 males, OPC). Apuseni Mts., Mt. Bihor, Gârda de Sus, tributary of Arieşul
Mare, N46°270.493’ E22°47.895’, 788m, 29.V.2013, singled leg. J. Oláh, E. Bajka,
Cs. Balogh, & G. Borics (1 male, OPC). Apuseni Mts. Munţii Gilăului, Staţiunea

Muntele Băişorii, Lupinus stream, 18.VI.2013, singled leg. J. Oláh, Cs. Balogh,
& S. Fekete (1 male, OPC). Apuseni Mts. Munţii Gilăului, Staţiunea Muntele
Băişorii, La Mocirle, spring streams, N46°30.241’ E23°15.550’, 1552m, 19–20.
VI.2015, singled leg. J. Oláh, Cs. Balogh, & P. Juhász (1 male, OPC). Apuseni
Mts., Vlădeasa Mt., Stâna de Vale, upper section of Ciripa stream, N46°40.546’
E22°38.515’, 1360 m, 6.VII.2016, leg. J. Kecskés (3 females, OPC). Stana de Vale,
3. VI. 1956, leg V. Cirligat (1 male, registration number TRH136/013, CCPC).”
Diagnosis – Oláh et al. (2017: 150–151): “The hump on the apical margin
of the paraproct, that is the only recognised paraproct divergence between D.
chrysotus and D. romaicus complexes, is very decisive and pronounced at D. ekes
sp. n. The lateral profile of the fused dorsal branches of paraproct has the flat top
sloping anterad and produced posterad, similar to D. ferdes, differing from the
lateral configuration of the other three species. The top configuration of the spe-
ciation trait of paraproct is rather stable in the sampled populations of the three
main montain ranges of the Apuseni Mts.: Vladeasa Mt., Bihor Mt., and Gilaului
Mt. The subapical spines, that is the true terminal of the paramere shaft long and
armed basad with a few small tertiary spine like structures; the apparent termi-
nalia of modified setal origin is very long and thin. This paramere pattern differs
from the paramere patterns of the other three species.”
Etymology – “Ekes”, from „ékes”, supplied with wedge in Hungarian, refers to
the flat top of the paraproct sloping anterad and produced posterad into a wedge
shape. But “ékes”, has the peculiar meaning of glory, an outstanding beauty, more
than beautiful, thanks to the fractal nature of the Hungarian language. The given
name refers to both characters, this animal is really elegant and beautiful.”
Drusus fortos Ibrahimi et Oláh, 2017

Drusus fortos Ibrahimi et Oláh, 2017 in Oláh et al. (2017: 155–157):
“Holotype: Kosovo: Çakor, streamlet along the border line between Kosovo
and Montenegro. 42.685542°N, 20.053636°E, 1289 m, 25.VIII.2015 leg. E.
Dimitrou & V. Dragobia (1 male, DBFMNSUP). Allotype: Lloqan, Krojet e
Gucise, 42.55143°N, 20.1335°E, 2110 m, 3.VIII.2016, leg. H. Ibrahimi (1 female,
DBFMNSUP). Paratypes: same as holotype (1 male hybrid, OPC). Haxhaj,
spring area of a tributary of Lumbardhi i Pejës River, 42°42’30N 20°2’32E,
1278 m, 25.VIII.2015 leg. E. Dimitrou & V. Dragobia (1 male, OPC). Lloqan,
Gurrat e Hasan Agës springs, Bjeshkët e Nemuna. 42.557155°N, 20.152696°E,
1991 m, 18.VIII.2015 leg. H. Ibrahimi (1 male, DBFMNSUP). Lloqan, Gurrat
e Hasan Agës springs, Bjeshkët e Nemuna. 42.557155°N, 20.152696°E, 1991 m,
3.VIII.2016, leg. H. Ibrahimi (3 males DBFMNSUP). Lloqan, Krojet e Gucise,
42.55143°N, 20.1335°E, 2110 m, 3.VIII.2016, leg. H. Ibrahimi (2 males OPC).”

148 J. Oláh
Diagnosis – Oláh et al. (2017: 155–157): “The lateral profile of the fused
dorsal branches of paraproct has short dorsoapical digitiform process with me-
dium thickness, most similar to name bearing species D. siveci, but differs by hav-
ing variously produced hump on the apical margin of the paraproct, completely
lacking at D. siveci; by the slightly anterad turning and tapering apex of the fused
digitiform dorsoapical process, vertical and not tapering at D. siveci; and the very
tip of the fused dorsal branches of the paraproct is entirely fused into a narrow
and straight pencil-like black process without any mesal suture, that is the vesti-
gium of the fusion surface completely disappeared as visible in caudal view, the
tip is bilobed and the mesal suture is discernible both at D. siveci and at D. vekon
sp. nov. The subapical spine of the paramere is robust bearing small dorsal spine
or corrugations. In the sampled habitats D. fortos sp. nov. lives together with D.
vekon sp. nov. with hybrid forms.”
Etymology – “Fortos”, from „összeforrt”, fused in Hungarian, refers to the
completely fused state of the dorsoapical digitiform process of the paraproct.”
Drusus gombos Oláh, 2013

Drusus gombos Oláh, 2013 in Oláh & Kovács (2013: 112–114): “Holotype:
Montenegro, Žabljak municipality, Sinjajevina Mts, Dobrilovina, forest stream
at the monastery, N43°01.652’, E19°24.086’, 765 m, 25. V .2013, leg. P. Juhász,
T. Kovács, G. Magos, G. Puskás (1², OPC). Allotype. Same as holotype (1³,
OPC). Paratypes. Same as holotype (2 ³, OPC).”
Diagnosis – Oláh & Kovács (2013: 112–114): “This castanean brown
species belongs to the species complex with large upward arching triangular
gonopods inhabiting the Balkan Mountains. Most close to Drusus klapaleki
Marinkovic, but differs by having cerci less slender with more bulky ventro-
apical corner; lateral profile of the paraproct different and female has trilobed
apical margin on the anal tube in dorsal view, not bilobed; the median lobe on
the vulvar scale present, not absent; the dorsal profile of the vaginal sclerite co-
plex different.”
Etymology – “Gombos, from “gombos” buttom-like in Hungarian, refers to
the rounded knob-shaped dorsoapical lateral processes of the paraproct.”
Drusus illyricus Oláh, 2010

Drusus illyricus Oláh, 2010: 102–104: “Holotype, male, HNHM. Albania:
Mat district, Kreshtës Mts, Vajkal, Fusha e Kaliut, brook N of the village, 1730 m,
N41°32.508’, E20°12.390’, 30.V.2008, leg. Z. Barina, D. Pifkó & B. Pintér.”
Diagnosis – Oláh (2010: 102–104): “This dark species belongs to the spe-
cies complex with large upward arching triangular gonopods and triangular or

bilobed cerci inhabiting the Balkan Mountains. Most close to Drusus pelasgus sp.
n., but differs by having (1) larger size; (2) sternal lateral suture of the fused IXth
segment with middle fork, not without; (3) segment IX very long ventrally, not
medium long; (4) cerci with dorsal lobe slender, downward curving, not blunt
rounded; (5) inner branch of paraproct with short and narrow dorsal apex as vis-
ible both in lateral, dorsal and caudal view, not long and broad.”
Etymology – “The name was given to remind one possible origin of the
Albanian people inhabiting the type locality. Three theories exist: the Illyrian,
Dacian-Thracian and Pelasgian origin of the Albanians.”
Drusus juliae Oláh, 2011

Drusus juliae Oláh, 2011a: 113–1174: “Holotype male. Albania: Mirdite
district, Oroshi area, Nanshene, open stream in the village, N41°51.848’,
E20°07.088’, 1175 m, 21.05.2010, leg. D. Murányi (1 male, HNHM). Paratypes:
same as holotype (4 males, 2 females, HNHM)”.
Diagnosis – Oláh (2011a: 113–1174): “This castanean brown species be-
longs to the species complex with large upward arching triangular gonopods in-
habiting the Balkan Mountains. Most close to Drusus radovanovici Marinković-
Gospodnetić, but differs by having sternal lateral suture of the fused IXth seg-
ment curving, not straight; cerci subquadratic, not subtriangular in lateral view;
inner branch of paraproct with rounded lateral lobes, not with triangular in dor-
sal view.”
“Etymology – “Patronym in honor of my wife Erzsébet Julia Tóth, who ac-
companies and supports my various activities on science.”
Drusus kerek Oláh, 2011

Drusus kerek Oláh, 2011a: 114–116: “Holotype male. Albania: Prokletije
Mts, creek above village, Sylbice, N42°31.326’, E20°05.919’ 2010 m, 8.07.2011,
leg. Z. Barina, A. Kovács, G. Puskás, B. Sárospataki (1 male, HNHM). Paratypes:
same as holotype (2 females, HNHM). Prokletije Mts, S slope of Mt maja e
Shpatit above village Doberdol, rocky creek covered with tall herbs, N42°33.040’,
E20°05.919’ 1940 m, 9.07.2011, leg. Z. Barina, A. Kovács, G. Puskás, B. Sárospataki
(1 male, 1 female, HNHM). Prokletije Mts, valley of stream Topoje, small side
creek in village Sylbice, N42°30.557’, E20°08.054’, 1580 m, 8.07.2011, leg. Z.
Barina, A. Kovács, G. Puskás, B. Sárospataki (1 female, HNHM). Prokletije Mts,
valley of stream Topoje below village Sylbice, tall herb community of the creek,
N42°30.283’, E20°08.917’, 1460 m, 8.07.2011, leg. Z. Barina, A. Kovács, G. Puskás,
B. Sárospataki (3 males, 2 females, HNHM).”

150 J. Oláh
Diagnosis – Oláh (2011a: 114–116): “This dark brow species belongs to the
species complex with large upward arching triangular gonopods inhabiting the
Balkan Mountains. Most close to Drusus juliae sp. n. but differs by having sternal
lateral suture of the fused IXth segment curving, not so deep; cerci rounded, not
subquadratic in lateral view; inner branch of paraproct more robust; apical third
of gonopods more tapering. Female: segment IX triangular in lateral view, not
subquadrangular, lateral setose lobe double long than high, not similar; supra-
genital plate not regular quadrangular in lateral view; median lobe of the vulvar
scale (lower vaginal lip) enterily lacking; lateral lobes of the vulvar scale differ-
ently shaped.”
Etymology – “Kerek, from “kerek” round in Hungarian, refers to the rounded
cerci.”
Drusus klapaleki Marinković-Gospodnetić, 1971

Drusus klapaleki Marinković-Gospodnetić, 1971a: 80: “Southeast Bos-
nia, mm and ff in small springs of tributary of the river Sutjeska, between Tjen-
tiste and Curevo.”
Drusus klapaleki Marinković-Gospodnetić, 1971: Marinković-Gospod-
netić (1971b: 144): “Southeast Bosnia, ²² ³³ in small springs of tributary of the
river Sutjeska.”
Drusus klapaleki Marinković-Gospodnetić, 1971 in Oláh & Kovács
(2013: 115–116): “Bosnia-Herzegovina: Jablanica, spring stream, 04.09.1988,
singled, J. OLÁH (1², 1³, OPC).” “The female of this species is unknown. We
have collected a single female from a spring area together with three males. Here
we describe the female and redraw the male in order to compare it with its close
relative, with Drusus gombos sp. n. Compared to male of D. gombos, D. klapaleki
has more slender cerci, differently shaped paraproct having apicolateral lobes
hook-shaped, and slightly different spine pattern of the paramere.”
Diagnosis – Marinković-Gospodnetić (1971a: 80): “This species is close-
ly related to D. bosnicus Klap. The greatest difference between these two species is
in the shape of the superior appendages and of the intermediate appendages.”
Drusus komanus Oláh, 2017

Drusus krusniki Malicky, 1981: Oláh & Kovács (2015: 109): misidentifi-
cation.
Drusus komanus Oláh, 2017 in Oláh et al. (2017: 110): “Holotype: Albania,
Shkodër district, Prokletije Mts, Mollë, Maljag Stream on the right bank of
Koman Lake, N42°11.673’, E19°49.063’, 185 m, 18.06.2012, leg. Z. Fehér, T.
Kovács, D. Murányi (1 male, OPC).

Diagnosis – Oláh et al. (2017: 110): “The D. komanus is closest to D. krus-

niki and D. kerek, but differs from both by having the dorsal branches of the
paraproct, that is the speciation trait, differently shaped. In lateral view both D.
krusniki and D. kerek characterized by having a hump subdorsad on the apical
margin of the paraproct. At D. komanus this hump structure is modiofied into
a triangular process well visible both in lateral and dorsal view. In caudal view
the dorsal margin of paraproct clearly V-shaped at the new species, not straigh
as at D. krusniki or shallow V-shaped as at D. kerek. The transversally plate form-
ing measally touching pair of the dorsal branches of the paramere, that is the
transversal plate characteristic for the species complex is differently formed at all
the three species. This plate configuration of the speciation trait visible in caudal
view is very stable. Stability is the result of selective, non-random, non-neutral
processes.”
Etymology – “Named after the locus typicus.”
Drusus kovacsi Oláh, 2017

Drusus dacothracus Oláh, 2010: 100: “Albania, Skrapar district, Ostrovicë
Mts, Ceremica, brook W of the village, 1820 m, N40°32.649’, E20°26.573’,
5.VII.2005, leg. Z. Barina, D. Pifkó & D. Schmidt (1 male).” Misidentification.
Drusus arbanios Oláh, 1910: Oláh & Kovács (2012: 90–91): female de-
scription and drawings. “Allotype female. Albania: Skrapar district, Ostrovicë
Mts, Backë, Krojmbret Spring and its outlet brook NE of the village, N40°31.753’
E20°25.152’, 1965 m, 12.10.2012, leg. P. Juhász, T. Kovács, D. Murányi, G.
Puskás (1³, OPC). Same as allotype (3², 6³, OPC). Skrapar district, Ostrovicë
Mts, Backë, brook and spring NE of the village, N40°31.346’ E20°25.096’, 1650
m, 12.10.2012, leg. P. Juhász, T. Kovács, D. Murányi, G. Puskás (1², 4³, OPC).”
Misidentification.
Drusus arbanios Oláh, 1910: Oláh & Kovács (2013: 112): “Albania,
Korçë district, Ostrovicë Mts, Çemerricë, open brook W (above) the village,
N40°32’38.9”, E20°26’33.5”, 1820 m, 28.V.2013, leg. P. Juhász, T. Kovács, G.
Magos, G. Puskás, (2², 7³, OPC). Albania, Skrapar district, Ostrovicë Mts,
Backë, Krojmbret Spring and its outlet brook NE of the village, N40°31.753’,
E20°25.152’, 1965 m, 28.V.2013, leg. P. Juhász, T. Kovács, G. Magos, G. Puskás,
(19², 7³, OPC). Albania, Skrapar district, Ostrovicë Mts, Backë, brook and
spring NE of the village, N40°31.346’, E20°25.096’, 1650 m, 29.V.2013, leg. P.
Juhász, T. Kovács, G. Magos, G. Puskás, (10², 13³, OPC). Albania, Skrapar
district, Ostrovicë Mts, open stream 3 km S of village Faqekuq, N40°32’19.3,
E20°24’22.0”, 1730 m, 29.V.2013, leg. P. Juhász, T. Kovács, G. Magos, G. Puskás
(3², 9³, OPC).” Misidentification.

152 J. Oláh
Drusus kovacsi Oláh, 2017 in Oláh et al. (2017: 98–100): “Holotype: male.
Albania: Skrapar district, Ostrovicë Mts, Backë, Krojmbret Spring and its out-
let brook NE of the village, N40°31.753’ E20°25.152’, 1965 m, 12.10.2012, leg. P.
Juhász, T. Kovács, D. Murányi, G. Puskás (3 males, 6 females; OPC). Allotype:
same as holotype (1 female, OPC). Paratypes: same as holotype (2 males, 6 fe-
males; OPC). Albania, Skrapar district, Ostrovice Mts, Ceremica, brook W of
the village, 1820m, N40°32.649’ E20°26.573’, 5. VII. 2005, leg. Z. Barina, D. Pifkó
& D. Schmidt (1 male, HNHM). Skrapar district, Ostrovicë Mts, Backë, brook
and spring NE of the village, N40°31.346’ E20°25.096’, 1650 m, 12.10.2012, leg.
P. Juhász, T. Kovács, D. Murányi, G. Puskás (1 male, 4 females, OPC). Albania,
Korçë district, Ostrovicë Mts, Çemerricë, open brook W (above) the vil-
lage, N40°32’38.9”, E20°26’33.5”, 1820 m, 28.V.2013, leg. P. Juhász, T. Kovács,
G. Magos, G. Puskás, (2 males, 7 females; OPC). Albania, Skrapar district,
Ostrovicë Mts, Backë, Krojmbret Spring and its outlet brook NE of the village,
N40°31.753’, E20°25.152’, 1965 m, 28.V.2013, leg. P. Juhász, T. Kovács, G. Magos,
G. Puskás, (19 males, 7 females; OPC). Albania, Skrapar district, Ostrovicë Mts,
Backë, brook and spring NE of the village, N40°31.346’, E20°25.096’, 1650 m,
29.V.2013, leg. P. Juhász, T. Kovács, G. Magos, G. Puskás, (10 males, 13 females;
OPC). Albania, Skrapar district, Ostrovicë Mts, open stream 3 km S of village
Faqekuq, N40°32’19.3, E20°24’22.0”, 1730 m, 29.V.2013, leg. P. Juhász, T. Kovács,
G. Magos, G. Puskás, (3 males, 9 females; OPC)”.
Diagnosis – Oláh et al. (2017: 98–100): “Drusus kovacsi is most close to D.
arbanios, but differs by having cerci with rounded dorsal lobe, not flat; lateral
profile of paraproct with convex vertical dorsoapical margin, not straight vertical
as well as dorsoapical tips, the diverging tips is rounded lobed, not truncate. The
erected primary spine differently shaped, however it is not known how stable is
this divergion having only the single holotype of D. arbanios available.”
Etymology – “We named this species after the collector Tibor Kovács, who
has performed a systematic and specialised collecting program to sample target
populations of Drusinae subfamily in the sky islands of high altitude crenon and
hypocrenon habitats in the Balkan mountan ranges during the years of 2010 and
2017.”
Drusus krpachi Kučinić, Graf et Vitecek, 2015

Drusus krpachi Kučinić, Graf et Vitecek, 2015 in Vitecek et al. (2015b: 81–
83): “Holotype. 1 male. Macedonia, Mavrovo National Park, Korab Mountains,
cesma Elem; N41.857, E20.625; leg. Kucinic, Krpac, Mihoci; 15. VIII. 2011.
Currently deposited in coll. WG, will be deposited in the Croatian Natural
History Museum, Zagreb, Croatia. Paratypes. 3 males: Macedonia, Mavrovo

National Park, Korab Mountains, Rec; leg. Krpac, Mihoci, Kucinic; 1. VIII.
2011. Currently deposited in coll. MK, two paratypes will be deposited in the
Macedonian Museum of Natural History, Skopje, Republic of Macedonia, one
paratype will be deposited in coll. WG.”
Diagnosis – Vitecek et al. (2015b: 81–83): “Males of the new species are
most similar to D. macedonicus.”
Drusus krusniki Malicky, 1981

Drusus krusniki Malicky, 1981a: 342–343: “Holotypus ²: Crna Gora,
Snjili Potok, Andrijevica, 25. V. 1979, leg. Sivec, coll. Malicky.”
Drusus kerek Oláh, 2011: 116: “Albania: Prokletije Mts, creek above village,
Sylbice, N42°31.326’, E20°05.919’ 2010 m, 8.07.2011, leg. Z. Barina, A. Kovács, G.
Puskás B. Sárospataki (2 females, HNHM). Prokletije Mts, valley of stream Topoje
below village Sylbice, tall herb community of the creek, N42°30.283’, E20°08.917’
1460 m, 8.07.2011, leg. Z. Barina, A. Kovács, G. Puskás, B. Sárospataki (3 males,
2 females, HNHM). Prokletije Mts, valley of stream Topoje, small side creek in
village Sylbice N42°30.557’, E20°08.054’ 1580 m, 8.07.2011, leg. Z. Barina, A.
Kovács, G. Puskás, B. Sárospataki (1 female, HNHM).” Misidentification.
Diagnosis – Malicky (1981a: 342–343): “Diese Art gehört in die Gruppe
der miteiander nahverwandten und durchwegs allopatrischen Arten um Drusus
bosnicus Klapálek, 1899, über die Marinković-Gospodnetić (1976) ausführ-
lich berichtet hat. Die Unterschide zwischen diesen Arten sind relativ gering und
liegen hauptsachlich in der Form der mittleren Anhänge.” Described as an al-
lopatric species of the Drusus bosnicus species group, and closest to Drusus kla-
paleki Marinković-Gospodnetić, 1971.”
Notes – Oláh et al. (2017: 111): “On Callumit Mountain we have detected a
contact zone where D. krusniki lives together with D. kerek. The two D. krusniki
males are smaller with significant modifications in the paramere spine pattern.
The subterminal perpendicular enlarged spine less perpendicular, almost hori-
zontal in one male.”
Drusus lakmos Oláh, 2017

Drusus graecus McLachlan, 1876: Malicky (2005a: 107): all specimens col-
lected by Sivec in Pindos and Lakmos Mts are misidentifications.
Drusus graecus McLachlan, 1876: Oláh & Kovács (2015: 109): misiden-
tification.
Drusus lakmos Oláh, 2017 in Oláh et al. (2017: 114–116): “Holotype:
Greece, Thessaly, Trikala peripheral unit, Lakmos Mts, Chaliki, springs on Verliga
Plateau, N39°40.825’, E21°07.551’, 2020 m, 09.05.2014, T. Kovács, D. Murányi (1

154 J. Oláh
male, OPC). Allotype: same as holotype (1 female, OPC). Paratypes: same as hol-
otype (2 males, 1 female, OPC). Greece, Thessaly, Trikala peripheral unit, Lakmos
Mts, Chaliki, open brook W of the village, N39°40.895’, E21°08.261’, 1840 m,
09.05.2014, T. Kovács, D. Murányi (1 male, 1 female, OPC). Thessaly, Trikala pe-
ripheral unit, Lakmos Mts, Chaliki, open stream SW of the village, N39°40.267’,
E21°09.176’, 1430 m, 09.05.2014, T. Kovács, D. Murányi (1 male, 2 females; OPC).”
Diagnosis – Oláh et al. (2017: 114–116): “The dorsal branch-es of the para-
proct fused forming simple, rounded hump-like, blunt apical arm in lateral view
at all the examined five specimens; not hump, more pointed at D. graecus. The
paraproct caudal profile is low, high at D. graecus; the apex of the fused dorsal
branches rounded with small pointed mesal structure, apex is straigt or rather
concave without any mesal structure at D. graecus. Cerci are rounded at all the
five specimens; elongated at D. graecus. The primary erect spine on the paramere
is highly reduced, almost vestigial at all the five males; this erect spine an an-
cestral character of the Drusus bosnicus species group is retained, almost fully
developed at D. graecus.”
Drusus lapos Oláh, 2017

Drusus lapos Oláh, 2017 in Oláh et al. (2017: 136–138): “Holotype: Italy,
Trento, Telve, torr. Maso T.L. c/o Malga Cagnon di Sotto, 1670 m, 8. VIII.
2001, leg. B. Lodovici (1 male, MCSNBG). Allotype: Italy, Bolzano, Moso in
Passiria, Ponte segheria, 1600 m, 19.VI.1993, leg M. Valle (1 female, MCSNBG).
Paratypes: Italy, Friuli Venazia Giulia, Tarvisio (UD), 870 m, Rio del Lago, light
trap, 21.VII.1996, leg Pantini & Valle (1 male, OPC). Austria, Weinebene, 46°
50.52 15° 00.33, 28.VII.2007. leg. W. Graf (1 male, OPC). Pinned specimen: 1st
label: Admont. 19.VIII.02; 2nd label: chrysotus Klapálek; 3rd label: chrysotus det
Kempny; 4th label: Drusus chrysotus ² det. Malicky 1983 (1 male, WM). Pinned
specimen: 1st label: Carinthia, Glockner, 7.VIII.20; 2nd label: Drusus chrysotus
² det. Dr. Döhler 1931; 3rd label: Drusus chrysotus ² det. Malicky 1983 (1 male,
WM). Pinned specimen: 1st label: Admont, 19.VIII.02; 2nd label: chrysotus det.
Kempny; 3rd label: Drusus chrysotus ²det. Malicky 1983 (1 male, WM). Pinned
specimen: 1st label: Stelzing, Jul.01; 2nd label: Coll Brauer, 3rd label: discolor ³
det. Brauer; 4th label: H. flavipennis; 5th label: Drusus chrysotus ²det. Malicky
1983 (1 male, WM). Pinned specimen: 1st label: Gesaues, subalpin H. Franz; 2nd
label: Coll. H. Franz; 3rd label: Drusus chrysotus Ramb. ² det. Dr. Döhler 1948;
Drusus chrysotus ² det. Malicky 1983 (1 male, WM). Pinned specimen: 1st label:
Strechengraben, Nied. Tauern, leg. H. Franz; 2nd label: coll. Franz; 3rd label:
Drusus chrysotus Ramb. ² det. Dr. Döhler 1952; 4th label: Drusus chrysotus ²

det. Malicky 1983 (1 male, WM). Pinned specimen: 1st label: Manu[?] Glockn.
Carin, M 856; 2nd label: flavipennis det Brauer; 3rd label: Drusus chrysotus ² det.
Malicky 1983 (1 male, WM). Pinned specimen: 1st label: Twong 12–1400 m;
2nd label: Salzburg, Radstadter Tauern, 3–10.VIII.’40, Zerny; 3rd label: Drusus
chrysotus Ramb. ³ det. Dr. Döhler 1941; 4th label: Drusus chrysotus det. Malicky
1983 (1 female, WM). Pinned specimen: 1st label: Austria Alp; 2nd label: Ullr.;
3rd label: flavipennis det Brauer; 4th label: Drusus chrysotus ² det. Malicky 1984
(1 male, WM). Pinned specimen: 1st label: Manu, Glockner, carin 856; 2nd label:
flavipennis det Brauer; 3rd label: Drusus chrysotus ² det. Malicky 1984 (1 male,
WM). Pinned specimen: 1st label: Austria Alp. 2nd label: flavipennis det Brauer;
3rd label: Drusus chrysotus ² det. Malicky 1984 (1 male, WM). Pinned speci-
men: 1st label: Gug 57. 2nd label: Drusus chrysotus Rbr. (1); 3rd label: Drusus
chrysotus ² det. Malicky 1986 (1 male, WM). Pinned specimen: 1st label: Austria
Alp; 2nd label: flavipennis det Brauer; 3rd label: Drusus chrysotus ² det. Malicky
1984 (1 male, WM). Pinned specimen: 1st label: Manu, Glocker 1861; 2nd label:
flavipennis det Brauer; 3rd label: Drusus chrysotus ² det. Malicky 1984 (1 male,
WM). Pinned specimen: 1st label: Triafoi, Handlirsch; 2nd label: Drusus chry-
sotus Ramb. ² det. Dr. Döhler 1931; 3rd label: Drusus chrysotus ² det. Malicky
1983 (1 male, WM). Pinned specimen: 1st label: Giglachseegeb. Schlaum Tauern,
leg. H. C. Franz; 2nd label: Drusus chrysotus Ramb. ² det. Dr. Döhler 1958; 3rd
label: Drusus chrysotus ² det. Malicky 1983 (1 male, WM). Czech Republic, S.
Bohemia, Sumava Mts. tributary of Cerne jezero lake, 15.V.2007 leg. P. Chvojka
(1 male, 2 females, NMPC). Switzerland, Valais Canton, Gd St Bernard, 2472 m,
7.VIII.1981, leg. C. Siegenthaler (1 male, CMZL). Switzerland, Valais Canton,
Gd St Bernard, Maringo, 1950 m, 14.VII.2004, leg. P. Stucki (1 male, CMZL).”
Diagnosis – Oláh et al. (2017: 136–138): “The fused dorsal branches of
paraproct robust with almost straith and vertical apical margin in lateral view.
Similar to D. chrysotus, but the dorsum of the fused dor-sal branches are flat hori-
zontal in lateral view, not sloping anterad. Periphallic organs of cerci and gonop-
ods are short. Subapical spine on the para-mere short and intact, not subdivided
like at D. chrysotus.”
Etymology – “Lapos”, from „lapos”, flat in Hungarian, refers to flat and hori-
sontal dorsum of the fused dorsal branches of the paraproct in lateral view.”
Drusus lepcos Oláh, 2011

Drusus lepcos Oláh, 2011: 116–118: “Holotype male. Albania: Mirdite
district, Shent Mts, Kurbnesh, Urake River and its sidespring NE of the city,
N41°47.711’, E20°06.703’, 800 m, 20.05.2010, leg. Z. Fehér, D. Murányi, Zs. Ujvári
(1 male, HNHM). Paratypes: same as holotype (1 male, 2 females, HNHM).”

156 J. Oláh
Diagnosis – Oláh (2011: 116–118): “This dark brown brow species belongs
to the species complex with large upward arching triangular gonopods inhabit-
ing the Balkan Mountains. Most close to Drusus dacothracus Oláh and D. illyricus
Oláh, but differs from both by having dorsum of paraproct stepwise formed in
lateral view. However these species are very close, moreover the lateral shape of
their cerci varying. Most easy to distinguish between the 3 species is to stretch
out the paraproct of the cleared genitalia out under the dark spinose tergite VIII
and compare the paraproctal dorsum in lateral view: flat sloping at D. dacothra-
cus, towering at D. illyricus and stepwise in D. lepcos sp. n.”
Etymology – “Lepcos, from “lépcsős”, stepwise in Hungarian, refers to the
shape of the dorsum of the paraproct in lateral view.”
Drusus macedonicus Schmid, 1956

Drusus macedonicus Schmid, 1956: 90–91: “Holotype ², allotype ³ et
paratypes ²²³³: Massif du Périster (Macédoine yougoslave), 10–12.VIII.1955.
L’espèce était assez commune le long des petits torrents d’alpage entre 1.500 et
2.000 m d’altitude.”
Diagnosis – Schmid (1956: 90–91): “Le Drusus de Macédoine est très voisin
de discolor comme en témoignent la coloration et les génitalia. Il s’en distingue
par sa coloration moins grise, par les appendices supérieurs du ² plus grand, plus
proéminents et rappelant ceux de transylvanicus et par les appendices inférieurs
pus élancés. Il est toujours bien distinct de discolor avec qui il cohabite.”
Drusus malickyorum Oláh, 2017

Drusus malickyi Oláh et Vitecek, 2015 in Vitecek et al. (2015b: 83–85):
“Material examined. Holotype. 1 male: Albania, Shkoder County, Shkoder
District, Prokletije Mts, beech forest with brook above Okol; N42.42258,
E19.76127; leg. Puskás 05. IX. 2013. Currently deposited in coll. WG, will be
deposited in János Oláh Private Collection under national protection of the
Hungarian Natural History Museum, Budapest, Hungary (JO).” Junior homo-
nym of Drusus malickyi (Sipahiler, 1992).
Drusus malickyorum Oláh, 2017 in Oláh et al. (2017: 145): new name for
Drusus malickyi Oláh et Vitecek, 2015.
Diagnosis – Vitecek et al. (2015b: 83): “The holotype of the new species is
most similar to D. macedonicus, but exhibits (1) a sharp mediocaudal protrusion of
segment IX; (2) a dorsally straight and rough tip of the intermediate appendage
distinctly separated by a proximal intendation; (3) a distinctly slender and con-
stricted distal half of the inferior appendage in lateral view. Drusus macedonicus
males have a mediocaudal and a ventrocaudal protrusion of segment IX, interme-

diate appendages with two rough rounded dorsal protrusions but lacking a dis-
tinct proximal indentation, and to a lesser degree constricted inferior appendages.”
Etymology – “We dedicate the new name to H. Malicky, to his wife and
their son.”
Drusus medianus Marinković-Gospodnetić, 1976

Drusus medianus Marinković-Gospodnetić, 1976: 80: “Localities in
the springs of the left tributaries of the river Bosna are: – Lasva (Plave vode),
30² 12³, 5.VI.1965; 8² 4³, 4.V.1973; Holotype ², allotype ³, paratypes ²²³³ in
author’s collection. – Lasva (Komar), 12² 4³, 13.V.1971; 6², 4.V.1972; 4² 3³,
15.V.1973. – Zujevina, 5² 2³, 10.V.1970. – Zujevina (Ljubovcica potok), 26²
2³, 14.V.1957; 5² 1³, 15.V.1958; 2² 23.IV.1958; 12² 3³, 15.V.1960. – Zujevina
(Krupa) 2² 28.VI.1957. – Fojnica (Pozarna) 22² 10³, 31.V.1967. Localities in the
springs of the right tributaries of the river Vrbas are: – Bistra (near Gornji Vakuf ),
1², 15.VI.1973. Krusnica (tributary of the river Bistrica), 1², 18.VI.1972.”
Diagnosis – Marinković-Gospodnetić (1976: 80): “Appendices interme-
diales differ from those of other Bosnian species of Drusus gr. bosnicus in being
small. They bear two narrow tips on the dorsal part. In dorsal view, appendices
intermediales are narrow and with a deep recess laterally. Appendices superiorers
are concave and similar to those of D. radovanovici septentrionis. The surface beset
with tubercules is also similar to that of D. radovanovic septentrionis, but its darker
semicircular zones are more separated by the light zone beset with few tubercules.”
Drusus meridionalis Kumanski, 1973

Drusus romanicus Murgoci et Botoşăneanu, 1954: Szczęsny (1970: 775):
Rila Mts, Malovitsa stream, 9.VIII.1969, 2 males; 12.VIII.1969, 4 females. Noted
numerous differences in genital structure between the Rila and the Romanian
specimens. Misidentification.
Drusus romanicus meridionalis Kumanski, 1973a: 108–110: “Die Material
der neuen Unterart is aus Pirin-Gebirge, Banderischki-Zirkus, Bergbach unter-
halb der Muratowi-Seen (2200 m Höhe), 21.IX.1967, 1² und 3³ und aus der
Umgebung der Berghütte “Demjanitza” (1900 m Höhe), 1.VIII.1970, 1², wie
auch aus dem Rila-Gebirge, Bergbach in dem Malöwitza-Zirkus, 9–12.VIII.1969,
2² (leg. M. Kownacka) und oberhalb der Hütte “Sawratschitza” (etwa 2200 m
Höhe), 10.VIII.1968, 2² (leg. Dr. I. Buresh).” Misidentification in part. Specimens
from the the Pirin Mts see Drusus pirinensis Oláh et Chvojka, 2017.
Drusus romanicus meridionalis Kumanski, 1973: Kumanski & Malicky
(1976: 105): Rila Mts, Nebenbach des Beli Iskar ob Borowez, 2300 m, 23–24.
VIII.1971, 1 male, 2 females. Rila Mts. Nebenbach des Beli Iskar ob Borowez,

158 J. Oláh
1200–1800 m, 24.VIII.1971, 1 female. Rila Mts. Nebenbach des Beli Iskar ob

Borowez, 2200–2300 m, 18–21.VII.1968, 3 males.
Drusus romanicus meridionalis Kumanski, 1973: Kumanski (1988: 42–44):
Rila Mts and Pirin Mts are given for distribution. No exact collecting data are
given, however the lateral profile of the paraproct as well as the paramere pattern
on the drawings are identical with specimens from the Rila Mts.
Drusus meridionalis Kumanski, 1973: Vitecek et al. (2015a: 258): raised to
species level by performance of morphological data in phylogenetic reconstruc-
tion of Drusinae.
Drusus meridionalis Kumanski, 1973: Vitecek et al. (2015c: 90–93): re-
drawn and redescribed from specimens collected in Pirin Mts. Misidentification.
Diagnosis – Kumanski (1973a: 108–110): “App. superiores bei den ² be-
merkenwert kürzer als bei der Nominatform (Drusus romanicus romanicus). App.
intermediales im Profil mit schwacher individualisiertem oberen (freien) Teil;
derselbe erhebt sich bei romanicus romanicus in seinem Vorderende unter einem
geraden Winkel.”
Drusus muranyorum Oláh, 2010

Drusus muranyorum Oláh, 2010: 104–106: “Holotype, male, HNHM.
Greece: Rodopi prefecture, Sapka Mts, Nea Sanda, torrent in an oak forest, E of
the village, 651 m, N41°07.672’ E25°53.223’, 4.IV.2007, leg. L. Dányi, Z. Erőss, Z.
Fehér, J. Kontschán & D. Murányi. Paratypes. Greece: Rodopi prefecture,, Sapka
Mts, Nea Sanda, torrent in an oak forest, E of the village, 651 m, N41°07.672’
E25°53.223’, 4.IV.2007, leg. L. Dányi, Z. Erőss, Z. Fehér, J. Kontschán & D.
Murányi (5 females).”
Diagnosis – Oláh (2010: 104–106): “This autumn collected dark fuscous spe-
cies belongs to the species complex of D. discophorus described from the Balkan.
Most close to Drusus bureschi Kumanski, 1973, but differs by having (1) almost
black body color, not light brownish yellow; (2) IXth segment long, not short in
lateral view; (3) lateral lobe on paraproctal complex in dorsal and caudal view nar-
row, not broad; (4) cerci subrectangular, not rounded; (5) gonopods S-shaped,
not upward arching triangular; (6) parameres single filament, not bifid.”
Etymology – “The name was given for the collectors Dávid Murányi and his
wife Szilvia Czigány, who have made together extensive collection activity along
alpine springs and streams on the Balkan.”
Drusus noricus Malicky, 1981

Drusus noricus Malicky, 1981b: 44: “Holotype male: Austria, Carinthia,
Saualpe 1900 m, Ladinger Alm, 12.VIII.1980, leg. et coll. Malicky.”

Diagnosis – Malicky (1981b: 44): “General appearance similar to Drusus

chrysotus Rambur, but smaller and paler coloration.” “The intermediate append-
ages are fused on their base only, but in chrysotus over their whole inner surface.
The upper edge of the intermediate appendages decreases caudally in D. noricus,
but increases in D. chrysotus.”
Drusus osogovicus Kumanski, 1980

Drusus osogovicus Kumanski, 1980: 204–205: “Holotype ² and 4 Paratypes
(2²² and 2³³): Osogovska Mt. (SW Bulgaria), hostel “Osogovo”, 1640 m a.s.l.,
18–19.VI.1979 (leg J. Ganev, at light), in the National Natural History Museum,
Sofia.”
Diagnosis – Kumanski (1980: 204–205): “Dr. osogovicus n. sp. belongs to
the group of discophorus, being in general one of the dark coloured species there.
The contrast between the yellow thoracic sclerites scutum and scutellum on one
hand and the darker rest of the insect on the other is one of its characteristics;
thus, the darkest species in the group, Dr. discophoroides Kum., as well as the dark
coloured population of Dr. discophorus Rad. found in the Rhodopes are monoto-
nous. The extremely narrow spinulate zone of 8th tergite and the turned inwards
position of the superior appendages are the main diagnostic features of the male.
As to the female, its genitalia are very similar to those of Dr. balcanicus Kum. and
the coloration remains its most distinctive feature.”
Drusus ostot Oláh, 2017

Drusus ostot Oláh, 2017 in Oláh et al. (2017: 101–102): “Holotype fe-
male. Albania: Prokletije Mts. Creek above village, Sylbice, 2010 m, N42°31.326’
E20°05.919’ 8. VII. 2011, leg. Z. Barina, A. Kovács, G. Puskás & B. Sárospataki (1
female HNHM). Paratype. Same as holotype (1 female, OPC).”
Diagnosis – Oláh et al. (2017: 101–102): “The very specialised tergit IX
makes it possible to distinguish this species from all the other known species.
Tergite of segment IX clearly bipartite well visible from both dorsal and lateral
view; on the middle of the tergite there is a traversal rim separating the basal
and distal parts of the segment; the lateral setose lobe of sternite IX rounded.
[…] The dorsal profile of the bipartite segment IX has some resemblence both
to D. klapaleki and D. medianus, but the the mesal excision on the apical mar-
gin of segment IX is deeper than at D. klapaleki and wider than at D. medianus.
Morover, the lateral profile of the segment IX differs very much from the other
two species.”
Etymology – “Ostot, from „osztott”, divided in Hungarian, refers to the shape
of segment IX divided by pronounced rim into basal and distal parts.”

160 J. Oláh
Drusus pallidus Kumanski, 1988

Drusus rectus McLachlan, 1868: Klapálek (1913: 16): “Jezero pod Musallou,
31. VII. 2², neobyčejné malé kusy, majici v rozpeti 16 mm.” P. Chvojka’s transla-
tion: “Lake below Musala Mt., remarkably small specimens, wing-span 16 mm).”
Musala is in the alpine zone of the Rila Mts.” Misidentification.
Drusus rectus rectus McLachlan, 1868: Schmid (1956: 61): established sub-
species D. rectus rectus and D. rectus nigrorectus and considered Klapálek’s record
from Bulgaria as probably a mistake.
Drusus discophorus Radovanović, 1942: Botoşăneanu & Sykora (1963:
22–24): revised and drawn Klapálek’s specimen from Rila Mts and compared
with Schmid’s drawings of Drusus discophorus. Misidentification.
Drusus discophorus Kumanski, 1979b: 68: misidentification.
Drusus discophorus pallidus Kumanski, 1989: 19–20: “Holotype chosen
among males of a large sample (20² and 5³) from the Rila Mountains, the Lower
Elensko Lake (ca. 2300 m alt.), 31.VII.1965, leg. A. Popov. The holotype and a
large series of paratypes (altogether 50² and 25³) from 20 localities in the three
above mentioned mountains (Pirin, Rila, Vitosha) is kept in the collection of the
National Museum of Natural History, Sofia.” Misidentification in part. Specimens
from the Pirin Mts belong to Drusus tovises Oláh et Chvojka, 2017.
Drusus pallidus Kumanski, 1989: Oláh et al. (2017: 165): elevated to spe-
cies rank.
Diagnosis – Kumanski (1989: 19–20): “Rather small insects, very variable
in size; length of forewing (², ³) 5.5–9.0 mm. Coloration pale-yellow to yellow-
brownish, in general much lighter than in the other subspecies (D. discophorus
rhodopaeus).”
Drusus paros Oláh, 2017

Drusus paros Oláh, 2017 in Oláh et al. (2017: 102): “Holotype female.
Albania: North Albanian Alps (Prokletije Mts.), Cerem, 42°29’48”N 19°56’55”E,
1225 m, 29. VII. 2016, light leg. Z. Varga (1 female, OPC).”
Diagnosis – Oláh et al. (2017: 102): “The very specialised tergit IX makes
it possible to distinguish this species from all the other known species. Tergite
of segment IX clearly bipartite well visible from both dorsal and lateral view;
lateral broadening of the basal part separating the basal and distal parts of the
seg.ment; apical margin of the segment IX is very specific, quadrifid; the lateral
setose lobe of sternite IX rounded continuing ventrad by setaless slightly pig-
mented ventrum.”
Etymology – “Paros, from „páros”, paired in Hungarian, refers to the bifid
“paired” shape of the mesal lobe on the apical margin of segment IX.”

Drusus pelasgus Oláh, 2010

Drusus pelasgus Oláh, 2010: 106–108: “Holotype male. Albania: Dibër
district, Korab Mts, torrent and wet meadow NE of the Mt. Korab, 2300 m,
N41°48.143’, E20°33.285’, 27.VI.2007, leg. L. Dányi, Z. Erőss, Z. Fehér, A. Hunyadi
& D. Murányi. Paratype, male, HNHM. ALBANIA: Dibër district, Korab Mts,
torrent and wet meadow NE of the Mt Korab, 2300 m, N41°48.143’, E20°33.285’,
27.VI.2007, leg. L. Dányi, Z. Erőss, Z. Fehér, A. Hunyadi & D. Murányi.”
Diagnosis – Oláh (2010: 106–108): “This dark species with almost ebony
black sclerites belongs to the species complex with large upward arching triangu-
lar gonopods and triangular or bilobed cerci inhabiting the Balkan Mountains.
Most close to Drusus plicatus Radovanović, 1942, but differs by having (1) poster-
odorsal spinate area on VIIIth tergite guadrangular in dorsal view, not triangular;
(2) rounded mesal lobe of the spinate area in lateral view, not rectangular; (3)
cerci not deeply bilobed; (4) cerci with dorsal lobe broad and short, not slender
and long; (5) inner branch of paraproct stepped in lateral view, not rounded tri-
angular; (6) inner branch with quadrangular lateral lobe in dorsal view, not with
triangular; (7) outer branch robust and straight vertical, not thin and arching;
(8) outer branch met mesad forming a closed structure around anus, not open.”
Etymology – “The name was given to remind the origin of the Albanian peo-
ple inhabiting the type locality. Pelasgus was the ancestor of the Pelasgians, the
son of Zeus, the Ancient Greeks even used to believe that he was the first man.
In a wider sense Albanians are Illyrians and Illyrians are Pelasgians, as a result
the Albanian language explains the names of the ancient Greek gods, the Greek
mythology originates from the Illyrian-Pelasgian.”
Drusus pirinensis Oláh et Chvojka, 2017

Drusus romanicus Murgoci et Botoşăneanu, 1954: Kumanski (1969a: 177):
“Bulgaria, Pirin Mts., Banderiski Circus. Ottoka na Muratovi Ozera. 21.IX.1967.”
Misidentification.
Drusus romanicus meridionalis Kumanski, 1973a: 108–110: “Die Material
der neuen Unterart is aus Pirin-Gebirge, Banderischki-Zirkus, Bergbach unter-
halb der Muratowi-Seen (2200 m Höhe), 21.IX.1967, 1 ² und 3³ und aus der
Umgebung der Berghütte “Demjanitza” (1900 m Höhe), 1.VIII.1970, 1², wie
auch aus dem Rila-Gebirge, Bergbach in dem Malöwitza-Zirkus, 9–12.VIII.1969,
2² (leg. M. Kownacka) und oberhalb der Hütte “Sawratschitza” (etwa 2200 m
Höhe), 10.VIII.1968, 2² (leg. Dr. I. Buresh).” Misidentification in part. Only
specimens from the the Rila Mts are Drusus meridionalis Kumanski, 1973.
Drusus romanicus meridionalis Kumanski, 1973: Kumanski & Malicky
(1976: 105): “Pirin Mts. Banderiza-Einzugsgebiet mit Nebenbachen, 1000–2000

162 J. Oláh
m, 24.VII.1968, 1 male. Pirin Mts. Banderiza-Einzugsgebiet mit Nebenbachen,

2000–2300 m, 26.VII.1971, 1 male, 3 females.” Misidentification.
Drusus meridionalis Kumanski, 1973: Vitecek et al. (2015c: 91–93):
“Bulgaria, Vihren, Pirin Mountains, Okotovo-Banserishka, marshy spring;
N41.7389, E23.4462; 23.VIII.2013 leg. Keresztes, Török, Kolcsár; 10 males: in
coll WG.” Misidentification.
Drusus pirinensis Oláh et Chvojka, 2017 in Oláh et al. (2017: 152–153):
“Holotype: Bulgaria, Pirin Mts. 2100 m, above Vichren challet, 27.VIII.1979,
leg. Beron (1 male, NMPC). Allotype. Same as holotype (1 female, NMPC).
Paratypes: Bulgaria, Blagojevgrad province, Pirin Mts, between Ribno Ezero and
Gorno Ezero, left side brook of Vasilashki Potok, 41°44’26.3”, 23°26’51.7”, 2191
m, 12. IX. 2016, leg. P. Juhász, T. Kovács, G. Szilágyi (2 females, OPC).”
Diagnosis – Oláh et al. (2017: 152–153): “The hump on the apical margin
of the paraproct, that is the only recognised paraproct divergence between D.
chrysotus and D. romaicus complexes, is less pronounced. The lateral profile of the
fused dorsal branches of paraproct has the rounded flat top low, sloping anterad,
differing from the lateral configuration of the other three species.”
Drusus plicatus Radovanović, 1942

Drusus plicatus Radovanović, 1942: 186–190: “Fundort: Labuniste, nord-
westlich von Struga, am Fussa, des Jablanitza-Gebirges. 13–16.VII.1939.”
Diagnosis – Radovanović (1942: 186–190): Compared to Drusus bosnicus
and D. graecus among the known species in the Balkan Peninsula. “Von dieser
Art [Drusus graecus] sowie von Drusus bosnicus Klap. Unterscheiden sich die
Exemplare von der Jablanitza (Drusus discophorus) und von Labuniste (Drusus
plicatus) nach einigen augenscheinlichen charakteristischen Merkmalen, vorw-
iegend nach der Form der Genitalanhange und nact der Gestalt des 8. Tergits.”
Drusus popovi Kumanski, 1980

Drusus popovi Kumanski, 1980: 203–204: “Holotype ² and 2³³ Paratypes:
Western Stara Planina Mts., Petrohan pass, stream Burzija (ca. 1000 m a.s.l.), 25.
V. 1979, (leg. Popov), in the National Natural History Museum, Sofia.”
Diagnosis – Kumanski (1980: 203–204): “This is another member of the
discophorus-group, related closely to Dr. bureschi Kum. It can be recognised from
the latter by the different shape of vthe intermediate appendages and by the long,
bnearing a series of thorns parameres of the male, as well as by the dorsal shape of
female genitalia and the longer central pice of the subgenital plate. These features
also complete the differential diagnosis of Dr. popovi n. sp.”

Drusus puskasi Oláh et Ibrahimi, 2017

Drusus puskasi Oláh et Ibrahimi, 2017 in Oláh et al. (2017: 157): “Holotype:
Bosnia & Herzegovina: Republika Srpska, Foča, Sutjeska NP, Zelengora Mts, S
of village Govza, brooks and outlets of Bijelo jezero, 1420 m, N43.380° E18.584°,
netting, leg. G. Puskás & G.Szövényi 8.VIII.2014 (1 male, OPC). Paratypes: same
as holotype (1 male, DBFMNSUP; 3 males, OPC).”
Diagnosis – Oláh et al. (2017: 157): “The lateral profile of the fused dorsal
branches of paraproct has short, stout and slightly anterad turning dorso-apical
digitiform process, most similar to the name bearing species D. siveci, but differs
by having small hump on the apical margin of the paraproct; by the anterad turn-
ing and tapering apex of the fused digitiform dorsoapical process; by the very tip
that is hardly bilobed just the mesal suture is discernible. The subapical spine of
the paramere is short robust, not long and bearing additional variously shaped
small spine-like for-mation on middle dorsum. The periphallic organ of gonopod
is elongated compared to D. siveci.”
Etymology – “We have dedicated this species to the collector Gellért Puskás,
who collects adult and juvenile Orthoptera on dry highlands, but devoted to visit
crenon area to collect caddisfl ies.”
Drusus radovanovici Marinković-Gospodnetić, 1971

Drusus radovanovici Marinković-Gospodnetić, 1971a: 80: ”Many
males and females in small springs of the tributaries of the river Sutjeska, on the
mountain Zelengora, up to 1400 m.”
Drusus radovanovici Marinković-Gospodnetić, 1971: Marinković-Gos-
podnetić, (1971b: 144): “Southeast Bosnia, ²² ³³ in small springs on the
mountain Zelengora.”
Drusus radovanovici radovanovici Marinković-Gospodnetić, 1971: Marin-
ković-Gospodnetić (1976: 78): Subspecies status created by the description
of Drusus radovanovici septentrionis Marinković-Gospodnetić, 1976.
Drusus radovanovici Marinković-Gospodnetić, 1971: Kučinić et al. (2011:
150): taxonomic status raised to species level based on larval and adult morphol-
ogy as well as by mtCOI gene sequences.
Diagnosis – Marinković-Gospodnetić (1971a: 80): “This species be-
longs to the group bosnicus too, but it is mostly similar to D. plicatus Rad. The
differences between D. radovanovici and D. plicatus is in the structure of the eight
tergit, in the shape of its front margin as well as in the shape of its zones with
tubercula. These characters of D. radovanovici correspond to these of bosnicus
and D. klapaleki.”

164 J. Oláh
Drusus ramae Marinković-Gospodnetić, 1971

Drusus ramae Marinković-Gospodnetić, 1971a: 80: “This species (mm
and ff ) is found only in the large karst springs of the river Rama.”
Drusus ramae Marinković-Gospodnetić, 1971: Marinković-Gospodne-
tić (1971b: 144): “Southeast Bosnia, ²² ³³ in karst spring of the river Rama.”
Diagnosis – Marinković-Gospodnetić (1971a: 80): “D. ramae is closely
related to D. radovanovici. It differs from it in the form of intermediate append-
ages. They are concave on the back side (lateral view). In dorsal view, they are
narrower, more pointed than by D. radovanovici. The differences appear in the
shape of superior appendages and inferior appendages, too.”
Drusus rhaeticus (Schmid, 1956)

Metanoea rhaetica Schmid, 1956: 69–70: “Elle habite les Alpes de la Suisse
orientale ou son aire de distribution prolonge sans discontinuité celle de flavi-
pennis. J’en ai étudié un grand nombre d’exemplaires de Suisse orientale.”
Drusus rhaeticus (Schmid, 1956): Oláh et al. (2017: 179): Metanoea syno-
nymised with Drusus.
Diagnosis – Schmid (1956: 69–70): “Cette espèce, très voisine de flavipen-
nis, a une aire repartition plus orientale que cette dernier. Cette espèce ne se dif-
férencie guère de flavipennis que par l’armature genitale; la coloration, la nervula-
tion et les proportions des différents articles des membres sont identiques chez
les deux espèces.”
Drusus rhodopeus Kumanski, 1989

Drusus discophorus Rad. ssp.?: Kumanski & Malicky (1976: 104):
“Die Tiere aus dem Rhodopen unterscheiden sich insofern leicht von der na-
menstypischen Form, als sie deutlich dunkler gefärbt sind. Da aber in den
Kopulationsarmaturen keine Unteschiede zu finden sind und auch die geogra-
phische Isolation von den anderen Populationen nicht sehr ausgeprägt ist, ver-
zichten wir darauf, dieser Form einen eigenen Namen zu geben. Es sei aber im-
merhin auf den Sachverhalt hingewiesen.”
Drusus discophorus rhodopeus Kumanski, 1989: 20: “Holotype ² and a
couple (² and ³) of paratypes (streamlets in the vicinity of the mountain hut
“Erkyupriya”, 1300–1700 m, 26. V. 1969, leg. D. Braasch) in the collection of
the National Museum of Natural History, Sofia; 1² paratype (same region, 1–2.
IX. 1971, leg. D. Braasch) and 1³ paratype (River Shirokolashka, upper stream,
1200–1600 m, 21.–31. VIII. 1971, leg D. Braasch) in coll. Malicky, Lunz am See,
Austria”.

Drusus rhodopeus Kumanski, 1989: Oláh et al. (2017: 166): elevated to spe-
cies rank.
Diagnosis – Kumanski (1989: 20): “Also small insects, but less variable than
D. d. pallidus ssp. n.; forewing length (²,³) 7.5–9.5 mm. General coloration rath-
er fuscous. Except for the smaller size, the new subspecies is morphologically less
different from the nominate form than D. d. pallidus. Its main distinctive feature
remains the geographic isolation of the population.”
Drusus romanicus Murgoci et Botoşăneanu, 1954

Drusus romanicus Murgoci et Botoşăneanu, 1954: 967–972: “Masivul
Retezat: circa 30 de exemplare (²² et ³³) în regiunea izvoarelor lacurilor Bucura,
Galešul, Stînišoara, Pietrele (Peste 2000 m altitudine) (leg. Botoşăneanu, 1950–
1952. Masivul Bucegi: 1 exemplar ² şi 1 exemplar ³ in Valea Căşăriei, în iunie
1951, la circa 900 m altitude.”
Drusus transylvanicus Schmid, 1956: 27–28: ”Holotype ²: Retyezat
(Transylvanie), déposé au musée de Budapest. Dr. transylvanicus est très voisin de
romanicus et surtout de chapmani; il s’en distingué principalement par la forme
de ses appendices intermediaires et superieurs.” Drusus transylvanicus has been
synonimized with Drusus romanicus by Botosaneanu (1967c: 300).
Diagnosis – Murgoci & Botoşăneanu (1954: 967–972): “În conclusie,
cele mai importante caractere permiţînd diferenţîierea speciei D. romanicus n.
sp., de toate celelalte descrise, sînt urmatoarele: ² – forma trapezoidală a ariei
spinigere pe tergitul abdominal VIII (observate dorsal); – aspectul apendicelor
preanale (privite lateral), – structura picioarelor genitale şi mai ales a regiunii lor
mediana şi ventrale; – forma titilatorilor. ³ – tubul anal egal în lungime cu apen-
dicele preanale; – forma pieselor laterale ale sternitului IX şi ale solzului vulvar.”
Drusus sarkos Oláh, 2017

Drusus sarkos Oláh, 2017 in Oláh et al. (2017: 138–139): “Holotype: Italy,
Friuli Venezia Giulia, Uccea (UD), 550 m, Uccea Rio Uccea, light trap, 24.V.1996,
leg. Pantini & C. Valle (1 male, MCSNBG)”.
Diagnosis – Oláh et al. (2017: 138–139): “Similar to D. slovenicus, but the
dorsum of the fused dorsal branches has different profile in lateral view; the dor-
so-apical corner or angle is more produced upward. The divergences in the shape
of the fused dorsal branches is more pronounced in the two species in the dorsal
and caudal profile. Periphallic organs of cerci and gonopods are short. The para-
mere has short subapical spine accompanied by two small secondary spines an-
terad. The most pronounced divergence between D. sarkos sp. nov. and D. sloveni-

166 J. Oláh
cus is well visible in the shape of the spinulose area on the VIII tergit. The shape
is rounded oviform at D. slovenicus and regular quadrangular at D. sarkos sp. nov.”
Etymology – “Sarkos”, from „sarkos”, corner in Hungarian, refers to the more
produced dorsoa-pical corner of the dorsal branches of the para-proct as well as
the to the quadrangular shape of the spinulose area on tergite VIII.
Drusus schmidi Botoşăneanu, 1960

Drusus schmidi Botoşăneanu, 1960a: 288–290: “6² et 1³ de Trescavica
et 3² et 1³ de Trnovo; le holotype ² et l’allotypes ³ ont été choisis parmis les ex-
emplaires de Trescavica (F. Schmid); 2 paratypes ²: Deutsches Entomologisches
Institut, Berlin; 4² et 1³ parat.: L. Botoşăneanu.”
Diagnosis – Botoşăneanu (1960a: 288–290): “En comparant minu-
tieusement les ² de D. tenellus qu’il avait capturé en Macédoine (Perister) et en
Bosnie (Trescavica et Trnovo), le Dr. Schmid constata qu’en réalité seulement les
premiers peuvent être considérés comme appartenant réellement à l’espèce de
Klapalek, en ce qui concerne les exemplaires bosniaques “...la zone de spinules du
VIIIe tergite est assez petite, indivise et située sur le ligne médiane. En comparant
les exemplaires bosniaques à celui de Macédoine et aux exemplaires de D. tenellus
provenant de Roumanie (Retezat et Banat) je suis moi-même arrivé à la conclu-
sion qu’il s’agit d’une nouvelle espèce, très proche de tenellus.”
Drusus septentrionis Marinković-Gospodnetić, 1976

Drusus radovanovici septentrionis Marinković-Gospodnetić, 1976: 78–
80: “Inhabits the north-western part of Bosnia. The localities are: – the source
of the river Pliva, 13² 2³, 8.VI.1972; 10² 4³, 4.V.1973. (Holotype ², allotype
³, paratypes ²²³³ are in the author’s collection, the Faculty of Natural Science
and Mathematics, Sarajevo.). – the source of the rivers in Livanjsko polje: river
Bistrica, 7² 2³, 8.VI.1966; 9² 1³, 19.X.1970; 1² 1³, 16.VII.1971; Sturba, 2² 2³,
8.VI.1966; 29² 19³, 17.X.1970; 4² 5³, 15.VII.1971. – the source on the left bank
of the river Vrbas near Bocac, 10² 2.VI.1966; 4² 4³, 8.VI.1972.”
Drusus septentrionis Marinković-Gospodnetić, 1976: Kucinic et al. (2011:
150): based on larval and adult morphology as well as by mtCOI gene sequences
taxonomic status raised to species level.
Drusus septentrionis Marinković-Gospodnetić, 1976: Kucinic et al. (2011:
150): taxonomic status raised to species level based on larval and adult morphol-
ogy as well as by mtCOI gene sequences.
Diagnosis – Marinković-Gospodnetić (1976: 78–80): “The difference
between D. radovanovici radovanovici and D. radovanovici septentrionis appears

in the form of appendices intermediales and appendices superior as well as in the

shape of tubercules zone of the 8th tergite.”
Drusus serbicus Marinković-Gospodnetić, 1971

Drusus serbicus Marinković-Gospodnetić, 1971c: 107–108: “Stream
which is an affluent of the river Glonsica (near Nova Varos, west Serbia), 13m,
1f, 30. V. 1970.”
Drusus sharrensis Ibrahimi, Vitecek et Previšić, 2016

Drusus sharrensis Ibrahimi, Vitecek et Previšić, 2016 in Ibrahimi et al.
(2016: 111–119): “Holotype. 1 male: Republic of Kosovo, Shterpce Municipality,
Sharr Mountains, tributary of the Lepenc River, 2 km above the main road
Prizren-Shterpce, 1558 m, 42.17228°N, 20.98823°E, 21.v.2014, leg. Halil Ibrahimi
(DBFMNSUP). Paratypes: same collection and locality data as holotype,
6 males, 3 females (DBFMNSUP), 2 males, 1 female (CNHM), 2 males, 1 fe-
male (BDOL); same except 8.v.2014, 2 males, 1 female (CNHM); same except
15.vi.2013, leg. Halil Ibrahimi and Joachim Milbradt, 3 males (DBFMNSUP);
Shtërpce Municipality, Sharr Mountains, small spring, a branch of the Lepenc
River 50 meters above the main road Prizren – Shtërpce, 1410 m, 42.17506°N,
20.97593°E, 08.vi.2010, leg. Halil Ibrahimi, 2 males (DBFMNSUP); Shtërpce
Municipality, Sharr Mountains, Lepenc River on the main road Prizren –
Shtërpce, 1465 m, 42.1813°N, 20.9781°E, 18.v.2010, leg. Halil Ibrahimi, 2 males
(DBFMNSUP); Prizren Municipality, Sharr Mountains, Lumbardhi i Prizrenit
River, Prevallë village 1664 m, 42.161°N, 20.99533°E, 08.vi.2009, leg. Halil
Ibrahimi, 1 male (DBFMNSUP); Prizren Municipality, Sharr Mountains, first
small lake above Prevallë village, 2142 m, 42.152402°N, 20.995024°E, 18.ix.2010,
leg. Halil Ibrahimi, 3 males, 1 female (DBFMNSUP).”
Diagnosis – Ibrahimi et al. (2016: 111–119): “Males of the new species are
most similar to Drusus krusniki, D. kerek and D. juliae. Females of the new species
are most similar to D. krusniki, D. kerek, D. juliae, and D. plicatus.”
Drusus siveci Malicky, 1981

Drusus siveci Malicky, 1981a: 343–344: “Holotypus ²: Crna Gora, Snjili
Potok, Andrijevica, 25. V. 1979, leg. Sivec, coll.Malicky.”
Diagnosis – Malicky (1981a: 343–344): “Diese Art gehört in die
Verwandtschaft von Drusus croaticus Marinkovic, 1971, Drusus macedonicus
Schmid, 1956, Drusus imrovisus McLachlan, 1884, Drusus biguttatus Pictet,
1834, und anderen.”

168 J. Oláh
Drusus slovenicus Urbanič, Krušnik et Malicky, 2002

Drusus slovenicus Urbanič, Krušnik et Malicky, 2002: 35: “Holotype
²: Slovenia,Triglav National Park, Bohinj, Studorski preval, 1740 m, 46°18’N,
13°54’E, 19.VII.1983, leg Sivec, in the collection of the first author.”
Diagnosis – Urbanič et al. (2002: 35): “It may be related with some en-
demic species of the Balkan peninsula such as D. croaticus Marinkovic 1971 or D.
siveci Malicky 1981. It is also somewhat similar to Drusus noricus Malicky 1981.”
Drusus tenellus (Klapálek, 1898)

Catadice tenella Klapálek, 1898a: 488–489: “Hungary, Vom Hochgebirge
Retyezát. 2², 1³”.
Catadice tenella Klapálek, 1898: Klapálek, 1899a: 432–434: “Retyezát.
2², 1³”.
Drusus tenellus (Klapálek, 1898): Mosely 1933: 499: Catadice synonymised
with Drusus.
Drusus tenellus (Klapálek, 1898): Botoşăneanu (1959: 71): “Retezat, 800
m altitudine 12 VIII. Valea Cernei: 1–2 VIII. Belareca spre vărsarea în Cerna
(de la Mehadia pînă la confluenţă): 31. VII. “Dintre toate speciile de Drusus din
materiale noastre, este cea care coboară cel mai jos (pînă la altitudini în jur de
150 m) şi care populează gama cea mai euritermă de ape (rîurile Belareca, Cerna,
probabil rîurile rezultate din unirea torenţilor Retezatului).”
Diagnosis – Klapálek (1898a: 488–489): “Körper rötlich gelbbraun,
mit gelbbraunen Harchen, denen aber auf dem Scheitel und neben den Augen
schwarze Härchen beigemischt sind. Die obere Anhänge schmal, länglich,
schwach gekrümmt, in der Ansicht von oben stumpf dreieckig und ausgehöhlt;
die mittleren Anhänge verwachsen; untere Anhänge stark, aber kurz, kegelför-
mig, daher in der Seiten- und Rückenansicht dreieckig, stumpf.”
Drusus tovises Oláh et Chvojka, 2017

Drusus tovises Oláh et Chvojka, 2017 in Oláh et al. (2017: 167): “Holotype:
Bulgaria, Blagojevgrad province, Pirin Mt, left side brook of Vasilashki Potok,
41°44’32.08”, 23°26’24.45”, 3.VII.2016, leg. K. Harmos, T. Kovács & G. Magos
(1 male, OPC). Allotype: same as holotype (1 female, OPC). Paratypes: same as
holotype (10 males, 1 femle; OPC). Bulgaria, Blagojevgrad province, Pirin Mt,
Bansko – Demianitsa hut, left side stream of Demianitsa Reka, 2.VII.2016, leg.
K. Harmos, T. Kovács & G. Magos (4 males, OPC). Bulgaria, Blagojevgrad prov-
ince, Pirin Mt, Bansko – Demianitsa hut, left side brook of Demianitsa Reka,
41°46’15.3”, 23°27’44.3”, 3.VII.2016, leg. K. Harmos, T. Kovács & G. Magos (3

males, OPC). Bulgaria, Pirin Mts., Bela Reka stream (1800+1900 m), 19. IX.
1990, leg. P. Chvojka (4 males, 4 females, NMPC; 2 males, 2 females; OPC).”
Diagnosis – Oláh et al. (2017: 167): “This new species is widely distributed
in the crenon and hypocrenon habitats of high elevations in the Pirin Mts. A
sibling species of D. pallidus and D. rhodopeus, but differs from both species by
the shape of paraproct; the apical arms of the dorsal branches of the paraproct is
slightly anterad angled, not rounded blunt of D. rhodopeus or simple hump of D.
pallidus in lateral view; in caudal view the apical arms are very low with concave
dorsum not very high of D. pallidus or high with straight dorsum of D. rhodopeus.”
Etymology – “Tovises”, from „tövises”, spiny in Hungarian, refers to the para-
mere with subapical spine bunch of many spines as well as to the paramere shaft
anterad of the subapical spine bunch covered with numerous tertiary spines.”
Drusus vargai Oláh, 2017

Drusus vargai Oláh, 2017 in Oláh et al. (2017: 187): “Holotype: Jugoslavia,
Durmitor, Zablják, 21-25. VII. 1965, leg. Z. Varga (1 male, OPC).”
Diagnosis – Oláh et al. (2017: 187): “This new species is close to Drusus sch-
midi, but differs by having the paraproct diverged, differently formed. In lateral
view the paraproct is slender, not robust; in dorsal view the slender apical arms
of the dorsal branches of the paraproct form a deepp V-shaped apical excision
that is lacking in D. schmidi; the lateral margins are smooth concave, not sup-
plied with lateral humps. In caudal view the ventral branch of paraproct is high
and narrow, not low and wide as well as the laterad directed arms are robust, not
tapering.”
Etymology – “We describe this new species and dedicate it to the collector,
Prof. Z. Varga.”
Drusus vekon Ibrahimi et Oláh, 2017

Drusus vekon Ibrahimi et Oláh, 2017 in Oláh et al. (2017: 158–159):
“Holotype: Kosovo: Çakor, streamlet along the border line between Kosovo and
Montenegro. 42.685542°N, 20.053636°E, 1289 m, 25.VIII.2015 leg. E. Dimitrou
& V. Dragobia (1 male, DBFMNSUP). Allotype: Lloqan, Gurrat e Hasan Agës
springs, Bjeshkët e Nemuna. 42.557155°N, 20.152696°E, 1991 m, 3.VIII.2016,
leg. H. Ibrahimi (1 female, DBFMNSUP). Paratypes: same as holotype (1 male,
1 female; DBFMNSUP; 1 male hybrid, OPC). Haxhaj, spring area of a tributary
of Lumbardhi i Pejës River, 42°42’30N 20°2’32E, 1278 m, 25.VIII.2015 leg. E.
Dimitrou & V. Dragobia (1 male, OPC). Lloqan, Gurrat e Hasan Agës springs,
Bjeshkët e Nemuna. 42.557155°N, 20.152696°E, 1991 m, 18.VIII.2015 leg. H.
Ibrahimi (2 males, DBFMNSUP, 1 male, OPC). Lloqan, Gurrat e Hasan Agës

170 J. Oláh
springs, Bjeshkët e Nemuna. 42.557155°N, 20.152696°E, 1991 m, 3.VIII.2016,

leg. H. Ibrahimi (62 males, OPC, DBFMNSUP). Lloqan, Krojet e Gucise,
42.55143°N, 20.1335°E, 2110 m, 3.VIII.2016, leg. H. Ibrahimi (12 males, OPC,
DBFMNSUP). Lloqan, Gurrat e Hasan Agës springs, Bjeshkët e Nemuna.
42.557155°N, 20.152696°E, 1991 m, 3.VIII.2016, leg. H. Ibrahimi (1 female,
OPC). Lloqan, Krojet e Gucise, 42.55143°N, 20.1335°E, 2110 m, 3.VIII.2016, leg.
H. Ibrahimi (1 female, DBFMNSUP).”
Diagnosis – Oláh et al. (2017: 158–159): “The lateral profile of the fused
dorsal branches of paraproct has long and slender dorsoapical digitiform proc-
ess, most similar to D. fortos sp. nov., but differs by the anterad turning slender,
long and tapering apex of the fused digitiform dorsoapical process, and the very
tip of the fused dorsal branches of the paraproct is not completely fused, the tip
is bilobed and the mesal suture is discernible that is the vestigium of the fusion
surface present and visible in caudal view, The subapical spine of the paramere
long and robust and frequently more complex, than the subapical spine of D. for-
tos. In the sampled habitats D. vekon sp. nov. lives together with D. fortos sp. nov.
with hybrid forms.”
Etymology – “Vekon”, from „vékony”, thin in Hungarian, refers to the slender
dorsoapical digiti-form process of the paraproct.”
Drusus vernonensis Malicky, 1989

Drusus veronensis Malicky, 1989: 303–304: “Holotypus ² und paratypen
(1², 1³): Griechenland, Vernon-Gebirge beit Bitsi, ca. 1700 m, 11.VI.1989, in
meiner Sammlung.”
Diagnosis – Malicky (1989: 303–304): “Kopulationsarmaturen ² ähnlich
wie bei Drusus Krušniki Malicky. Drusus Krušniki aus Crna Gora ist etwas gröss-
er, im Habitus und in der Färbung gleich, das hyaline Dorsalfeld des 8. Tergits ist
breiter, der Endteil der unteren Anhänge in Ventralansicht viel plumper, und die
Form der freien Dorsalteile der mittleren Anhänge ist anders.”
Drusus vespertinus Marinković-Gospodnetić, 1976

Drusus vespertinus Marinković-Gospodnetić, 1976: 80: “Has been
found only in a large karstic spring of the river Ribnik, a tributary of the river
Sana: 68² 5³, 26.III.1968 (Holotype ², allotype ³, paratypes ²²³³ are in the
author’s collection); 20² 11³, 25.V.1968; 12² 1³, 28.III.1975”
Diagnosis – Marinković-Gospodnetić (1976: 80): “The structure of ap-
pendices intermediale is complex. The shape of the dark part of appendices inter-
mediales is very similar to appendices intermediales of D. radovanovici septentri-

onis. Appendices superiores are concave and similar to those of D. radovanovici

septentrionis and D. medianus.”
Drusus zivici Kučinić, Previšić, Stojanović et Vitecek, 2017

Drusus zivici Kučinić, Previšić, Stojanović et Vitecek, 2017 in Vitecek et al.
(2017: 8–12): “Holotype. 1 male: Serbia, Stara Planina Mountains, spring of the
river Tovarnička Reka; N43.3362367 E22.583983; 1493 m a.s.l.; 19.vi.2013–21.
vi.2013; leg. M. Kučinić, K. Stojanović, M. Živić; specimen identifier f Dpo0202M.
Paratypes: 1 male, 1 female, 5 larvae: same data; specimen identifiers: 1 female:
f Dpo0201F, 1 male: f Dpo0203M, 5 larvae: f Dpo0204L–f Dpo0208L. 2 males:
Serbia, Stara Planina Mountains, spring of the river Rekička Reka; N43.372150
E22.625333; 1540 m a.s.l.; 19.vi.2013; leg. M. Kučinić, K. Stojanović, M. Živić;
specimen identifiers: f Dpo0301M, f Dpo0302M. 1 female, 3 larvae: Serbia, Stara
Planina Mountains, spring of Kaluđerske vode; N43.388690 E22.677934; 1930 m
a.s.l.; 20.vi.2013; leg. M. Kučinić, K. Stojanović, M. Živić; specimen identifiers:
1 female: f Dpo0401F, 3 larvae: f Dpo0402L–f Dpo0404L. 1 female: Serbia, Stara
Planina Mountains, spring of the river Javorska Reka; N43.386150 E22.689817;
1890 m a.s.l.; 20.vi.2013; leg. M. Kučinić, K. Stojanović, M. Živić; specimen iden-
tifier: f Dpo0501F.”
Diagnosis – Vitecek et al. (2017: 8–12): “Males of the new species are most
similar to Drusus popovi but exhibit (1) subcircular, elongate superior append-
ages in lateral view; (2) high tips of the intermediate appendages in lateral view,
high and wide tips of intermediate appendages in caudal view; (3) suboval, elon-
gate, approximately straight inferior appendages in lateral view; and (4) a high
segment IX with a distinct, rounded, caudad medial protrusion in lateral view.
Drusus popovi males have subcircular, short superior appendages; short and nar-
row tips of the intermediate appendages in lateral and caudal view; suboval, dor-
sadly curved inferior appendages; a wide segment IX lacking a distinct medial
indentation in lateral view. Females of the new species are most similar to females
of D. popovi but exhibit (1) in dorsal view distinct, rounded lateral shoulders of
segment X, (2) in dorsal view a ragged outline of the lateral lobes of segment X.
Drusus popovi females have an evenly rounded lateral outline of segment X in
dorsal view and evenly rounded lateral lobes of segment X. Larvae of the new
species are most similar to D. serbicus Marinković-Gospodnetić as larvae of both
species have an intermittent lateral line ([98, 99]), but exhibit a pronotum with a
distinct, rounded pronotal ridge (type B sensu [98]). Larvae of D. serbicus have an
annular pronotal ridge (type E sensu [98]).”
Etymology – “Named for Miroslav Živić, biophysicist, for his continuous
support of faunistic surveys.”

172 J. Oláh
Ecclisopteryx Kolenati, 1848

Ecclisopteryx asterix Malicky, 1979
Ecclisopteryx asterix Malicky, 1979: 4–5: “Holotype ² and two paratypes
²: Austria, Carinthia, northern slope of Loiblpass, 800 m, 18. VI. 1977, leg. &
coll. Malicky.”
Diagnosis – Malicky (1979: 5): “This species resembles Metanoea rhaetica
in colour and general appearance. In the genus Ecclisopteryx (Schmid, 1956) it is
somewhat isolated because segment 9 is closed ventrally, whereas in the other
three species interrupted.”
Ecclisopteryx alkon Oláh et Oláh, 2017

Ecclisopteryx alkon Oláh et Oláh, 2017 in Oláh et al. (2017: 199–202):
“Holotype: Romania, Lotru Mts, Obirsia Lotrului, dawn swarm along Lotru River,
30.VI.2016, leg. J. Oláh & J. Oláh jr. (1 male, OPC). Allotype: same as holotype (1
female, OPC). Paratypes: Bulgaria Western Rhodopi, Sumena Reka near crossroar
to Zmeitza, 1245m, N41°39’20” E24°13’44”, 16.VI.2012, at lamps, light traps leg.
S. Beshkov M. Beshkova & V. Gashtarov (1 male, 1 female, NMNHBAS). Central
Stara Planina Mts. Elenova Gora Reserve near Skobelevo Village, Mazalat Forestry,
872 m, N42°44’34” E025°08’50”, 1.VIII.2014, leg. S. Beshkov (1 male, 1 female;
OPC). Montenegro: Mojkovac municipality, Sinjajevina Mts, Gornja Polja, Zoljski
Ljevak Stream above the village, N42°57.808’, E19°31.597’, 880 m, 14.VI.2012, leg. Z.
Fehér, T. Kovács, D. Murányi (1 male, OPC). Romania, same as holotype (4 males,
3 females, OPC). Rodna Mts. Borşa, Staţiunea Borşa, garden, 878m N47º36’48.0”
E24º46’55.8”, 28–29.VI.2005, light trap J. Kontschán, D. Murányi, K. Orci (2 males,
NHMB). Maramaros Mts. tributaries at Visó spring area, 15.VI.1993 leg. J. Oláh (6
females, OPC). Apuseni Mts, Direction to Scărişoara, 30.V.2006, leg. M. Bálint (14
males, OPC). Apuseni Mts. Munţii Gilăului, Muntele Bâisorii, stream Valea Gera,
N46°33.001’ E23°20.014’, 1055m, 18.VI.2013, light leg. J. Oláh, Cs. Balogh, & S.
Fekete (1 male, 1 female; OPC). Apuşeni Mts, Bihor Mts, Crişul Pietros, Boga, Valea
Bulz and Valea Galbena, N: 46°35’ 23,25” E: 22°37’ 54,74”, 450m, 4.VII.2013, light
leg. Cs. Balogh (1 male, OPC). Apuşeni Mts, Bihor Mts, Bubeşti-Cobleş, tributary
P. Cobleş, N: 46°29’ 56.08” E: 22°43’ 48.64” 902 m, 14.V.2014, leg. Cs. Balogh &
B. V. Béres (1 male, OPC). Cupaş Mts, Cupaş Valley, Lacu Roşu, 14.VII.1981 leg.
Peregovits & Ronkay (4 males, 2 females, HNHM). Bucegi Mts, Cocora stream,
25.443993, 45.394969, 1564 m, 15.VII.2015, light trap, leg. Z. Baczó & J. Kecskés (14
males, 182 females, OPC). Bucegi Mts, Cocora stream, 25.443147, 45.3402125, 1680
m, 16.VII.2015, leg. Z. Baczó & J. Kecskés (1 male, 1 female; OPC). Vâlcea county,
Parâng Mts, Obrâşia Lotrului, open spring area, 500 m along Transalpina (67C) road,
downstream from 45°22’27.7”, 23°39’4.0”, 1915 m, 30.VI.2016, leg. J. Oláh & J. Oláh

jr. (1 female, OPC). Lotru Mts, Obirsia Lotrului, 1578 m, 45.463 23.620, 29.VI.2016,
singled leg. J. Oláh & J. Oláh jr. (1 female, OPC). Lotru Mts, Obirsia Lotrului, 1578
m, 45.463 23.620, 29.VI.2016, light trap leg. J. Oláh & J. Oláh jr. (5 females, OPC).
Lotru Mts, Obirsia Lotrului, 1578 m, 45.463 23.620, 30.VI.2016, light trap leg. J. Oláh
& J. Oláh jr. (3 females, OPC). Lotru Mts, Obirsia Lotrului, side stream of River
Lotru, 29.VI.2016, dawn swarm, leg. J. Oláh & J. Oláh jr. (5 males, 105 females; OPC).
Lotru Mts, Obirsia Lotrului, side stream of River Lotru, 30.VI.2016, light trap, leg.
J. Oláh & J. Oláh jr. (19 females; OPC). Retezat Mts., Cheile Butii, 936m, N: 45°18’
07,30” E: 22°58’ 27,92” 9.VII.2013, leg. E. Bajka, Cs. Balogh, G. Borics, P. Borics (3
males, 2 females, OPC). Retezat Mts., Cheile Butii, 910m, N: 45°18’ 06,96” E: 22°58’
31,48” 9.VII.2013, light leg. E. Bajka, Cs. Balogh, G. Borics, P. Borics (28 males, 41
females; OPC). Munţii Sibiului, Rau Sadu, 700 m N45.64 E24.06, 3.VII.2007, leg. M.
Bálint (1 male, OPC). Caras-Severin county, Tarcu Mts. Poiana Marului, upper sec-
tion of Sucu Stream, S of the village, 955m, N45°20.907’ E22°31.073’, 8.VI.2011, leg.
T. Kovács, D. Murányi & G. Puskás (16 females, HNHM).”
Diagnosis – Oláh et al. (2017: 199–202): “This new species is a close incipi-
ent sibling species of E. dalecarlica, but differs by having different lateral profile
of the gonopods; gonopod apex is monolobed, not bilobed; produced mono-lobe
on the ventral corner of the gonopod apex is a decisive character state of E. alkon
sp. nov.; E dalecarlica has bilobed gonopod apex, i.e. there is an additional, smaller
lobe on the dorsal corner of the gonopods; this different pattern of the lateral
profile of the gonopods is the result of the modified position of the heavily pegged
vertical ridge of the stimulatory organ; actually the small lobe of the dorsal corner
is present also on E. alkon sp. nov. but the vertical ridge moved or shifted higher
masking or decreasing the apparent lobeness of the dorsal corner of the gonop-
ods.” “The difference in the spine patterns between the two siblings is less pro-
nounced, but E. alkon sp. nov. has smaller subapical spine cluster and more heavily
developed subbasal scattered spine pattern, com-pared to E. dalecarlica.”
Etymology – “Alkon”, from „alkony”, nightfall in Hungarian, refers to the
mass swarming habit of this new species in the dusk of the nightfall. We have
experienced clouds of heavy swarmings at nighfall in Apuseni, Bucegi, Lotru and
Retezat mountains.”
Ecclisopteryx ivkae Previšić, Graf et Vitecek, 2014

Ecclisopteryx ivkae Previšić, Graf et Vitecek, 2014 in Previšić et al. (2014:
317–319): “Holotype ²: Cetina River, Glavaš spring, N 43.976697 E 16.430150,
386 m asl, 02.vi.2011, leg. Previšić A.; deposited in the Biology Centre,
Oberösterreichisches Landesmuseum, Linz, Austria. Paratypes: 4 ² and 2 ³, same
data; 1 ² and 1 ³ 31.V.2005, leg Previšić A.; 1 ³ 07.vi. 2007, leg Graf W.; 2 ³

174 J. Oláh
02.vi.2012, leg. Previšić A.; deposited in the first author’s collection at the Faculty
of Science in Zagreb. 8 5th instar larvae, same location, 04.x. 2013 (N=4, leg.
Kučinić M.) and 07.xi.2013 (N=4, leg. Previšić A.).”
Diagnosis – Previšić et al. (2014: 317–319): “Posterior edge of tip of infe-
rior appendages more or less straight in lateral view, lacking a clear ventral elon-
gation, tips in dorsal view with distinct shoulder.”
Ecclisopteryx keroveci Previšić, Graf et Vitecek, 2014

Ecclisopteryx keroveci Previšić, Graf et Vitecek, 2014 in Previšić et al.
(2014: 317–319): “Holotype ²: Bosnia and Herzegovina, mouth of Jabučica
River, N 43.29022 E 18.61733, 765 m asl, 04.vii.2012, leg. Previšić A., Ivković M.,
Mihaljević Z., Miliša M.; deposited in the Biology Centre, Oberösterreichisches
Landesmuseum, Linz, Austria. Paratypes: 30 ² and 49 ³, same data; deposited
in the first author’s collection at the Faculty of Science in Zagreb. 10 5th instar
larvae, same location, 14.v.2008 and 02.vi.2009, leg. Previšić A., Graf W.”
Diagnosis – Previšić et al. (2014: 317–319): “Posterior edge of tip of in-
ferior appendages ventrally elongated and arched dorsally, tips in dorsal view
lacking distinct shoulder.”
Ecclisopteryx loudai Oláh, 2017

Ecclisopteryx loudai Oláh, 2017 in Oláh et al. (2017: 204): “Holotype:
Greece, Metsovo, 1100 m, 39.77oN 21.18oE, 13. VII. 2012 leg. J. Louda (1 male,
MCSNBG). Allotype: same as holotype (1 female, MCSNBG). Paratypes: same
as holotype (3 females, MCSNBG; 1 male, 1 female, OPC).”
Diagnosis – Oláh et al. (2017: 204): “This new species is a close incipient
sibling species of E. alkon sp. nov. and E. dalecarlica but differs by having differ-
ent lateral profile of the gonopods; the smaller lobe on the dorsal corner of the
gonopods has completely reduced, that is disappeared at both the holotype and
at the single male paratype.”
Etymology – “The species was named after Josef Louda, Czech entomologist,
who has collected this new species.”
Limnephilinae
Limnephilini
Limnephilus Leach, 1815
Limnephilus petri Marinković-Gospodnetić, 1966
Limnephilus petri Marinković-Gospodnetić, 1966a: 112: “Holotype ²,
allotype ³, paratypes ²² ³³: Sar-Planina, August 1954.”

Chaetopterygini
Annitella Klapálek, 1907
Annitella apfelbecki (Klapálek, 1899)
Chaetopterygopsis apfelbecki Klapálek, 1899: 329–330: “Ledici kod
Sarajeva (Apfelb.).” Klapálek (1900: 676–677): “Ledici bei Sarajevo (Apfelb.).”
Annitella apfelbecki (Klapálek, 1899): Klapálek (1907: 29–30): Annitella
gen. n. erected with type species Annitella kosciuszkii. Chaetopterygopsis apfel-
becki Klapálek transferred to the new genus.
Annitella ostrovicensis Oláh et Kovács, 2012

Annitella ostrovicensis Oláh et Kovács, 2012: 92–93. “Holotype: Albania:
Skrapar district, Ostrovicë Mts, Backë, Krojmbret Spring and its outlet brook
NE of the village, N40°31.753’, E20°25.152’, 1965 m, 12.10.2012, leg. P. Juhász,
T. Kovács, D. Murányi, G. Puskás (1², OPC). Allotype: same as holotype (1³,
OPC). Paratypes: same as holotype (24², 10³; OPC; 4², 1³, MM).”
Diagnosis – Oláh & Kovács (2012: 92–93): “This spring brook dwell-
ing new species collected on high elevation is a sister species of Annitella triloba
Marinković-Gospodnetić, 1955 but differs in male by having tergite VIII with-
out median spinate lobe, paraproct without median process, cerci reduced to an
almost indiscernible pair of warts, bifid distal sclerite of aedeagus very narrow.
Also differs in female by having sternite IX (setosa lateral lobes) with very short
ventrum, dorsal black region of segment X simple rounded, not with ventral pair
of oblique rounded ridges. A. ostrovicensis sp. n., probably a parapatric or peripat-
ric species occurs not far from the southernmost populations of its sister species
A. triloba.”
Etymology – “The new species is named after the Ostrovicë Mts, where the
type locality is found. These mountains are rich in valuable autumnal Trichoptera
sapecies (even our one-day collecting demonstrates it).”
Annitella jablanicensis Oláh, 2014

Annitella jablanicensis Oláh, 2014 in Oláh & Kovács (2014: 110–112):
“Holotype. Macedonia: Southwestern region, Jablanica Mts, 6.5 km W of
Labuništa, open brook at Labuniško Lake, N41°16.069’, E20°31.242’, 1905 m,
10.10.2014, leg. P. Juhász, T. Kovács, G. Puskás (1², OPC). Allotype. Same as
holotype (1³, OPC).”
Diagnosis – Oláh & Kovács (2014: 110–112): “This new species col-
lected on high elevation of the Jablanica Mts. in Macedonia is a sister species of
Annitella triloba Marinkovic, 1955 and A. ostrovicensis Oláh & Kovács, 2012. It

176 J. Oláh
differs in male by having tergite VIII with reduced, mintituarized median spinate
lobe, not without any such lobe like A. ostrovicensis and not with large spinate
lobe dominating over the dorsum of tergite VIII like A. triloba; needle-pointed
paraproct without median process, with well developed median process in all
the 29 population in Albania Bulgaria, Montenegro; cerci present, vestigial at A.
ostrovicensis. Also differs in female by having sternite IX (setosa lateral lobes) and
dorsal black region of segment X differently formed. Probably an “island” allo-
patric species occurs not far from the southernmost populations of its sister spe-
cies A. triloba. However a detailed fine structure analysis of several populations
of all the three sibling species will give us more details about the early stages of
their speciation. A comparative analysis of the phallic and periphallic organs of
cerci, paraproct as well as the vaginal sclerite complex, the female sternite IX and
the black region of segment X is recommended. We have examined and recorded
very high stability of the paraproct fine structure of A. triloba in the 8 Bulagtian,
12 Montenegro and 9 Albanian populations.”
Note – Oláh & Kovács (2014: 110–112): “This isolated mountain range of
Jablanica Mts. is an endemic hot-spot. All the representatives of caddisfly groups
exhibiting pleistocen divergence evolved an incipient species in spring or lake
inflow and outflow habitats of high elevation of Jablanica mountain: Allogamus
zugor sp. n., Annitella jablanicensis sp. n., Drusus discophorus Radovanović 1942,
Potamophylax alsos sp. n. Other insect groups have also evolved endemic species in
this mountain range: a short-winged herbivorous bushcricket Poecilimon jablani-
censis Chobanov & Heller, 2010; a stonefly, Isoperla vevcianensis Ikonomov, 1980;
a high-altitude ground beetle Trechus (Trechus) nezlobinskyi Hristovski, 2014.”
Etymology – “The new species is named after the Jablanica Mts., where the
type locality is found.”
Annitella singularis (Klapálek, 1902)

Chaetopteryx singularis Klapálek, 1902: 162–164: “Vares, 8/10 1900 1²”.
Vareshiana singularis (Klapálek, 1902): Marinković-Gospodnetić
(1966b: 206): new genus erected for Chaetopteryx singularis.
Anitella singularis (Klapálek, 1902): Malicky (2005: 572): Vareshiana syn-
omymised with Annitella.
Annitella triloba Marinković-Gospodnetić , 1955

Annitella triloba Marinković-Gospodnetić , 1955: 128–129: “Na izvoru
Miljacke nađeni su primerci jedne vrste Annitella koja po svokim karakterima iz-
gleda još primitivna.” (“At the spring of River Miljacka Annitella specimens were
found which appear to be primitive.”)

Annitella kosciuszkii species complex

Examination of newly collected materials as well as the historical materials
deposited in the Klapálek’s Collection in NMP and in the Dziędzielewicz’s col-
lection in SMNHL initiated to establish the A. kosciuszkii new species complex
with four species: A. chomiacensis, A. kosciuszkii, A. lateroproducta, and A. wolo-
satka (Oláh et al. 2015b: 67).
Annitella chomiacensis (Dziędzielewicz, 1908)

Heliconis chomiacensis Dziędzielewicz, 1908 in Dziędzielewicz &
Klapálek (1908a: 22–23, 1908b: 250–255): “Jawi się w dostatocznej ilości przy
brzegach rzek: Prutec i Gnilec i przy dolnym biegu potoku: Barani na poludnio-
wych stokach góry Chomiak za Tatarowem we wschodnich Karpatach od
początku po koniec października (X). Samica o wiele rzadsza od śamca”.
Annitella chomiacensis (Dziędzielewicz, 1908): Raciecka (1934: 240–241):
Heliconis synonymised with Annitella. A. chomiacensis redescribed and redrawn.
Annitella chomiacensis chomiacensis (Dziędzielewicz, 1908): Szczęsny
(1979: 260): reduced to subspecies status based on supposed crossing between A.
chomiacensis and A. lateroproducta. Synonymised with A. kosciuszkii Klapálek,
1907: 30–31, with A. Dziędzielewiczi Schmid, 1952: 157–158. “Holotype ² et
un paratype ²: Worochta, Okolice, 12.X.1908 (Carpathes). Ils sont déposés dans
la collection de Ris. Ces specimens ont été envoyés à Ris par Dziędzielewicz, qui
les à confondus avec d’autres spécimens de kosciuszkii”. With Annitella transyl-
vanica Murgoci in Murgoci & Botoşăneanu (1957: 139–142). “Un exemplar
² (Holotip, in col. A. Murgoci), in Munţii Rodnei, pe Valea pîrîului Fîntina, in
apropierea cabanei Borşa, leg. Eleonora Erhan.”
Annitella chomiacensis (Dziędzielewicz, 1908): Szczęsny (1980: 473–
474): lectotype and paralectotype designation. In NHM-ISEA: “9²², 2³³;
East Carpathians (Gorgany to the North of Worochta); 7 specimens were la-
belled with “Heliconis Klapaleki Dziedz.”. Dziędzielewicz (Dziędzielewicz &
Klapálek 1908a) described this species from a series of specimens collected
“by the stream Gnilec, by the lower reaches of Barani stream on the southern
slopes of the summit of Chomiak and also by the rivers Prutec and Blotek” in the
autumn of 1907, beginning from the end of September. 7²² and 2³³ from the
collections in Cracow come from this time and place given by Dziędzielewicz,
thus the status of these specimens as syntypes is certain. They have the following
inventory numbers: 82/23 – (6²², 1³), 69/24 (1², 1³. From them is designated
the lectotype of male – 82/23 – with the label “Chomiak, Blotek 20.X.1907”.
Annitella chomiacensis (Dziędzielewicz, 1908): Botosaneanu (1995: 82):
reinstated the species status.

178 J. Oláh
Annitella chomiacensis (Dziędzielewicz, 1908): Szczęsny & Godunko

(2007: 35): in SMNHL: “8²², 1³; East Carpathians, Gorgany Massif (mainly
on slope of Chomiak Mt), Czarnohora Massif, (only 1² on slope of Polonina
Kozmieska); 6²² and 1³ collected by Dziędzielewicz during 4–23.X.1907
on slope of Chomiak Mt belong to the series of specimens on basis of which
Dziędzielewicz described this species, and match the criteria for paralectotypes
according to the ICZN Article 74.1, Recommendation 74.6 (No E24.12.010)
01–07). Several paralectotypes are stored in NMP (P. Chvojka, pers. comm.). The
lectotype designated by Szczęsny (1980) is stored in MP ISEZ.”
Annitella chomiacensis (Dziędzielewicz, 1908): Szczęsny & Chvojka
(2008: 156): in NMP: “Paralectotypes of 2²² and 1³ specimens were collected in
14–16.X.1907 in the East Carpathians, Chomiak, Blotek from the type locality
and deposited in NMP.”
Annitella chomiacensis (Dziędzielewicz, 1908): Szczęsny & Godunko
(2008: 65): “Streams on southern slope of the Khomiak Mts. in Gorgany are locus
typicus for the taxon.”
Annitella chomiacensis (Dziędzielewicz, 1908): Oláh et al. (2015b: 67–68):
“Ukraine: original label: “Chomiak, Blotek, 14.X.1907, leg. J. Dziędzielewicz” (1
male, Klapalek’s Collection in NMPC: K383). Ukraine: original label: “Chomiak,
Blotek, 15.X.1907, leg. J. Dziędzielewicz” (1 male, Klapalek’s Collection in
NMPC: No. 48). Ukraine: original label: “Chomiak, Blotek, 16.X.1907, leg. J.
Dziędzielewicz” (1 female, Klapalek’s Collection in NMPC: No. 41). Ukraine:
original label: “Chomiak, Blotek, X.1907” (1 male, Dziędzielewicz’s collection
in SMNHL: BS.024, E24.12.14.01/09). Ukraine: original label: “Czarnohora,
Kozmieska, 16.X.1908” (1 male, Dziędzielewicz’s collection in SMNHL: BS.012).
Ukraine: original label: “Chomiak, Blotek, 15.X.1907” (1 female, Dziędzielewicz’s
collection in SMNHL: BS.029, E24.12.14.01/04). Ukraine: original label:
“Tatarow, (Prutec), 7.X.1905” (1 male, Dziędzielewicz’s collection in SMNHL:
BS.004). Ukraine: original label: “Chomiak, Potok Barani, 16.X.1907” (1 male,
Dziędzielewicz’s collection in SMNHL: BS.028, E24.12.14.01/06). Ukraine: orig-
inal label: “Chomiak, Potok Barani, 16.X.1907” (1 male, Dziędzielewicz’s collec-
tion in SMNHL: BS.022, E24.12.14.01/05). Ukraine: original label: “Chomiak,
Blotek, 6.X.1907” (1 male, Dziędzielewicz’s collection in SMNHL: BS.025,
E24.12.14.01/01). Ukraine: original label: “Chomiak, Blotek, 4.X.1907” (1 male,
Dziędzielewicz’s collection in SMNHL: BS.023, E24.12.14.01/03).”
Diagnosis – Oláh et al. (2015b: 67–68): “The lateral lobe of the paraproct
is shorter than the mesal lobe, and the mesal lobe is very robust as visible both
in lateral and ventral view. The sclerotized tip of the aedeagus is most developed
and diverted laterad. Female anal tube with wide mesal excision in dorsal view,
lateral apodemes of the vaginal sclerite complex slender and laterad directed.

The locus typicus of this species is on the southern slope of the Khomiak Mts in
Gorgany, Ukraine.”
Annitella kosciuszkii Klapálek, 1907

Annitella kosciuszkii Klapálek, 1907: 30–31: “Naleziště: Tatarów, Błotek
(vých. Karpaty) v řijnu 1906 1². Leg. Józ. Dziędzielewicz”.
Annitella kosciuszkii Klapálek, 1907: Dziędzielewicz (1911: 46–47): fe-
male description.
Annitella kosciuszkii Klapálek, 1907: Raciecka (1934: 241–243): rede-
scribed and redrawn.
Annitella kosciuszkii Klapálek, 1907: Szczęsny (1979: 260): reduced to a hy-
brid status formed in hybridisation zone by Annitella lateroproducta × Annitella
chomiacensis.
Annitella kosciuszkii Klapálek, 1907: Malicky (2005b: 572): a hybrid status
formed in hybridisation zone by Annitella lateroproducta × Annitella chomiacen-
sis reconfirmed.
Annitella dziedzielewiczi Schmid, 1952b: 157–158: Schmid distinguished
A. Dziędzielewiczi from A. kosciuszki mostly or even exclusively by the shape of
the posterior process of tergite VIII.
Annitella dziedzielewiczi Schmid, 1952: Szczęsny (1979: 260): reduced
to a hybrid status formed in hybridisation zone by Annitella lateroproducta ×
Annitella chomiacensis.
Annitella dziedzielewiczi Schmid, 1952: Malicky (2005b: 572): hybrid sta-
tus formed in hybridisation zone by Annitella lateroproducta x Annitella chomia-
censis reconfirmed.
Annitella transylvanica Murgoci, 1957 in Murgoci & Botoşăneanu
(1957: 139–142): “Un exemplar ² (Holotip, in col. A. Murgoci), in Munţii Rodnei,
pe Valea pîrîului Fîntina, in apropierea cabanei Borşa, leg. Eleonora Erhan.”
Annitella transylvanica Murgoci, 1957: Botosaneanu (1973: 132–134):
female described, species status reconfirmed.
Annitella transylvanica Murgoci, 1957: Szczęsny (1979: 260): reduced
to a hybrid status formed in hybridisation zone by Annitella lateroproducta ×
Annitella chomiacensis.
Annitella transylvanica Murgoci, 1957: Botosaneanu (1995: 82): species
status resurrected.
Annitella transylvanica Murgoci, 1957: Malicky (2005b: 572): hybrid sta-
tus formed in hybride zone by Annitella lateroproducta × Annitella chomiacensis
reconfirmed.

180 J. Oláh
Annitella dziedzielewiczi Schmid, 1952: Oláh et al. (2015b: 69): a synonym

of A. kosciuszkii.
Annitella transylvanica Murgoci 1957: Oláh et al. (2015b: 69): a synonym of
A. kosciuszkii.
Annitella kosciuszkii Klapálek, 1907: Oláh et al. (2015b: 69): species sta-
tus restituted. “Romania: Maramures county, Rodna Mts. Borsa-Staţiunea Borsa,
stream along the road towards Prislop Pass, 1014 m, N47° 37’ 34.0’’ E24° 49’ 13.0’’,
26.ix.2006 leg. Dányi, J. Kontschan D. Murányi, (1² HNHM). Romania, Rodna
Mts. small spring streamlets on the Bistrita Aurie spring area, N47°34’23.8”
E24°48’43.9”, 1654m, 28. IX. 2014, leg. J. Oláh & Cs. Balogh (1 male, OPC).
Rodna Mts. Complex Borsa, small side spring stream of Fantana Stream, 29. IX.
2014, leg. J. Oláh & Cs. Balogh (1 male, 1 female; OPC). Ukraine: original la-
bel: “Chomiak, Blotek, 13.X.1907, leg. Lesmitz” (1 male, Klapalek’s Collection
in NMPC: No. 14). Ukraine: original label: “Worochta, Okolice, 9.X.1908, leg.
J. Dziędzielewicz (1 male, Klapalek’s Collection in NMPC: No. 13). Ukraine:
original label: “Worochta, Okolice, 23.X.1908, leg. J. Dziędzielewicz” (1 female,
Klapalek’s Collection in NMPC: No. 16). Ukraine: original label: “Worochta,
Okolice, 24.X.1908, leg. J. Dziędzielewicz” (1 male, Klapalek’s Collection in
NMPC: K 384). Ukraine: original label: “Worochta, Okolice, 26.X.1908, leg.
J. Dziędzielewicz” (1 male, Klapalek’s Collection in NMPC: No. 15). Ukraine:
original label: “Worochta, Okolice, 12.X.1908” (1 male, Dziędzielewicz’s col-
lection in SMNHL: No. 1240). Ukraine: original label: “Worochta, Okolice,
14.X.1908” (1 male, Dziędzielewicz’s collection in SMNHL: No. 1238). Ukraine:
original label: “Czarnohora, Kozmieska, 12.X.1908” (1 male, Dziędzielewicz’s
collection in SMNHL: No. 1250). Ukraine: original label: “Czarnohora,
Foreszczynka, 8.X.1910” (1 male, Dziędzielewicz’s collection in SMNHL: No.
1248). Ukraine: original label: “Worochta, 5.X.1909” (1 male, Dziędzielewicz’s
collection in SMNHL: No. 1246). Ukraine: original label: “Worochta, 9.X.1910”
(1 male, Dziędzielewicz’s collection in SMNHL: No. 1247). Ukraine: original
label: “Worochta, Okolice, 5–11.X.1908” (1 male, Dziędzielewicz’s collection
in SMNHL: No. 1246). Ukraine: original label: “Worochta, 22.X.1909” (1 fe-
male, Dziędzielewicz’s collection in SMNHL: No. 1242). Ukraine: original
label: “Czarnohora, Zawojela, 2.X.1908” (1 female, Dziędzielewicz’s collec-
tion in SMNHL: No. 1252). Ukraine: original label: “Worochta, 17.X.1909”
(1 male, Dziędzielewicz’s collection in SMNHL: No. 1253). Ukraine: origi-
nal label: “Worochta, 22.X.1909” (1 male, Dziędzielewicz’s collection in
SMNHL: No. 1244). Ukraine: original label: “Tatarow, Blotek, X.1906” (1
male, Dziędzielewicz’s collection in SMNHL: No. 1243). Ukraine: original la-
bel: “Czarnohora, Kozmieska, 17.X.1908” (1 male, Dziędzielewicz’s collec-
tion in SMNHL: No. 1249). Ukraine: original label: “Czarnohora, Kozmieska,

13.X.1909” (1 male, Dziędzielewicz’s collection in SMNHL: No. 1241). Ukraine:

original label: “Worochta, Okolice, 8.X.1908” (1 male, Dziędzielewicz’s collec-
tion in SMNHL: No. 1245). Ukraine: original label: “Worochta, 9.X.1910” (1
male, Dziędzielewicz’s collection in SMNHL: No. 1255).
Diagnosis – Oláh et al. (2015b: 69): “The lateral lobe of the paraproct has
almos equal length with the mesal lobe, and the mesal lobe is slender as visible
both in lateral and ventral view. The sclerotized tip of the aedeagus has some flat
apical surface. Female anal tube without wide mesal excision in dorsal view, lat-
eral apodemes of the vaginal sclerite complex rounded. The locus typicus of this
species is in the Czarnohora Mts. Ukraine.”
Annitella lateroproducta (Botoşăneanu, 1952)

Carpathopsyche lateroproducta Botoşăneanu, 1952c: 1–15: “Les materi-
aux dont nous disposons jusqu’ à présent provinnent exclusivement du massif
du Retezat, region importante de la chaîne des Carpathes Méridionaux de la
R. P. R. Les localités qui ont fourni le matériel prelévé par nous des collections
de l’expédition hydrobiologique de 1946, sons les suivantes: 1. Lac de Bucura
(Viorica). Littoral, au niveau de la cascade. 2. Lac de Stanisoara. Littoral Nord.
3. Torrent droit de la source du lac de Galesul. 4. Embouchure de la source
d’evacuation du lac de Bucura (lac principal)”
Annitella lateroproducta (Botoşăneanu, 1952): Murgoci & Botoşăneanu
(1957: 142): Carpathopsyche is synonymised with Annitella.
Annitella lateroprocucta lateroproducta (Botoşăneanu, 1952): Szczęsny
(1979: 260): reduced to subspecies status based on supposed crossing between A.
lateroproducta and A. chomiacensis.
Annitella lateroproducta (Botoşăneanu, 1952): Botosaneanu (1995: 82):
reinstated the species status.
Annitella lateroproducta (Botoşăneanu, 1952): Oláh et al. (2015b: 71):
“Romania: Apuseni Mts. Vadul Crisului, Crisul Rapide, 29. X. 1997, leg. L. Újvárosi
(1², OPC). Apuseni Mts. Ic Ponor, spring area of Somesul Cald, 6. XI. 1998, leg. L.
Újvárosi (5²,4³, OPC). Apuseni Mts. Arieseni, Alboc, 6. X. 1999, leg. L. Theodor
(1²,1³, OPC). Gilau Mts. Jerii Valley, 8. X. 2000, leg. L. Újvárosi (1², OPC).
Apuseni Mts. Doda Pilii, 3. XII. 2006, leg. L. Újvárosi (6²,4³, OPC). Apuseni
Mts, Valea lui Dragan, 650m, N46.83119 E22.77093, 20.xi.2008 leg. M. Bálint &
Tasnádi (1², OPC). Apuseni Mts. Sebes Körös valley, Suncuius, near Izbandis
spring, 26. X. 2009 singled leg. J. OLÁH & M. BÁLINT (7², OPC). Apuseni Mts,
Padis, open stream near pine forested sphagnum bog, N46° 35’ 20.632 E22° 45’
54.857, 5.XI.2011, leg. Gy. Monori, J. Oláh & L. Szél (8²,6³, OPC). Hargitha Mts.
Sincraieni, Valea Mare, 6–14. IX. 1993, light trap, (23², OPC). Ciucaş Mts. 3 km

182 J. Oláh
S of Dălghiu, Dălghiu stream, N45°33’00.2”, E25°54’43.5”, 970 m, 13.10.2011, leg.

Á. Ecsedi, T. Kovács, G. Puskás, (1²,1³, OPC). Caraş-Severin county, Ţarcu Mts.,
open stream with Salix bushes 6 km S of Poiana Mărului, 1000 m, N45°20’47.5”,
E22°31’04.6”, 14.10.2011, leg. Á.Ecsedi, T. Kovács, G. Puskás, (1³, OPC). Caraş-
Severin county, Ţarcu Mts., left side brook of open stream on the N slope of Mt.
Ţarcu, 1500 m, N45°17’40.7”, E22°31’44.5”, 14.10.2011, leg. Á.Ecsedi, T. Kovács,
G. Puskás, (1²,1³, OPC). Caraş-Severin county, Ţarcu Mts., open stream on
the N slope of Mt. Ţarcu, N45°17’46.2”, E22°31’41.5”, 1500 m, 14.10.2011, leg.
Á.Ecsedi, T. Kovács, G. Puskás, (5²,2³, OPC). Caraş-Severin county, Semenic
Mts., open brook E of Mt. Piatra Goznei, N45°10’55.4”, E22°04’01.4”, 1340 m,
15.10.2011, leg. Á. Ecsedi, T. Kovács, G. Puskás, (1³, OPC). Gurghiu Mts. near
Bucin Pass, Tárnava Mica springs and stream, N: 46°39’ 16,63”E: 25°16’ 42,46”,
1290, 30.X.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (1 male; OPC).
Hargitha Mts. Filio stream, N: 46°27’ 03,90” E: 25°30’ 20,10”, 940m, 31.X.2014
leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (1 male, 1 female; OPC). Dâmbovia
county, Bucegi Mts, Hotel Peştera, Ialomiţa, 45°23’54.5”, 25°26’25.1”, 1610 m,
07.11.2014, leg. T. Kovács & G. Magos (1 male, OPC). Sibiu county, Făgăraş Mts,
Cârţişoara, Bâlea Stream below the Bâlea Lake, 45°36’30.4”, 24°37’14.6”, 1940 m,
08.11.2014, leg. T. Kovács & G. Magos (1 male, OPC).”
Diagnosis – Oláh et al. (2015b: 71): “The lateral lobe of the paraproct is
longer than the mesal lobe, and the mesal lobe is reduced digitiform as visible
both in lateral and ventral view. The sclerotized tip of the aedeagus is blunt.
Female anal tube tappering in dorsal view, lateral apodemes of the vaginal scle-
rite complex rounded laterad directed. The locus typicus of this species is in the
Retezat Mts. Romania.”
Annitella wolosatka Oláh et Szczęsny, 2015

Annitella chomiacensis (Dziędzielewicz, 1908): Szczęsny (1966: 344–346):
misidentification.
Annitella wolosatka Oláh et Szczęsny, 2015 in Oláh et al. (2015b: 72–73):
“Holotype: Poland, East Carpathians, Bieszczady Mts. at Wolosatka brook, 9oom,
28.X.2010, leg B. Szczęsny (1 male, OPC). Allotype: same as holotype (1 female,
OPC). Paratypes: same as holotype (2 males, OPC). Poland, East Carpathians,
Bieszczady Mts. at Wolosatka brook, 850–1000m, 22.X.2014, leg B. Szczęsny (3
males, OPC).”
Diagnosis – Oláh et al. (2015b: 72–73): “The lateral lobe of the paraproct
is very short and directed laterad; the mesal lobe is very robust with rounded
lateral margin as visible in lateral view. The sclerotized tip of the aedeagus is less
developed, very blunt. Female anal tube with wide mesal excision and produced

an addition small mesal lobe on the mesal tip of the lateral lobes in dorsal view;
lateral apodemes of the vaginal sclerite complex straight. Reduced male tibial
spur formula differs from the other member of the species complex.”
Etymology – “This new species was named after the Wolosatka Stream valley
in the Beszczady Mts. where the type material was collected.”
Chaetopteroides Kumanski, 1987

Chaetopteroides bulgaricus (Kumanski , 1969)
Chaetopteryx bulgaricus Kumanski, 1969b: 21–27: “Material und Fundort:
10.X.1967, Pirin-Gebirge, Banderitza-Tal, Bach über der Berghütte “Wichren”,
2²², 1³ und im Abfluss der Muratowi-Seen (gleicher Bezirk), 1², 2³³. Höhe
über dem Meeresspiegel entsprechend 2100 und 2200 m.” Holotypus und die
Paratypen (2²², 3³³) in Sammlung (in Alcohol) des Zoologischen Museums der
Bulgarischen Akademie der Wissenschaften”.
Chaetopteroides bulgaricus (Kumanski, 1969): Kumanski (1987b: 15):
Chaetopteroides gen. n. erected.
Chaetopteroides bulgaricus (Kumanski, 1969): Oláh et al. (2013b: 99–
100): “Additional female was collected in Pirin Mts below Bezbog on 2240 m in
18.IX.1968 (Kumanski 1971). Later a single female in Rila Mts. at tributary of
Beli Iskar ob Borowez, 2300 m, 23–24.VIII.1971 and a single male at the tribu-
tary of Beli Iskar ob Borowez 1200–1800 m, 24.VIII.1971 have been collected
(Kumanski & Malicky 1976).”
Chaetopteroides bulgaricus (Kumanski, 1969): Oláh & Kovács (2012):
Rila Mts. Beli Iskar, 1900 m, 23.VIII.1971 leg Braasch, (1², 1³, OPC present
from MPC). Oláh & Kovács (2012): Rila Mts. Borovets, Zavrachitsa hut, Prava
Maritsa, N42°10’04.9”, E23°38’28.1”, 2200 m, 05.10.2011, leg. Á. Ecsedi, T.
Kovács, G. Puskás (1³, OPC). Pirin Mts, 950 m S of Demianitsa hut, left side
brook of Valyavitsa stream, N41°44’02.6” E23°28’03.1”, 2020 m, 07.10.2011, leg.
Á. Ecsedi, T. Kovács, G. Puskás (1², 6³, OPC). Pirin Mts. 1.5 km E of Begovitsa
hut, Begovitsa stream, N41°40’32.6” E23°26’38.8”, 1930 m, 08.10.2011, leg.
Á. Ecsedi, T. Kovács, G. Puskás (1², OPC). ”Bulgaria, Blagoevgrad province,
Pirin Mts, Bansko, stream in pine shrub above the Vihren hut, N41°45.293’
E23°24.933’, 1995 m, 24.X.2013, leg. J. Kontschán, D. Murányi, T. Szederjesi
(1², HNHM).”
Diagnosis – Oláh et al. (2013b: 99–100): “Cerci simple without addi-
tional setose process of subdivion. Paamere apex multidenticulate. Anal tube
with short dorsolateral setose lobe and ventrolateral lobe-like setose surface.
Sternite IX high. Vulvar scale simple rounded in lateral view, median lobe short
rounded.”

184 J. Oláh
Chaetopteroides kosovarorum Ibrahimi et Oláh, 2013

Chaetopteroides kosovarorum Ibrahimi et Oláh, 2013 in Oláh et al. (2013b:
101): “Holotype. Kosovo, Mitrovicë Municipality, Bajgorë area, entrance into
the Kaçandoll village from Mitrovicë side, sidespring of the Kaçandoll River by
the main road, N42.979° E21.0509°, 1262 m, 29.10.2013, H. Ibrahimi, F. Asllani
Ibrahimi, Irsa Ibrahimi & Idlir Ibrahimi (1², DBFMNSUP). Paratypes. Same as
holotype (2², DBFMNSUP). Same place as holotype, 18.09.2012, light trap, H.
Ibrahimi (2², DBFMNSUP); 25.10.2013, H. Ibrahimi (1², OPC).DBFMNSUP”
Diagnosis – Oláh et al. (2013b: 101): “This large sized species with few light
spots on male forewing and with subdivided cerci is more similar to C. veges sp.
n., but differs by the modified parameres. Parameres became elongated thin-slen-
der with bulbous basement, apical teeth reduced in size, almost minituarized and
their number multiplied up to 8–10.”
Etymology – “Kosovarorum from “kosovar”, the inhabitants of Kosovo.”
Chaetopteroides maximus (Kumanski, 1968)

Chaetopteryx maximus Kumanski, 1968b: 59–61: “Fundort: Vitosa-
Gebirge, beim Bergbach, nich weit von der Berghütte “Bor”, 1620 m Höhe,
1.X.1958, 7²² (leg. N. Vihodcevski). Holotypus und 1 paratypus ² in der
Kollection von F. Schmid (Ottawa); 5²² Paratypen in der Insektensammlung
des Zoologischen Museums der Bulgarischen Akademie der Wissenschaften.”
Chaetopteroides maximus (Kumanski, 1968): Chaetopteroides gen. n. erected
by Kumanski (1987b: 15).
Chaetopteroides maximus (Kumanski, 1968): Oláh et al. (2013: 102–103):
“In Vitosha Mts Kumanski (1971) has collected 3 males in Zlatnite bridge and 1
male below Rodina in 19.10.1968. A single female was collected on Vitosha Mts
on 19.10.1974, near the type locality (Kumanski 1987). Oláh & Kovács (2012):
Vitosha Mts, spring and brook 200 m E of Rodina hut, N42°37’09.6”, E23°15’32.3”,
1600 m, 03.10.2011, Á. Ecsedi, T. Kovács, G. Puskás (4³, OPC). Vitosha Mts,
Lavchemo, Boyanska Reka, N42°34’34.6” E23°16’57.7”, 2050 m, 04.10.2011,
Á.Ecsedi, T. Kovács, G. Puskás (1², 3³, OPC; 1², 1³, MM). Marinković-
Gospodnetić (1980) has reported the species from Serbia: spring area of Lisinska
River, Kopaonik Mts, 1², 2³, 08.10.1978. Marinkovic’-Gospodnetić’s specimens
have been lost: the entire collection was destroyed during the Bosnian war be-
tween 1992 and 1995 (personal communication by H. Malicky). New collection
is required to confirm its real taxonomic position. It is probably not C. maximus.
C. maximus is known as endemic to Vitosha Mts. Bulgaria.
Diagnosis – Oláh et al. (2013: 102–103): “Cerci with additional setose
process of subdivion. Paramere apex linear denticulate laterad. Anal tube with

long dorsolateral setose lobe, ventral setose surface divided. Sternite IX rounded
low. Vulvar scale humped rounded in lateral view; median lobe less developed
triangular.”
Chaetopteroides tunik Oláh, 2013

Chaetopteroides tunik Oláh, 2013 in Oláh et al. (2013b: 103–104):
“Holotype. Macedonia, Vardar region, Kožuf Mts, open brook in alpine grass-
land towards Ski Kožuf, N41°12.560’, E22°13.170’, 1670 m, 04.10.2013, T. Kovács,
D.Murányi (1², OPC).”
Diagnosis – Oláh et al. (2013b: 103–104): “This medium sized species with
narrow and long forewing without any pattern and having subdivided cerci is
more similar to C. kosovarorum, but differs by having cerci, elongated not stalked;
gonopod without vertical ridge; paramere less slender, straight in dorsal view,
not arching mesad and the minute teeth limited to terminal position and their
number are reduced to 4. This description is based on a single holotype male;
female is required to confirm its position.”
Etymology – “Tunik from “tűnik” appear and disappear in Hungarian, refers
to just appearing, almost disappearing setae on the parameres.”
Chaetopteroides veges Oláh, 2013

Chaetopteroides veges Oláh, 2013 in Oláh et al. (2013b: 104–106):
“Holotype. Bulgaria, Kyustendil province, Osogovska planina, spruce forest, for-
est brook below Trite buki hut, N42°10.463’, E22°38.066’, 1520m, 23.10.2013, J.
Kontschán, D. Murányi, T. Szederjesi (1², HNHM). Allotype. Same as holotype
(1³, HNHM). Paratypes. Same as holotype (1³, HNHM). Kyustendil province,
Osogovska planina, beech forest and forest brook at Iglika hut, N42°13.783’,
E22°38.842’, 1325m, 23.10.2013, J. Kontschán, D. Murányi, T. Szederjesi (1 fe-
male, HNHM).
Diagnosis – Oláh et al. (2013b: 104–106): “This large sized species with
narrow and long light-spotted male forewing and brachypterous fused-spotted
female forewing having subdivided cerci is more similar to C. maximus, but dif-
fers by having male with paramere setae limited to terminal position and their
number are reduced to 2–3. Differs by female by having apical lateral lobes much
longer than dorsolateral lobes; ventral setose surface on the anal tube fused, not
divided; sternite IX higher; vulvar scale hooked in lateral view, not humped; me-
dian lobe of the vulvar scale more developed.”
Etymology – “Veges from “véges” meaning terminal/apical in Hungarian, re-
ferring to the few setae on the parameres present and limited to the terminal/
apical area.”

186 J. Oláh
Chaetopterygopsis Stein, 1874

Chaetopterygopsis sisestii Botosaneanu, 1961
Chaetopterygopsis sisestii Botosaneanu, 1961b: 61–64: “Vallée du Delghiu,
torrent a 800 m. alt. environ, bassin superieur du Buzea, au pied du Massif Ciucas,
Carpates Orientales, 24.X.1960, 11 ², 1³ (holotype ², allotype ³, 10 paratypes
²); Rivière Motru à Closani, Monts de l’Olténie du Nord, 17.X.1960 (1². Leg. I.
Tabacaru). Holotype ² et allotype ³ dans les collections du Musée Zoologique de
Lausanne; le rest du materiel se trouve dans ma collection.”
Chaetopteryx Stephens, 1829

Chaetopteryx aproka Oláh, 2011
Chaetopteryx aproka Oláh, 2011b: 9–10: “An unusually small-sized Chaeto-
pteryx species with reduced tibial spur number and open anal tube was collected first
by David Murányi.” “Holotype. Romania: Maramureş county, Munţii Ignis, Deseşti-
Staţiunea Izvoare, open brook with spring bog on the Valhani Plateau, 1020m,
N47°43’01.0” E23°44’32.1”, 24.09.2005, leg. J. Kontschán, D. Murányi, J. Nédli (1
male NHMB). Paratypes. Romania: Maramureş county, Munţii Ignis, Deseşti-
Staţiunea Izvoare, open brook with spring bog on the Valhani Plateau, 1020m,
N47°43’01.0” E23°44’32.1”, 24.09.2005, leg. J. Kontschán, D. Murányi, J. Nédli (2
males, NHMB). Maramureş county, Munţii Ignis, Deseşti-Staţiunea Izvoare, open
brook with spring bog on the Valhani Plateau, 1020m, N47°43.015’ E23°44.547’,
07.10.2010, leg. P. Barcánfalvi, D. Murányi & J. Oláh (15 males, 2 females, OPC).
Maramureş county, Munţii Ignis, Deseşti-Staţiunea Izvoare, forest spring at settle-
ment, 920m, N47°45.167’ E23°43.013’, 08.10.2010 leg. P. Barcánfalvi, D. Murányi
& J. Oláh, (28 males, 10 females, OPC). Maramureş county, Munţii Ignis, Deseşti-
Staţiunea Izvoare, forest spring and spring brook at settlement, 920m, N47°45.167’
E23°43.013’, 20.X.2010 leg. Á. Ecsedi, J. Oláh & I. Szivák (12 males, 10 females,
OPC). Maramureş county, Munţii Ignis, Deseşti-Staţiunea Izvoare, forest spring
and spring brook at settlement, 920m, N47°45.167’ E23°43.013’, 21.X.2010 leg. Á.
Ecsedi, J. Oláh & I. Szivák (21 males, 14 females, OPC). Maramureş county, Munţii
Ignis, Deseşti-Staţiunea Izvoare, forest spring and spring brook at settlement, 920m,
N47°45.167’ E23°43.013’, 22.X.2010 leg. Á. Ecsedi, J. Oláh & I. Szivák (14 males, 6
females, OPC). Maramureş county, Munţii Ignis, Deseşti-Staţiunea Izvoare, small
spring brook on the Valhani Plateau, 1020m, N47°43.015’ E23°44.547’, 21.X.2010,
leg. Á. Ecsedi, J. Oláh & I. Szivák (25 males, 8 females, OPC).” “The paratypes
are deposited in the following collections: Oláh Private Collection (Debrecen,
Hungary), Malicky Private Collection (Lunz-am-See, Austria), Museo Civico di
Scienze Naturali (Bergamo), Naturhistoriska Riksmuseet (Stockholm), National
Museum of Natural History, Smithsonian Institution (Washington, DC).”

Diagnosis – Oláh (2011b: 9–10): “Despite its reduced spur number this
species is a true Chaetopteryx and despite its modified female anal tube it belongs
to the Chaetopteryx rugulosa species group.”
Etymology – “The name refers to the unusually small size of this species, tiny
“aproka” in Hungarian.”
Chaetopteryx biloba Botoşăneanu, 1960

Chaetopteryx biloba Botoşăneanu, 1960b: 116–118: “Ogasul Ulmului,
source affluente du ruisseau Frasincea, à Cornereva bassin de Belareca, Banat,
Roumania), 10.X.1956. Holotype ², allotype ³, dans les collections du British
Museum (Nat. Hist.), Department of Entomology. Un paratype ² dand le collec-
tion de l’auteur.”
Chaetopteryx bosniaca Marinković-Gospodnetić, 1955

Chaetopteryx bosniaca Marinković-Gospodnetić, 1955: 125–128:
“Primerci uhvaćeni na vrelu Bosne, Večerici (prvoj desnoj pritoci Bosne) i na
Treskavici (i to uz potok koji spaja Platno sa Velikim jezerom) razlikuju se od
dosad opisanih vrsta.” “U zbirci Biološkog instituta pri Zemaljskom muzeju u
Sarajevu nađeni su primerci Ch. bosniaca sa sledećih lokaliteta: Vojkovići, Ledići,
Pazarić i Vrelo Bosne. Svi ti primerci su bili određeni kao Chaetopteryx vilosa
[sic!] Fabr.” (“The specimens collected at Vrelo Bosne, Večerica (the first right
tributary of the River Bosna) and at Treskavica (and along the creek connecting
Platno and Big lakes) differ from the species described earlier.” “In the collection
of the Biological Institute of the National Museum in Sarajevo there are spec-
imens from Vojkovići, Ledići, Pazarić and Vrelo Bosne. These specimens were
identified as Chaetopteryx villosa Fabr.”)
Chaetopteryx cissylvanica Botoşăneanu, 1960b: 118–120: “Ogasul lui Ro-
set, affluent gauche de la Cerna, en amont de Bâile Herculane, Banat, Roumanie,
13.X.1957. Holotype ² et al.lotype ³, dans la collection F. Schmid (Lausanne). 2
² paratypes, dans les collections de l’auteur.”
Chaetopteryx cissylvanica Botoşăneanu, 1960: Malicky (2005: 573):
synonymised with Chaetopteryx bosniaca Marinković-Gospodnetić, 1955.
Chaetopteryx fontisdraconis Botosaneanu, 1993: 399–402: “1² (holo-
type), and 3³ (allotype and paratypes), in alcohol; 22.X.1971, Romania, Oltenia,
district Gorj: Village Runc (Runcu), Izvoarele (= the springs) La Balaure”.
Deposited in the Zoological Museum of the University of Amsterdam.”
Chaetopteryx fontisdraconis Botosaneanu, 1993: Malicky (2005: 573):
synonymised with Chaetopteryx bosniaca Marinković-Gospodnetić, 1955.
Chaetopteryx gonospina Marinković-Gospodnetić, 1966

188 J. Oláh
Chaetopteryx gonospina Marinković-Gospodnetić, 1966a: 110–112:

“Holotype ², Bosnia, Olovo, 7.X.1964. in author’s collection; 2 paratypes ²,
Bosnia, Sarajevo, Crepoljsko, one in colection of Zemaljski muzej, Sarajevo, the
other in Schmid’s collection, Ottawa.”
Chaetopteryx polonica Dziędzielewicz, 1889

Chaetopteryx polonica Dziędzielewicz, 1889: 112, 117–118: “Znajdowa-
łem 3 i 16 października w Młodiatynie, na podgórzu koło Kołomyi przy potoku w
borze po burzanach i podrostach jodłowych w małej ilości (5², 1³).”
Chaetopteryx polonica Dziędzielewicz, 1889: Szczęsny (1980: 472): in
NHM-ISEA: “4²², 1³; the specimens undoubtedly come from the typical se-
ries from which Dziędzielewicz described this species. In the publication, in
which the author describes the species (Dziędzielewicz, 1889) is the informa-
tion that the specimens were caught “at Mlodiatyn near Kolmyja by a stream
3.X. and 16X.” Two males with inventory number 8/6 have the following la-
bels: “Mlodiatyn 1887” and 2.X.Ml.”. The other two males with inventory
number 75/9 have identical labels “16.X.Ml.” and the female with inventory
number 33/8 “Mlodiatyn 16.X. (Kolom.). A male from no 75/9 I am designat-
ing as lectotype.”
Chaetopteryx polonica Dziędzielewicz, 1889: Szczęsny & Godunko
(2007: 35): in SMNHL: “18²², 9³³; West Carpathians, Beskid Wyspowy; East
Carpathians, Gorgany Massif (Chomiak); 1³ from Gorgany Mts was wrongly
determined “Chaetopteryx major”. Although the species was discovered and de-
scribed by Dziędzielewicz at Mlodiatyn village, at border between East Carpathian
Foothills and East Carpathians (the Pokucie-Marmarosch Carpathians), none of
specimens deposited in SMNHL belongs to the type series which is housed in
Cracow.”
Chaetopteryx polonica Dziędzielewicz, 1889: Szczęsny & Chvojka
(2008: 157): in NMP: Specimens of 4²² and 1³ were collected in 22–25.
IX.1906, in 29. IX.1907 and in 5.X.1907 in the East Carpathians, Chomiak
as well as in 11.IX.1908 in Pod Dancerz and deposited in NMP. Szczęsny &
Godunko (2008): a stream (probably Kobylytsia) at Molodiatyn village (bor-
der line between the Carpathian foothills and the Carpathians) is locus typicus
of the species.
Chaetopteryx stankovici Marinković-Gospodnetić, 1966

Chaetopteryx stankovici Marinković-Gospodnetić, 1966a: 112:
“Holotype ² and allotype ³: Serbia, mountain Kopaonik, 29. X. 1952. Leg. D.
Filipovic.”

Chaetopteryx subradiata Klapálek, 1907

Chaetopteryx subradiata Klapálek, 1907: 27–28: “Jeden párek sbíral Józ.
Dziędzielewicz, 22 a 23. IX. na Chomiaku (Błotek). Samička chycena v myslivně
na okně.”
Chaetopteryx uherkovichi Oláh, 2011

Chaetopteryx uherkovichi Oláh, 2011a: 118–119: “Holotype: Croatia,
Krndija Mts., 6 km N of Kutjevo, Velika rijeka, springs, N45˚28’59”, E17˚51’33”,
580 m, 4.11.2011, leg. Á. Uherkovich & I. Szivák (1 male, OPC).”
Diagnosis – Oláh (2011a: 118–119): “This new species belongs to the
Chaetopteryx major species group and is a close relative of the Chaetopteryx major
McLachlan. They live together in the Papuk Mountains. C. uherkovichi new spe-
cies can be easily distinguished from C. major by having spur number 133, not
033; posterodorsal spinate area on segment VIII vestigial, not well-developed;
dorsum of segment IX long, not very short; cercus sallowly excited in dorsal view,
not deeply; paraproct modified having outer arm shifted laterad; subanal plate
extremely broadened; gonopods differently shaped.”
Etymology – “Patronym in honor of the collector Ákos Uherkovich, who
has contributed most significantly to the knowledge of the Hungarian caddisfly
fauna, today Hungary is one of the most studied area in our biosphere.”
Chaetopteryx rugulosa species group

This species group was established by Malicky et al. (1986) with four
known and four new taxa. After a quarter of century, applying the phylogenetic
species concept and the sexual selection theory we have revised the species group,
established three subgroups, two species clusters and described seven new spe-
cies (Oláh et al. 2012). Malicky (2014) synonymised our three species and
questioned all the others, while arguing against the application of the phyloge-
netic species concept and the sexual selection theory, but without arguments. His
nomenclatural acts were rather autocratic contradicting also to his earlier state-
ments (Malicky et al. 1986). Malicky’s taxonomic actions have been realised
without any factual explanations and criticisms regarding the divergence diag-
noses of the new clades and species and without giving his own new diagnosis
explaining the divergences remained in his new synonymised combined taxa.
Based upon (1) new population sampling, re-examination of the old ma-
terials; (2) relying on the theoretical progress in species delineation, taxa de-
limitation, reproductive isolation, and phylogenetic species; (3) exploring the
discovery of speciation traits, as a powerful tool in phenomics we have revised

190 J. Oláh
the species group and described new species (Oláh et al. 2015b). At this stage
the Chaetopteryx rugulosa species group is comprised of 21 species: Chaetopteryx
balcanica Oláh, 2015, C. clara McLachlan, 1876, C.euganea Moretti et Malicky,
1986, C. giuliensis Oláh et Kovács, 2012, C. goricensis Malicky et Krušnik, 1986,
C. idriensis Oláh et Urbanič, 2012, C. irenae Krušnik et Malicky, 1986, C. kam-
nikensis Oláh et Urbanič, 2012, C. karima Oláh, 2015, C. kozarensis Oláh, 2015,
C. marinkovicae Malicky et Krušnik, 1988, C. mecsekensis Nógrádi, 1986, C. no-
ricum Malicky, 1976, C. papukensis Oláh et Szivák, 2012, C. pohorjensis Oláh et
Urbanič, 2012, C. prealpensis Oláh, 2012, C. psunjensis Oláh, 2015, C. rugulosa
Kolenati, 1848, C. schmidi Botoşăneanu, 1957, C. tompa Oláh, 2015, C. zalaensis
Oláh, 2012.These species are diverged in clades of three subgroups and two spe-
cies clusters.
Chaetopteryx schmidi species subgroup

Chaetopteryx balcanica Oláh, 2015
Chaetopteryx schmidi Botoşăneanu, 1957: Oláh et al. (2012: 62): all the
specimens from Serbia and Bosnia-Herzegovina were misidentified.
Chaetopteryx balcanica Oláh, 2015 in Oláh et al. (2015b: 78–81): “Holotype:
Serbia: Derdap Mts. Donji Milanovac, Grgeci spring and its outlet in a beech
forest, 500 m, N44° 28’ E22° 02’, 13.X.2006, leg. L. Dányi, J. Kontschán & D.
Murányi (1 male, in copula with the allotype, HNHM). Allotype: same as holo-
type (1 female in copula with the holotype; HNHM). Paratypes: same as holo-
type (1 male, 1 female, OPC). Miroc, D. Milanovac, Stream Supljanka, 8.X.1984,
leg. Branceli (1 female, PMS). Pesaca, Donji Milanovac, 9.X.1986, leg. I. Sivec &
B. Horvát (1 male, PMS). Popadija, Donji Milanovac, 9.X.1986, leg. I. Sivec & B.
Horvát (1 male, PMS). Derdap Mts. Golubinje, stream valley with young forest,
N of the village, 88 m, N44° 30’59.6’ E22° 12’41.5”, 13.X.2006, leg. L. Dányi, J.
Kontschán & D. Murányi (6 males, HNHM). Derdap Mts. Dobra, Reka Pesaca,
beech forest with stream, 386 m, N44° 34,670 E21° 59,250, 28.X.2010, leg. L.
Dányi, J. Kontschán & Zs. Ujvári. Murányi (2 males, 1 female; HNHM). Bosnia-
Hercegovina: Kravica, Zvornik, 5.X.1986, leg. I. Sivec & B. Horvát (1², PMS).
Dobrovci,Gracanica, 430m, 11.X.1990, leg. B. Horvát & I. Sivec (2 males, 4 fe-
males; PMS). Blagijevici, Ozren Planina, 390 m, 12.X.1990, leg. B. Horvát & I.
Sivec (1 female, PMS). Kamensko, River Krivaja, 15.X.1990, leg. B. Horvát & I.
Sivec (2 males, PMS). Cunista, River Krivaja, 450m, 15.X.1990, leg. B. Horvát &
I. Sivec (1 male, 1 female; PMS). Skender, vakuf, 820m, 19.X.1990, leg. B. Horvát
& I. Sivec (2 males, PMS).”
Diagnosis – Oláh et al. (2015b: 78–81): “This new species having stout
spine-like terminal shaft on the paramere as well as having no setose lateral lobes

on female tergite IX belongs to the C. schmidi subgroup, but differs from all the
know species by having combined character state: (1) Shallow curvature on para-
proctal hook, not deep curving more anterad as at C. schmidi; (2) Variously sized
lateral process, but without any sclerotized basal tube; only a sclerotized basal
ring is present permitting a longitudinal position of the inflated and protruded
membranous process; in many specimens the membranous process is withdrawn
inside the cylinder, not visible at all; at C. schmidi the sclerotized basal tube is
present and producing and supporting a perpendicular position of the protrud-
able and inflatable membranous posterior part of the lateral process; at C. pa-
pukensis the sclerotized basal tube is present, but almost longitudinal and the
membranous process is not retractable entirely; there is no any specimens with-
out lateral process. (3) Primary spine long and curved like at C. papukensis, but
paramere shaft is not triangular in dorsad view. (4) Female anal tube quadrangu-
lar with less protruded inner sclerite. We presume that C. balcanica sp. n. is the
ancestral species of the C. schmidi subgroup widely distributed from East Serbia
through Bosnai-Herzeovina, but we need to examine more specimens from more
populations to confirm its relations.”
Etymology – “This new species was named after its wide distribution in the
Balkan Peninsula.”
Chaetopteryx karima Oláh, 2015

Chaetopteryx papukensis Oláh et Szivák, 2012 in Oláh et al. (2012: 62) (par-
tim): misidentification.
Chaetopteryx karima Oláh, 2015 in Oláh et al. (2015b: 81): “Holotype:
Bosnia & Herzegovina: Banja Luka region, Kozara Mts, forest brook below the
Vrbaška – Kozarac road, 45°02.480’, 16°54.266’, 560 m, 07.11.2012, leg. T. Kovács
& G. Magos (1², OPC).”
Diagnosis – Oláh et al. (2015b: 81): “This single male was collected in a
small forest stream where an unknown Leuctra species was also collected in pre-
vious spring, still waiting to describe. Earlier without experiences on speciation
strait and the application of fine structure analysis we have determined this speci-
men and listed as a paratype of C. papukensis (Oláh et al.. 2012). Although the fe-
male is unknown, the strongly developed paramere spine clearly relates this new
species into the C. schmidi subgroup. The lack of setose lateral lobes on female
tergite IX would further confirm the phylogeny of this interesting species. The
presence of well elaborated supporting sclerite system of flange and ridge for-
mation on the aedeagus is present only in C. irenae subgroup and in C. noricum
species cluster. The detection of this much specialised trait of deeper/older di-
vergence in C. schmidi subgroup seems discordant and explainable by incomplete

192 J. Oláh
lineage sorting. Having this specific supporting sclerite system on the aedeagus
present this is a well diverged new species, however female traits of lateral setose
lobe and anal tube formation would give more information about its relations.”
Etymology – “Karima from “karima” flange in Hungarian, refers to the dou-
ble pairs of flange ridge formation on the aedeagus.”
Chaetopteryx kozarensis Oláh, 2015

Chaetopteryx papukensis Oláh et Szivák, 2012 in Oláh et al. (2012: 62) (par-
tim): misidentification.
Chaetopteryx kozarensis Oláh, 2015 in Oláh et al. (2015b: 81–83): “Holo-
type: Bosnia & Herzegovina: Banja Luka region, Kozara Mts, forest edge spring 1
km S of peak Lisina, 44°57.773’, 16°58.342’, 680 m, 7.XI.2012, leg. T. Kovács & G.
Magos (1 female, OPC). Allotype: same as holotype (1 male, OPC). Paratypes:
same as holotype (4 males, 5 females; OPC).”
Diagnosis – Oláh et al. (2015b: 81–83): “Most close to C. papukensis, but
differs by having apical hook formation of the dorsal branch of the paraproct
blunt, not pointed and the curvature is very shallow. The most striking diver-
gence from all the known female anal tube in the species group is detected in
the dorsal configuration. This is why the female was designated as holotype. The
unique dorsal profile of the anal tube is characterized by a deep V-shaped exci-
sion; however we do not know how the anchoring pressure may modify the shape
of the excision by moving the internal sclerite backward. It is actually not known
whether this internal sclerite is movable at all or not. The bilobed posterior end
of the internal sclerite is in protruded state in all of the examined hundreds of C.
papukensis. This may suggest that the internal sclerite, that is the vestigial tergite
X fixed withdrawn inside the anal tube is not movable even under the long an-
choring presssure of the paraproctal hook.”
Etymology – “This new species was named after the mountain range where
the type material was collected.”
Chaetopteryx mecsekensis Nógrádi, 1986

Chaetopteryx schmidi mecsekensis Nógrádi, 1986 in Malicky et al. (1986:
8–10): “Die Adulten findet man an Quellen und entlang der Oberlaufe von klein-
en Bachen im Mecsek-Gebirge sowohl auf Kalk als auch auf Sandstein. Sie sind
flugunfähig, laufen aber entlang der Bäche rasch herum, manchmal noch auf
Schnee. Die Aktivitätsperiode dauert von Anfang Oktober bis Anfang Januar.
Die Tiere sind auch noch bei Lufttemperaturen von minus 2–3 Grad Celsius aktiv
und überleben minus 10–12 Grad.” “Untersuchtes Material (alles vom Mecsek-
Gebirge in Südungarn): Holotypus ²: Tal “Meleg-mány”, 20. 12 1983, leg. A.

Uherkovich, in coll. Janus Pannonius Museum, Pécs, als Flüssigkeitspraparat,

Genitalpräparat Nr.282. Paratypen: Mánfa, bei der Höhle “Kőlyuk”, 26. 11. 1982:
1 ²; Tal “Meleg-mány”, 8. 10. 1983: 31 ²², 12 ³³, 22. 10. 1983: 10 ²², 10 ³³, 7. 11.
1983: 2 ²², 20. 12. 1983: 1 ², 6 ³³, 29. 12. 1983: 2 ²² (diese alle im Nordwesten
des Mecsek-Gebirges); Kisújbánya, Pásztor-Quelle, 8. 10. 1983: 11 ²², 5 ³³, 15.
10. 1983: 8 ²², 5 ³³, 16. 10. 1983: 2 ²², 2 ³³, 28. 10. 1983: 10 ²², 7 ³³, 27. 11.
1983: 23 ²², 3 ³³, 7. 12. 1983: 14 ²², 1³, 10. 12. 1983: 8 ²², 17. 12. 1983: 25 ²²,
26. 12. 1983: 7 ²², 1³, 1. 1. 1984: 6 ²², 6. 1. 1984: 3²; Vékény, Tal “Vár-völgy”, 8.
10. 1983: 9 ²², 5 ³³, 15. 10. 1983: 3 ²², 1³, 28. 10. 1983: 9²², 8 ³³, 27. 11. 1983:
12²², 7. 12. 1983: 7 ²², 2 ³³, 17. 12. 1983: 4²², 1³, 1. 1. 1984: 1 ², 1³; Pécs-Vasas,
Hármasbükk-Quelle, 30. 12. 1983: 2 ²² (diese alle im östlichen Mecsek-Gebirge,
etwa 20 km von den ersten Orten entfernt). Die Tiere wurden gesammelt von
Sára U. Nógrádi, A. Uherkovich und G. Vágner und befinden sich in coll. Janus
Pannonius Museum, Pécs, coll. Naturwissenschaftliches Museum Budapest, coll.
Ujhelyi, Budapest, und coll. Malicky.”
Chaetopteryx rugulosa mecsekensis Nógrádi, 1986: Malicky (2005b: 573):
transferred from C. schmidi to C. rugulosa.
Chaetopteryx mecsekensis Nógrádi, 1986: Oláh et al. (2012: 59–60): raised
to species status. “Hungary: Mecsek Mts. Pécs, Meleg-mány, 22.X.1983, leg. S.
Nógrádi (5², 5³, OPC). 1.XI.1985, leg. S. Nógrádi (5², 3³, OPC). 20.XII.1983,
leg. Á. Uherkovich (1², 3³, OPC). 28.X.1987, leg. Á. Uherkovich (10², 5³, OPC).
Mecsek Mts. Kisújbánya [Hosszúhetény], Pásztor-spring, 436m, N 46°13’04”,
E 18°21’27”, 12.XI.1986, leg. S. Nógrádi (3², 1³, OPC); 10.XII.1983, leg. S.
Uherkovich (4², OPC); 26.XII.1983, leg. S. Nógrádi (4², 1³, OPC); 1.I.1984, leg.
S. Nógrádi (3², OPC). 7.XII.1983, leg. Á. Uherkovich (7², 1³, OPC); 6.XI.2009,
leg. I. Szivák & Á. Uherkovich (3²,1³, OPC); 5.XI.2010, singled leg. I. Szivák, J.
Oláh & Á. Uherkovich (11², 11³, OPC). Mecsek Mts. Hosszúhetény, Takanyó-v,
24.X.1984, leg. Á. Uherkovich (2², 2³, OPC); Mecsek Mts. Hosszúhetény,
Hidasi-v. Csurgó, 11.XI.1984, leg. Á. Uherkovich (2², 2³, OPC); Mecsek Mts.
Mánfa, Kőlyuk, 12.XI.1985, leg. S. Nógrádi (8², 1³, OPC). Vékény, Vár-v. Iharos-
spring, 28.X.1983, leg. S. Nógrádi (3², 2³, OPC). 12.XI.1986, leg. S. Nógrádi
(4², 1³, OPC). Magyaregregy, Iharos-kút, 327m, N46°13’21.90’’, E18°20’06.80’’,
6.XI.2009, leg. I. Szivák & Á.Uherkovich (2², OPC). Magyaregregy, Máré-
forrás, N46°13’39.98’’, E18°19’19.39’’, 6.XI.2009, leg. I. Szivák & Á.Uherkovich
(1², 1³ in copula, OPC). Mecsek Mts. Pécs, Nagy-Mély-völgy, Kánya-forrás,
347m, N46°08’05.16’’, E18°12’43.75’’, 14.XI.2009, leg. I. Szivák (3², 1³, OPC).
Mecsek Mts. Pécs, Nagy-Mély-völgy, Sziklás-forrás, N46°08’26.7’’, E18°12’39.96’’,
14.XI.2009, leg. I. Szivák (3², OPC). Mecsek Mts. Pécs, Melegmányi-völgy, Anyák-
kútja, N46°08’08.55’’, E18°13’31.46’’, 14.XI.2009, leg. I. Szivák (2², 1³, OPC).
Mecsek Mts. Pécs, Melegmányi-völgy, Mésztufa lépcső, 352m, N46°08’12.89’’,

194 J. Oláh
E18°13’29.92’’, 14.XI.2009, leg. I. Szivák (3², OPC). Mánfa, Nagy-Mély-völgy,

Cserkész-forrás, N46°08’56.73’’, E18°12’38.08’’, 14.XI.2009, leg. I. Szivák (4²,
OPC). Magyaregregy, Vár-v. Réka-spring, N 46°13’39”, E 18°19’19” 6. XI. 2011,
leg. S. Nógrádi & Á. Uherkovich (7²,3³, OPC).
Distinguishing traits – Oláh et al. (2012: 59–60): “Male: cerci higher than at
C. schmidi and C. papukensis; apical flap of gonopod less developed, resulting in
gonopod apex rounded in lateral view, not with pointed or projected blunt apex
like at C. schmidi and C. papukensis; dorsal hook of paraproct medium turned, less
turned at C. papukensis, highly turned at C. schmidi; aedeagus with long, almost
filiform lateral processes, not gemmiform of C. schmidi or short digitiform of C.
papukensis; paramere shaft short rod-shaped with medium long straight primary
spine, not short straight of C. schmidi or long curved of C. papukensis. Female:
anal tube medium long; supragenital plate very sharp triangular in ventral view.”
Chaetopteryx papukensis Oláh et Szivák, 2012

Chaetopteryx rugulosa mecsekensis Nógrádi, 1986: Oláh (2010: 98): misi-
dentification.
Chaetopteryx papukensis Oláh et Szivák, 2012 in Oláh et al. (2012: 60–62):
Holotype. Croatia: Papuk Mts, Slatinski Drenovac, Jankovac, Jankovac spring,
45°31’08.1”, 17°41’11.9”, 510 m, 06.11.2012, T. Kovács, G. Magos (1², OPC).
Allotype. Same as holotype (1³ OPC). Paratypes. Same as holotype (14², 10³
OPC, 5², 3³ MM). Krndija Mts, 3 km N of Kutjevo, Velika rijeka, small trib-
utary, 424 m (YL23), N45°27’55”, E17°52’37”, 04.11.2011, I. Szivák (1², OPC).
Krndija Mts, 6 km N of Kutjevo, Velika rijeka, springs, 580 m, N45°28’59”,
E17°51’33”, 04.11.2011, I. Szivák, Á. Uherkovich (4², 4³ OPC); 06.11.2012,
T. Kovács, G. Magos (9², 9³ OPC). Krndija Mts, Kutjevo, Mala rijeka, 402 m
(YL23), N45°27’48”, E17°51’53”, 04.11.2011, I. Szivák, Á. Uherkovich (4², 1³
OPC). Papuk Mts, forest brook below the Slatinski Drenovac – Velika road,
45°29’32.4”, 17°39’10.9”, 480 m, 06.11.2012, T. Kovács, G. Magos (7², 10³ OPC).
Papuk Mts, Jankovac, 13.10.1986, B. Horvat, I. Sivec (2², 2³ PMS). Papuk Mts,
Jankovac spring, cave and the surrounding beech forest, 456 m, N45°31.126’,
E17°41.198’, 01.10.2007, L. Dányi, J. Kontschán, D. Murányi (7², HNHM).
Papuk Mts, Slatinski Drenovac, 1.5 km S, Jankovac stream, 350 m, N45°32’01”,
E17°42’08”, 19.10.2012, Á. Uherkovich (1³ OPC). Papuk Mts, Slatinski Drenovac,
Jankovac, Jankovački potok, 351 m (YL14), N45°31’31”, E17°41’25”, 04.11.2011,
I. Szivák (3², 1³ OPC). Papuk Mts, Slatinski Drenovac, Kovačica Potok, 541 m,
N45°31’08”, E17°39’54”, 03.11.2012, Á. Uherkovich (1², OPC).”
Chaetopteryx papukensis Oláh et Szivák, 2012: Malicky (2014: 52): syno-
nymised with C. schmidi.

Chaetopteryx papukensis Oláh et Szivák 2012: Oláh et al. (2015b: 83–84):

“Here we reinstate the specific status of Chaetopteryx papukensis Oláh & Szivák
2012 stat. restit.”
Diagnosis – Oláh et al. (2012: 60–62): “This new species belongs to the C.
schmidi New Species Subgroup of the C. rugulosa species group. Most close to C.
mecsekensis but differs by having cerci lower than at C. mecsekensis, apical flap of
gonopod present, not lacking; the well developed flap producing gonopod apex
with pointed or blunt projection in lateral view, not rounded like at C. mecseken-
sis; dorsal hook of paraproct less turned, medium turned at C. mecsekensis, highly
turned at C. schmidi; aedeagus with medium digitiform lateral processes, not
long filiform of C. mecsekensis or gemmiform of C. schmidi; paramere shaft trian-
gular, not digitate; primary spine long curved, not short straight of C. schmidi or
long straight of C. mecsekensis. Anal tube of female is long.”
Etymology – “The new species is named after the Papuk Mts, where the type
locality is found.”
Chaetopteryx psunjensis Oláh, 2015

Chaetopteryx papukensis Oláh et Szivák, 2012 in Oláh et al. (2012: 62):
misidentification.
Chaetopteryx psunjensis Oláh, 2015 in Oláh et al. (2015b: 84–86): “Holotype:
Croatia: Sumetlica Strmac, Psunj Mts. Creek on sandstone, 663m, 45˚22’32”N,
17˚21’40”E, 23.X.2012, leg. Á. Uherkovich (1², OPC). Allotype: Croatia,
Sumetlica Strmac, Psunj Mts. Small creek on crystalline rock, 722m, 45˚22’43”N,
17˚22’04”E, 23.X.2012, leg. Á. Uherkovich (1 female, OPC). Paratype: same as
allotype (1 female, OPC).”
Diagnosis – Oláh et al. (2015b: 84–86): “This new species has elongated
dorsal branch of the paraproct with blunt apex. The lateral process of the ae-
deagus is enforces by a short double S-shaped ridge connected directly to the
basal sclerotized tube of the lateral process. Such a sclerotized basal tube has
been evolved in C. kamnikensis of the C. rugulosa subgroup. This unique struc-
tural modification of the simple sclerotized cylinder of the aedeagus is an adap-
tive product of the sexual selection together with the modification of the apical
hook formation on the paraproct. The dorsal profile difference in the female
anal tube between the allotype and paratype is resulted probably by the long
coupling pressure of anchoring paraproctal apical hook on the internal scler-
ites, as anchor substrate during copulation. During copulation the hook pair is
inserted into the membranous anterior margin of the internal sclerite inside the
anal tube exerting pressure to move the entire internal sclerite backward. The
hook and sclerite interlocking keeps the male and female together for several

196 J. Oláh
days may move the internal sclerite backward exposing the bilobed very sclero-
tized apical part more free as well as slandering and elongating the entire anal
tube accordingly.”
the type material was collected.”
Chaetopteryx schmidi Botoşăneanu, 1957

Chaetopteryx schmidi Botoşăneanu, 1957b: 191–193: “15.XII.1956, pet-
it ruisseau affluent gauche du torrent Plavisevitza a environ 8 km en amont de
sa confluence avec le Danube (Banat, région du Défilé de Cazane) 8 ², en al-
cool. Holotype ² dans les collections du Rijksmuseum van Natuurlijke Historie
de Leyde). Paratypes ² dans les collections F. C. J. Fischer, D. E. Kimmins, F.
Schmid, ainsi que dans celle auteur.”
Chaetopteryx rugulosa schmidi Botosaneanu, 1957: Malicky (2005b: 573):
reduced to subspecies status.
Chaetopteryx schmidi Botosaneanu, 1957: Oláh et al. (2012: 62–63): resur-
rected to species status.
Chaetopteryx schmidi Botosaneanu, 1957: Kučinić et al. (2013: 21–22):
species status confirmed.
Chaetopteryx schmidi Botosaneanu, 1957: Oláh et al. (2015b: 87–88):
“Published material (Oláh et al. 2012) is re-examined and diverged fine structure
elements redrawn to compare them with C. papukensis.”
Chaetopteryx tompa Oláh, 2015

Chaetopteryx schmidi Botoşăneanu, 1957: Oláh et al. (2012: 62) (partim):
misidentification.
Chaetopteryx tompa Oláh, 2015 in Oláh et al. (2015b: 88): “Holotype:
Bosnia-Hercegovina: Skender, vakuf, 820m, 19.X.1990, leg. B. Horvát & I.
Sivec, (1 male, OPC). Allotype: same as holotype (1 female, OPC).”
Diagnosis – Oláh et al. (2015b: 88): “This new species is close to C. balcanica
sp. n. but differs by having paramere shaft triangular, not digitate; primary spine
short, stout and straight, not long and curved; dorsal branch of the paraproct,
the apical hook formation is different, longer with blunt apex; female anal tube
longer and not quadrangular. A more detailed examination on several specimens
from more population is required to establish its relations.”
Etymology – “Tompa from “tompa” blunt in Hungarian, refers to the blunt
apex of the hook formation on the dorsal branch of the paraproct.”

Chaetopteryx rugulosa species subgroup

Chaetopteryx noricum species cluster
Chaetopteryx noricum Malicky, 1976
Chaetopteryx schmidi noricum Malicky, 1976: 98–99: “Holotypus ²,
Allotypus ³ und Paratypen: Carinthia, Saualpe oberhalb Wieting, Umgebung
der Weisbürgerhütte, 1600–1800 m, 20.–28. 9. 1975, leg. et coll. Malicky. – Ein
weiteres ² befindet sich him Naturhistorischen Museum Admont, es wurde im
Veitlgraben bei Admont (Steiermark) gefunden und von Strobl (1906: 232) unter
Namen Ch. rugulosa publiziert.”
Chaetopteryx rugulosa noricum Malicky, 1976: Malicky (2005b: 573):
transferred from C. schmidi to C. rugulosa.
Chaetopteryx noricum Malicky, 1976: Oláh et al. (2012: 63): raised to spe-
cies status. “Austria: Carinthia, Saualpe ob Wieting, 1600m, 28.IX.1988, leg H.
Malicky (3³, OPC from MPC). Katschbach, STMKz, 3.XI.2000, leg. W. Graf
(1², OPC). Saualpe, Kliening, middle reach of stream Klieningbach, 932m,
N46°56’49.4’’, E14°46’24.2’’, 21.XI.2009, leg. A. Déry & I. Szivák (1², OPC).
Saualpe, Geierkogel Klippitztörl, springs of stream Klippitzbach, 1584m,
N46°55’53.8’’, E14°40’49.3’’, 21.XI.2009, leg. A. Déry & I. Szivák (5², OPC).
Saualpe, Hinterberg Löllinggraben, a spring in the middle reach of stream
Löllingbach, 802m, N46°54’38’’, E14°34’03’’, 21.XI.2009, leg. A. Déry & I. Szivák
(6², OPC).”
Chaetopteryx pohorjensis Oláh et Urbanič, 2012

Chaetopteryx pohorjensis Oláh et Urbanič, 2012 in Oláh et al. (2012:
63–65): “Holotype. Slovenia, Pohorje Mts, below Pesek, spring area of river
Oplotnica, 1345m, N46°28’24.8’’, E15°20’55.9’’, 08.11.2012, T. Kovács, G. Magos,
I. Sivec (1², OPC). Allotype. Same as holotype (1³, OPC). Paratypes. Same as
holotype (2², 6³, OPC; 1², 1³, MM). Same locality as holotype: 20.10.1981, I.
Sivec (2², 2³, OPC); 15.10.1984, B. Horvat, I. Sivec (16², 9³, PMS); 27.09.2008,
I. Sivec (1², 1³, OPC); 10.11.2008, I. Sivec (3², 1³, OPC); 28.09.2012, I. Sivec
(1², OPC); 19.10.2012, B. Horvat, I. Sivec (18², 7³, OPC); 08.11.2012, I. Sivec
(3², 1³, OPC). Dravograd, Ogleja puša, Vrački stream, 1170 m, N38°08’09”,
E15°05’24,30”, 04.10.2012, I. Sivec, G. Urbanič (4², 1³, OPC). Kamnik,
Volovljek, N46°18’59.7”, E14°42’03.1”, 14.11.2010, I. Sivec (1³, OPC). Kozji Vrh
nad Dravogradom, sidestream of stream Brelejev potok, 1530 m, N46°38’26.35”,
E15°04’52.35”, 04.10.2012, I. Sivec, G. Urbanič (3², OPC). Kozji Vrh nad
Dravogradom, sidestream of stream Velka, 570 m, N46°37’26.0”, E15°04’16.4”,
04.10.2012, I. Sivec, G. Urbanič (7², OPC); 07.11.2012, G. Urbanič (1³, labo-

198 J. Oláh
ratory reared, OPC). Kozji Vrh nad Dravogradom, stream Brelejev potok, 1170
m, N46°38’39.25”, E15°02’52.41”, 04.10.2012, I. Sivec, G. Urbanič (5², 1³, OPC).
Litija, Konjski graben, 16.10.1985, I. Sivec (1³, PMS). Mislinja, Mislinjski jarek,
29.10.2010, I. Sivec (2², 2³, OPC). Pohorje Mts, Pesek, 28.10.1989, B. Horvat, I.
Sivec (1², PMS). Pohorje Mts, Pesek, N46°28’26.3” E15°20’55.9”, 09.10.2010, I.
Sivec (2², OPC).”
Chaetopteryx pohorjensis Oláh et Urbanič, 2012: Malicky (2014: 52): dubi-
ous remarks on species status and synonymy with “Chaetopteryx rugulosa nori-
cum” are presented without any factual arguments or nomenclaturial acts.
Chaetopteryx pohorjensis Oláh et Urbanič, 2012: Oláh et al. (2015b: 89–
90): “Malicky (2014) has not synonymized C. pohorjensis with C. noricum, but
his position was uncertain and confusing, moreover he maintained without any
explanation that C. noricum is a subspecies of C. rugulosa and C. pohorjensis does
certainly not merit a specific name. Above we have summarised the older diver-
gences in the C. rugulosa species group forming the subgroup and species cluster
structures as was detailed earlier (Oláh et al.. 2012). Subgroups and cluster struc-
tures are differentiated by neutral traits of periphallic organs of older divergenc-
es, by gross phallic structures and by specific divergences in the speciation trait of
the lateral process on the aedeagus. C. noricum species cluster is diverged from C.
rugulosa species cluster by having entirely different aedeagus with large inflated
and rigid flexible lateral processes enforsed and supported by a pair of heavily
sclerotized ventral flanges. At higher magnification this type of aedeagus seems
clearly diverged far from the species of the C. rugulosa cluster. This magnitude
of shape divergence may realize dramatic changes in copulatory processes and
mating signals. C. pohorjensis compared to C. noricum, as detailed in the original
species description, has well diverged paramere shaft and paramere spine pat-
tern, very enlarged tube of lateral processes supported by a short, heavily sclero-
tized pair of ventral flanges. Moreover the ventral lip of the female anal tube of
C. pohorjensis is short, not long. The female of C. noricum has diverged signifi-
cantly from all members of the entire species group by having elongated ventral
lip on female anal tube. It is unique for the entire species group that the ventral
lip is longer than the dorsal lip. This old divergence of the ventral lip is stable and
well visible under lower magnification. The elongated ventral lip of C. noricum is
highly sclerotized, as usual in all the females of the C. rugulosa species group, its
distinct divergence from C. pohorjensis is easily recognised at first glance.”
Diagnosis – Oláh et al. (2012: 63–65): “Described and drawn, but not
named by Malicky et al. (1986). Failed to relate it clearly to any of the known taxa.
This new species belongs to the Chaetopteryx rugulosa species group, C. rugulosa
subgroup and C. noricum species cluster. Close to C. noricum sp. n. but differs by

having paraproct wide and angled laterad, not narrow in caudal view; paramere
shaft shorter than at C. noricum; number of paramere spines usually 3; position
of paramere spines nested, not with a tendency to be arranged in horizontal row
with laterad located primary spine and gradually mesad shortening secondary
spines. There are significant differences in the genital structures of the females:
the lower lip of the anal tube is shorter than the upper lip, not equal as at the C.
noricum; the dorsal apical profile of the anal tube characterized by deep V-shaped
excision due to the highly protruded position of the internal scerites.”
Etymology – “The new species is named after the Pohorje Mts, where the
type locality is found.”
Chaetopteryx rugulosa species cluster

Chaetopteryx kamnikensis Oláh et Urbanič, 2012
Chaetopteryx kamnikensis Oláh et Urbanič, 2012 in Oláh et al. (2012: 66–
68): “Holotype. Slovenia: Tržič, Bistrica, Blajšnica stream, 689 m, N46°21’50.02”,
E14°16’55.70”, 03.12.2011, A. Déry, I. Szivák (1², OPC). Allotype. Same as holo-
type (1³, OPC). Paratypes. Same as holotype (1², OPC). Same locality as holo-
type: 13.10.2011, A. Déry, I. Szivák (1², OPC). Dolž, Gorjanci, stream Klampfer,
660 m, 25.10.1990 B. Horvat, I. Sivec (1³, OPC). Golovec, brooklet near Rakovnik
distinct (Ljubljana), 335 m, N46°02’27.49”, E14°31’46.12”, 05.12.2011., A. Déry, I.
Szivák (4², 1³, OPC); 08.11.2012, T. Kovács, G. Magos (1², 1³, MM). Ig. Želimlje,
potok Želimeljščica, 330 m, N45°53’35”, E14°35’43”, 27.10.1989, B. Horvat, I.
Sivec (1³, PMS). Kamniške alpe, [rna pri Kamniku, Volovljek, 1016 m, N46°24’50”,
E14°54’10”, 26.10.2012, B. Horvat, I. Sivec (10:, 4³, OPC); 1028 m, N46°16’13,5”,
E14°41’20,8”, 26.10.2012, B. Horvat, I. Sivec (11:, 6³, OPC). Litija, Janče, stream
Gostinca, 350 m, N46°03’39.19”, E14°40’48.46”, 12.10.2012, G. Urbanič (2², 1³,
OPC). Rakovnik at Ljubljana, 20.10.1983, C. Krušnik (l², 1³, MPC). Šklendrovec
Podkum, stream Šklendrovec, 493 m, N46°04’52.6”, E15°01’10.5”, 25.10.2012, B.
Horvat, I. Sivec (2², 2³, OPC). Tržič, Brezje at Tržič, stream Blajšnica, 646 m,
N46°21’46.06”, E14°17’00.31”, 16.11.2012, B. Horvat, I. Sivec (4², 5³, OPC).
Tržič, Grahovše, potok Lomščica, 860 m, N45°22’00”, E14°22’03”, 01.10.1990,
B. Horvat, I. Sivec (1³, PMS). Tržič, Hudi Graben, stream Hudi Graben, 683 m,
N46°21’41.35”, E14°15’46.64”, 16.11.2012, B. Horvat, I. Sivec (1², 1³, OPC).
Chaetopteryx kamnikensis Oláh et Urbanič, 2012: Malicky (2014: 52): du-
bious remarks on the species status are presented indirectly without any factual
arguments or nomenclaturial acts.
Chaetopteryx kamnikensis Oláh et Urbanič, 2012: Oláh et al. (2015b: 90):
“Malicky (2014) has not synonymized this species, but his position was vague

200 J. Oláh
or even confused. Here we do not detail our original description and diagnosis,
simply we repeat that the divergence of paramere spine pattern and the female
anal tube distinguish this species both from C. prealpensis and from C. rugulosa.
Moreover the lateral process is not platiform- shaped, the platiform process is
characteristic for C. prealpensis. It is digitiform like the lateral process of C. ru-
gulosa, but with the evolution of the very specific sclerotized basal tube lacking
at C. rugulosa. This is why the oblique direction of the process is fixed at C. kam-
nikensis does not depend on the erection state of the endophallus. Without this
supporting sclerotized basal tube on the lateral process of C. rugulosa, the posi-
tion or oblique direction of the lateral process depends on the erection state of
the endophallus.”
Diagnosis – Oláh et al. (2012: 66–68): “Described and drawn, but not
named by Malicky et al. (1986). Due to insufficient material failed to relate it
clearly to any of the known taxa. This new species belongs to the Chaetopteryx ru-
gulosa species group, C. rugulosa subgroup and C. rugulosa species cluster. Close
to C. prealpensis sp. n. but differs by having subapical lateral processes on the
aedeagus digitiform, not platiform; 5–6 parameter spines present and gradually
decreasing in length from apicad to subapicad in sagittal plane, not 2–3 spines
nested. The anal tube of the female with rounded apical lobes and rounded mesal
excision formed by the tergite IX, not triangular and created by the protruded
segment X.”
Etymology – “The new species is named after the town Kamnik, the type
locality is not far away.”
Chaetopteryx prealpensis Oláh, 2012

Chaetopteryx prealpensis Oláh, 2012 in Oláh et al. (2012: 68–71):
“Holotype. Hungary: Kőszeg Mts, Hörmann-forrás, 18.10.1986, Á. Uherkovich
(1², OPC). Allotype. Same as holotype (1³, OPC). Paratypes. Same as holo-
type (4², 2³, OPC; 1², 1³, MM). Austria: Ausserneuwald, stream, 817 m,
N47°34’01.9”, E16°01’10.5”, 19.11.2009, A. Déry, I. Szivák (1², 1³, ²³ in copu-
la, OPC). Gleinalpe, GH Krautwaschl & Gleinalm Sulmhütte, 1100–1300 m,
08.10.2012, D. Stradner (1², 2³, OPC). Hochegg bei Grimmenstein, spring
and its outlet, 621 m, N47°36’44”, E16°05’52.7”, 19.11.2009, A. Déry, I. Szivák
(3³, OPC). Koralpe, Handalm, springs near Gösler Hütte (Weinebene), 1784
m, N46°50’35.89”, E15°01’18.53”, 21.10.2012, J. Oláh, I. Szivák (13², 7³, OPC).
Koralpe, St. Oswald, Wildbach, 30.09.2007, D. Stradner (4², 2³, OPC). Lafnitz
Quelle, 16.10.2012, W. Graf (1², 1³, OPC). Mitterneuwald, Hermann spring,
956 m, N47°32’56.3”, E15°58’56.1”, 19.11.2009, A. Déry, I. Szivák (2², OPC).
Packalpe, spring near Knödelhütte, 1440 m, N46°59’31.20”, E14°56’20.02”,

20.10.2012, J. Oláh, I. Szivák (2², 3³, OPC). Sommeralm, upper reach of stream
Mixnitz Bach, 1327 m, N47°20’57”, E15°32’56.5”, 20.11.2009, A. Déry, I. Szivák
(1², 1³, ²³ in copula, OPC). Styria, 14 km above restaurant Krautwaschl,
Gleinalm, N47°12’16”, E15°08’47”, 1187 m, 25.09.2011, D. Stradner (2², OPC).
Styria, NW Stainz, near Marhof, N46°54’, E15°13’, 10.11.2006, W. Graf (1³, OPC).
Croatia: Ivanščica Mts, Potok Slugovina, 15.12.2002, K. Žganec (1³, OPC).
Medvednica Mts, Bliznec, pilana, stream, 09.12.2009, M. Kučinić (1³, OPC).
Medvednica Mts, Izvor Mrzlak, 18.11.2006, A. Popijač (3², 2³, OPC). 18.11.2008,
A. Popijač (1², 1³, OPC). Medvednica Mts, Kraljičin Zdenac, 19.11.2009, M.
Kučinić, I. Vučkavić (1³, OPC). Medvednica Mts, Veliki Potok, N45°51’28.52”
E15°56’08.19”, 18.10.2011, A. Previšić (1², OPC). Žumberačka Mts, small stream
near River Slapnica, 03.11.2012, M. Kučinić (2², 1³, OPC). Žumberačka Mts,
Vlašić Brdo, River Slapnica, N45°42’35.7”, E15°29’40.1”, 215 m, 07.11.2012, T.
Kovács, G. Magos (1², 1³, OPC). Žumberačka Mts, Žumberak, River Slapnica,
28.10.2009, M. Kučinić (1³, OPC). Hungary² Kőszeg Mts, Hörmann-forrás, 694
m, N47°27’34.2”, E16°27’34.2”, 18.11.2009, I. Szivák (2², 1³, OPC). Kőszeg Mts,
Stájer-házak, 05.10.1991, Á. Uherkovich (2², OPC); 18.10.1986, Á. Uherkovich
(1², 1³, OPC). Velem, Borha-forrás, 04.11.1984, S. Nógrádi (1², 1³, OPC).
Slovenia: Brdo, Kranj, brooklet to the pond IX, 14.10.2003, G. Urbanič (2²,
PMS). Kališe, Črna pri Kamniku, 13.10.1990, I. SIVEC (1², 3³, PMS). Kamniška
Bistrica, 27.11.1969, B. Horvat, I. Sivec (1², 1³, PMS). Kamniške alpe, [rna pri
Kamniku, Volovljek, 1016 m, N46°24’50”, E14°54’10”, 04.10.2012, I. Sivec, G.
Urbanič (3², 3³, OPC). Kozje, stream Bistri graben, 01.10.1986, B. Horvat, I.
Sivec (1², PMS). Ljubno, Smrekovec, potok Pod Krumpaško Planino, 1390 m,
10.09.1997, B. Horvat, I. Sivec (1³, PMS). Ljubno, Smrekovec, potok Robanšek,
Pod Komnom, 1200 m, N46°24’41”, E14°51’02”, 25.09.1997, B. Horvat, I. Sivec
(1², PMS). Ljubno, Smrekovec, stream below Kugovnik, N46°25’13”, E14°52’20”,
1450 m, 25.09.1997, B. Horvat, I. Sivec (2², PMS). Lukovica, Trnjava, stream
Drtijščica, 340 m, 12.11.1996, B. Horvat, I. Sivec (1², PMS). Moravče, Vinje,
stream Drtijščica, 360 m, 12.11.1996, B. Horvat, I. Sivec (4², 1³, PMS). Pečice,
Bre¸ice, stream Močnik, 15.10.1988, B. Horvat, I. Sivec (1², 1³, PMS). Pohorje
Mts, brooklet near Rogla, 1350 m, N46.448280°, E15.339671°, 03.12.2011, A.
Déry, I. Szivák (1², 1³, OPC). Pohorje Mts, brooklet near Sne¸inka (Rogla), 1097
m, 46.435143 N, 15.368489 E, 03.12.2011., A. Déry, I. Szivák (3², 1³). Pohorje
Mts, Padeški vrh, source of Gradiški graben, 1020 m, N46°25’54.1”, E15°22’18.0”,
08.11.2012, T. Kovács, G. Magos, I. Sivec (1², OPC). Pohorje Mts, Pesek,
N46°28’26.3”, E15°20’55.9”, 10.11.2008, I. Sivec (1³, OPC); 12.09.2009, I. Sivec
(1³, OPC); 09.10.2010, I. Sivec (1², OPC). Pohorje Mts, Vel. Vrh, Osankarica,
1300 m, 04.11.1984, D. Šere (1², 1³, PMS). Pri koritu Ob Litijski cesti, 50 m, pod

202 J. Oláh
hišo Sp. Besnica 1, 25.11.1984, B. Horvat, I. Sivec (2², PMS). Smrekovec Mts,
below Krumpaška planina, 1390 m, N46°24’50”, E14°54’10”, 26.09.2012, I. Sivec,
G. Urbanič (4², OPC); 05.10.2012, B. Horvat, I. Sivec (2², 3³, OPC). Smrekovec
Mts, Tračka Planina, source of stream ˜ep, 11.09.1987, B. Horvat, I. Sivec (1³,
PMS). Tepe, Zagorje ob Savi, 16.10.1985, B. Horvat, I. Sivec (1³, PMS). Zg.
Velka, spring of the Ščavnica River, 01.10.1998, G. Urbanič (1², 1³pupae, PMS);
01.12.1998, G. Urbanič (1², PMS).”
Chaetopteryx prealpensis Kolenati, 1848: Malicky (2014: 52): synonymised
with C. rugulosa.
Chaetopteryx prealpensis Oláh, 2012: Oláh et al. (2015b: 90–91): species
state reinstated. “This species with very large distributional area is the putative
ancestral species of the C. rugulosa species cluster. Malicky (2014) has syno-
nymized C. prealpensis with C. rugulosa, declaring that the lateral processes are
variable in their length, only slightly sclerotized, and may easily be deformed
during maceration. In 2014 we have resampled three more populations in the
Kőszeg Mts. and one population in the Őrség NP in Hungary and re-exam-
ined with care and sophistication our old materials from Croatia, Slovenia and
Austria (population details in Oláh et al. 2012). We have found, as earlier, the
opposite what Malicky wrote. The lateral process is very stable and consistent-
ly structured in so many populations from the entire large distributional area.
It is sagittaly flat, not digitiform like at C. rugulosa or gemmiform like at C.
zalaensis, it is vertically plate-shaped or platiform (a terminus technicus stand-
ardization following the terms of digitiform and gemmiform). It is natural that
there are individual variations, no two animals are identical even in the diverged
adaptive traits, but the platiform shape is in the range of the basic architecture.
The process is not much sclerotized, but seems rather rigid. At least if we com-
pare the lateral view of the process drawn in the diverged trait matrix it is well
visible that the different specimens have differently erected endophallus, but
the shape of the lateral process is similar, stable. The divergence of the lateral
process represents the speciation trait evolved in sexual selection processes and
manisfests itself as a possible reproductive barrier in prezygotic phase realized
by cryptic female choice, sperm competion or by other unknown mechanisms.
Based on this divergence we reinstate the incipient phylogenetic species state of
Chaetopteryx prealpensis.”
Diagnosis – Oláh et al. (2012: 68–71): “Specimens of this widely distribut-
ed species collected from several populations in Austria, Bosnia & Herzegovina,
Croatia, Hungary and Slovenia formerly were determined as C. rugulosa.
However it clearly differs from the holotype of C. rugulosa Kolenati, 1948. This
new species belongs to the Chaetopteryx rugulosa species group, C. rugulosa sub-
group and C. rugulosa species cluster. Close to C. rugulosa Kolenati, 1848 but dif-

fers by having subapical lateral processes on the aedeagus platiform and directed
oblique upward, not digitiform and not horizontal. The anal tube of the female
broad, not slender.
Variability – Oláh et al. (2012: 68–71): “Similarly to most species in Chae-
topteryx rugulosa species group the non-intromittent periphallic structures, the
cerci, the paraprocts and the gonopods are highly variable. The number and
length of paramere spines are less variable; the spine pattern varies especially in
peripherial area with two tendencies: (1) reducing spine number down to the sin-
gle primary spine that is accompanied by 1–2 very short, almost tertiary spines;
(2) shortening the primary spine with increasing number of secondary spines up
to 3–4. The intromittent part of the phallic organ that is the aedeagus and espe-
cially its head with the spatulate, platiform lateral processes is rather stable even
in populations of peripherial area: Zumberacka Mts in Croatia and Kőszeg Mts.
in Hungary.”
Etymology – “The new species is named after the Prealpine region, where
this widely distributed ancestral species lives.”
Chaetopteryx rugulosa Kolenati, 1848

Chaetopteryx rugulosa Kolenati, 1848: 73: “Habitat in Dalmatia (Stenz!).”
Chaetopteryx rugulosa Kolenati, 1848: Malicky et al. (1986: 3–5): holo-
type redrawn. Locus typicus uncertain: “Da die Art sonst unseres Wissens
nie in Dalmatien gefunden wurde, könnte man an der Richtigkeit der
Herkunftsbezeichung zweifeln. In der erste Halfte des vorigen Jahrhunderts
nahm man es mit der Etikettierung nicht so genau”
Chaetopteryx rugulosa Kolenati, 1848: Oláh et al. (2012: 71): “Austria:
Gleinalpe, springs and springbrook 1.4 km above restaurant Krautwaschl,
1172 m, N47°12’15.31”, E15°08’22.14”, 22.10.2012, J. Oláh, I. Szivák (3² 6³,
OPC). Plenzengreith, upper reach of stream Schöcklbach, 954 m, N47°12’37.2”,
E15°29’00.8”, 20.11.2009, A. Déry, I. Szivák (2² 1³, OPC). Stiftingtal, Graz,
19.10.1998, W. Graf (1² 1³, OPC); 25.09.2005, W. Graf (3², 1³, 7 pupae, OPC);
10.2006, W. Graf (6², 8³, OPC; 2², 1³, MM).”
Chaetopteryx rugulosa Kolenati, 1848: Malicky (2014: 52): contradict-
ing to his earlier statement (Malicky et al. 1986), the author now argues “that
Chaetopteryx rugulosa was described from Dalmatia”.
Chaetopteryx rugulosa Kolenati, 1848: Oláh et al. (2015b: 91–93): “Austria,
Graz, Stiftingtal stream tributaries, 25. X. 2014, leg. M. Máté, D. Stradner, J.
Oláh & M. Oláh (17 male, 10 females; OPC). Hungary: Őrség National Park,
Kétvölgy, N 46°53’ 12.41” E 16°13’ 42.00”, 30. X. 2014, leg. M. Máté (3 males, 3
females, OPC).”

204 J. Oláh
Remarks – Oláh et al. (2012: 71): “Specimens from Austria, Bosnia &
Herzegovina, Croatia, Hungary and Slovenia were determined as C. rugulosa,
however we have found that specimens from only a very small area in Austria has
the aedeagal structure identical with the aedeagus of the holotype, other speci-
mens belong to four species: C. schmidi from Bosnia & Herzegovina, C. kam-
nikensis sp. n. from Slovenia, C. zalaensis sp. n. from Hungary and to the widely
distributed C. prealpensis sp. n.”
Diagnosis – Oláh et al. (2015b: 91–93): “The holotype of C. rugulosa has
slender digitiform lateral process (Malicky et al. 1986). In 2014 we have exam-
ined several populations newly sampled in side valleys of the main Stift ingtal
valley near Graz, near the headwater regions of the River Rába. In our new
collection trial we have found a small C. rugulosa population also in the Őrség
NP in Hungary again in a small spring stream belonging to the system of River
Rába. In diverged trait matrix we have presented diagrammatic lateral drawings
of aedeagus with the lateral process and with the endophallus of 22 specimens
representing one Hungarian and 8 Austrian populations. The finger-like lateral
process of C. rugulosa is consistently different from the vertically flat, plate-
shaped, platiform lateral process of C. prealpensis. Moreover the lateral process
of C. prealpensis is double or even triple sized. Similarly to C. prealpensis the size
of the lateral process has no significant relationship to the erection state of the
endophallus.
Malicky (2014) emphasized that C. rugulosa was described from Dalmatia
based on Kolenati description: “Habitat in Dalmatia (STENZ!)”. However in
an earlier paper Malicky et al. (1986) have questioned the reality of the habitat
data of the profit oriented insect dealer Stenz: “In der ersten Hälfte des vorigen
Jahrhunderts nahm man es mit der Etikettierung nicht so genau”. We have sam-
pled real Dalmatian costal area in right time and in right habitats several times,
but we have not collected any specimens from the C. rugulosa species group. We
have collected members of this species group in internal mountain ranges in
Bosnia-Herzegovia and Serbia, but all belong to the C. schmidi subgroup. No C.
rugulosa or C. prealpensis live in Dalmatia or even nearby Dalmatia. The holotype
is in good condition, aedeagus perfectly preserved, and all the specimens, with
exatly the same lateral process, was collected in a very restricted area near around
Graz, except the single Hungarian population just discovered.”
Chaetopteryx zalaensis Oláh, 2012

Chaetopteryx zalaensis Oláh, 2012 in Oláh et al. (2012: 71–73): “Holotype.
Hungary: Hegyhátszentjakab, Vadása-tó, források, N46°52’32”, E16°33’03”,
04.11.2010, singled, J. Oláh, Á. Uherkovich (1², OPC). Allotype. Same as hol-

otype (1³, OPC). Paratypes. Same as holotype (15², 11³, OPC, 3², 1³, MM).
Vas Megye, Szőce, 05.11.1985, Á. Uherkovich (12², 7³, OPC); 17.10.1986, Á.
Uherkovich (10³, OPC).”
Chaetopteryx zalaensis Oláh, 2012: Malicky (2014: 52): synonymised with
C. rugulosa. This nomenclatural act was based on putative identity of the lateral
shape of cerci. It was suggested that the stalked shape of cerci at C. zalaensis
is plane dependent: in slightly different view it is rectangular and parallel-sided
similarly to the lateral shape of cerci at C. rugulosa. The cerci are neutral, non-
adaptive trait in the contemporary divergences; highly exposed to various sto-
chastic processesare; therefore rather variable in most member of Chaetopteryx
rugulosa species group. Nevertheless on population level and in proper compara-
ble observational view, the cerci are characterized with stalked lateral shape at C.
zalaensis and with parallel-sided shape at C. rugulosa. Moreover the divergence
between the two species is evident and realised in non-neutral adaptive trait, that
is in the gemmiform (C. zalaensis) and digitiform (C. rugulosa) diverged state of
the lateral process on the aedeagus.
Chaetopteryx zalaensis Oláh, 2012: Oláh et al. (2015b: 93–95): “Hungary,
Örség, Szőce stream, Eastern arm, Biczó springs, N 46°54’ 16.60” E 16°34’ 42.36”
26.X.2014, leg. M. Máté, J. Oláh & M. Oláh (36 males, 11 females; OPC).
Örség, Szőce stream, Eastern arm, Biczó springs, N 46°54’ 16.60” E 16°34’ 42.36”
23.XII.2014, leg. M. Máté, (2 males, 2 females; OPC). Örség, Hegyhátszentjakab,
Vadása-tó Spring, 18.X.2014, leg. M. Máté (9 males, 8 females; OPC). Örség,
Hegyhátszentjakab, Vadása-tó Spring, N 46°52’ 33.37” E 16°33’ 06.13” 23.X.2014,
leg. M. Máté, J. Oláh & M. Oláh (6 males, 5 females; OPC). Örség, Szőce stream,
Dam springs, 26.X.2014, leg. M. Máté, J. Oláh & M. Oláh (7 males, 7 females;
OPC). Örség, Szőce stream, spring at bridge, 26.X.2014, leg. M. Máté, J. Oláh &
M. Oláh (1 male, OPC).” “Malicky (2014) has synonymized C. zalaensis having
gemmiform lateral process with C. rugulosa having long digitiform lateral proc-
ess, repeating again without documentation that the lateral process is variable.
[…] Here we reinstate the specific status of the incipient phylogenetic species
Chaetopteryx zalaensis stat. restit.”
Diagnosis – Oláh et al. (2012: 71–73): “This new species belongs to the
Chaetopteryx rugulosa species group, C. rugulosa subgroup and C. rugulosa spe-
cies cluster. Close to C. rugulosa Kolenati, 1848 but differs by having subapical
lateral processes on the aedeagus short and pointed gemmiform, not long digiti-
form; cerci stalked, not parallel-sided. The anal tube of the female very long and
slender, almost tapering apicad.”
Etymology – “The new species is named after the Zala region, where the type

206 J. Oláh
Chaetopteryx irenae species subgroup

Chaetopteryx clara McLachlan, 1876
Chaetopteryx clara McLachlan, 1876: 197: “Carniola (Schmidt; one ² in
Hagen’s collection); Görz (one ² in the Vienna Museum). Remarkable for its uni-
formly pale colour, and thoroughly distinct in its anal characters, the penis being
nearly obsolete.”
Chaetopteryx clara McLachlan, 1876: Oláh et al. (2012: 73): “Slovenia:
Bormes, stream Grabnarica, 21.10.1995, B. Horvat (1², PMS). Ljubljana, Mostec,
1989, H. Malicky (2², 2³, OPC from MPC). Ljubljana, Mostec, Pržanec stream,
Medvode, Osolnik, 440 m, N46°07’24.5”, E14°20’54.8”, 25.11.2012, I. Sivec (1²,
OPC).”
Notes – Oláh et al. (2012: 73): “Its species group position needs further
studies. The supplementary digitiform processes on the superanal complex is ves-
tigial. Paraproct spine pattern rather peculiar; characterized by the presence of
primary, secondary and tertiary spines arranged in anteriad shortening row in
sagittal plane; primary spine slender and undulate; location and number of peg-
like tertiary spines variable.”
Chaetopteryx euganea Moretti et Malicky, 1986

Chaetopteryx euganea Moretti et Malicky, 1986 in Malicky et al. (1986):
“Untersuchtes Material: Veneto, Fontanella del Mottolo, 130 m, Vo Euganeo,
21. 12. 1967: 2², 1³ (leg. Satori, coll. Moretti): Holotypus ², Allotypus ³, Para-
typus ².”
Chaetopteryx euganea Moretti et Malicky, 1986: Oláh et al. (2012: 73):
“Italy: Colli Euganei, 19.10.1987, H. Malicky (2², 2³, OPC from MPC).”
Chaetopteryx giuliensis Oláh et Kovács, 2012

Chaetopteryx giuliensis Oláh et Kovács, 2012 in Oláh et al. (2012: 73–
75): “Holotype. Italy: Alpi Giulie, Sella Carnizza, spring area of River Uccea,
N46°20’11.4”, E13°19’46.8”, 1105 m, 09.11.2012, T. Kovács, G. Magos (1², OPC).
Allotype. Same as holotype (1³, OPC). Paratypes. Same as holotype (2², OPC;
1², MM). Alpi Giulie, between Uccea and Resia, left side brook of River Uccea,
N46°18’54.2”, E13°23’37.6”, 725 m, 09.11.2012, T. Kovács, G. Magos (1², 1³,
OPC). Sorgente del T. Uccea (1050 m), Parco Naturale delle Prealpi Giulie, Com
Resia prov. Udine, crenal, 09.10.1999, S. Paradisi, F. Stoch (1², OPC).”
Chaetopteryx giuliensis Oláh et Kovács, 2012: Malicky (2014: 52): reduced
the divergence to the number of spines of the parameres between C. giuliensis

and C goricensis. However the basic pattern of paramere spines are the result of
older divergences and may be exposed to significant fluctuating asymmetry of
matching type indicative of developmental instabilities caused by adverse envi-
ronmental condition or by genetic challenges (Oláh et al. 2015b). In this species
group the real divergences have been evolved in the non-neutral, adaptive specia-
tion trait that is in the structure and shape of the aedeagus head.
Chaetopteryx giuliensis Oláh et Kovács, 2012: Oláh et al. (2015b: 95):
“Malicky (2014) has not synonymized these species (giuliensis and idriensis), but
his position was vague. We agree with him that these species are close to C. gori-
censis, but disagree with his other statements: “Except for the usual individual
variability of the structures, the only difference is the number of the spines of
the parameres, both having a small bunch of them”. “C. goricensis normally has
only one spine, but some specimens from the type locality Deskle have two”. The
number of spine-like modified apical setae nested on the tip of the parameres is
stable in the examined 15 specimens of C. idriensis from three populations. There
was a single specimen of C. giuliensis with left paramere having only two well de-
veloped modified setae out of the 6 examined specimens from three populations.
We have found also a specimen of C. goricensis from the type locality having 2
spines on the right paramere, but the second spine was very small vestigial.
As we have explained above in details the pattern of modified paramere
setae are not a contemporary speciation trait in Chaetopteryx rugulosa species
group, not diverging consistently on species level. They exhibit older divergences
at subgroup level. Nevertheless C. goricensis has specific paramere spine pattern
compared to C. giuliensis and C. idriensis. The three species have diverged signifi-
cantly in several neutral traits, we do not list them here, and the differences are
explained in details in the original species descriptions. In the C. rugulosa species
group, as explained before, the speciation trait is the lateral process and the as-
sociated substructures on the aedeagus. The lateral processes are rather rigid and
evolved into completely different shapes in the three species. C. idriensis with
the smallest lateral process has no any sclerotized ridge or flange evolved to sup-
port the function of the lateral process. Simply it is not required; this small proc-
ess may function perfectly without additional support of sclerotized structures.
Here we can realise againg that the divergence of the lateral process, the specia-
tion trait in the building process of reproductive barriers is governed by sophis-
ticated complex cooperation of several quantitative trait loci in concerted evolu-
tion. C. goricensis has larger lateral process and a small sclerotized ventral flange
supporting its function. C. giuliensis evolved an extremely large lateral process in
the form of a vertical subquadrangular plate. Its specific function is supported by
a well developed pair of ventral flange.”

208 J. Oláh
Diagnosis – Oláh et al. (2012: 73–75): “This new species belongs to the
Chaetopteryx rugulosa species group, C. irenae new species subgroup. Close to C.
irenae Krušnik & Malicky, 1986 but differs by having cerci downward directed
ventroapicad; paraproct more slender in apical view; supplementary processes
free, not fused to the paraproctal triangle; gonopods with apical margin less un-
dulate; lateral subapical processes platiform, not digitifom; supporting sclerite
broad, almost semicircular, not long and narrow. Female has apical profile of the
anal tube angulate, not rounded in dorsal view.”
Etymology – “The new species is named after the Alpi Giulie, where the type
Chaetopteryx goricensis Malicky et Krušnik, 1986

Chaetopteryx goricensis Malicky et Krušnik, 1986 in Malicky et al. (1986:
13): “Untersuchtes Material: Holotypus ² und Allotypus ³: Slowenien, Deskle
nördlich von Gorica, an einem kleinen Wiesenbachlein, 25. 10. 1982, leg. Krušnik
& Malicky, in coll. Malicky. Paratypen: 1³ mit den selben Daten sowie 1² vom
selben Platz vom 29. 10. 1983, leg. Krušnik, in coll. Krušnik.”
Chaetopteryx goricensis Malicky et Krušnik, 1986: Oláh et al. (2012:
75): “Slovenia: Ajdovščina, Predmeja, one spring of Lokavšček stream, 695 m,
N45°56’21.8”, E13°52’17.8”, 06.12.2009, A. Déry, I. Szivák (9², OPC; 1², MM).
Ajdovščina, Predmeja, stream Lokavšček, 15.10.1992, B. Horvat, I. Sivec (1²,
PMS). Čekovnik, Blašk, spring, brooklet, 631 m, N45°59’04.6”, E13°58’11.0”,
05.12.2009, A. Déry, I. Szivák (4², OPC). Čekovnik, Hleviše, spring, brooklet,
Deskle, 1986, H. Malicky (2², 2³, OPC).”
Chaetopteryx idriensis Oláh et Urbanič, 2012

Chaetopteryx idriensis Oláh et Urbanič, 2012 in Oláh et al. (2012: 75–
77): “Holotype. Slovenia: Idrijsko hribovje, Čekovnik, Blašk, spring, brooklet,
N45°59’04.6”, E13°58’11.0”, 650 m, 09.11.2012, T. Kovács, G. Magos (1², OPC).
Allotype. Same as holotype (1³, OPC). Paratypes. Same as holotype (2², OPC;
1², MM). Same locality as holotype: 04.12.2011, A. Déry, I. Szivák (10², OPC);
05.12.2009, A. Déry, I. Szivák (4², OPC). Idrijsko hribovje, Čekovnik, Hleviše,
spring, brooklet, 640 m, N45°58’48.18”, E13°59’9.52”, 05.12.2009, A. Déry, I.
Szivák (1², OPC). Idrijsko hribovje, Krekovše, spring, brooklet at the Idrijca
River, N45°59’01.4”, E13°56’57.4”, 460 m, 31.10.2003, G. Urbanič (1³, MPC);
08.10.2002, G. Urbanič (1², MPC).”
Chaetopteryx idriensis Oláh et Kovács, 2012: Malicky (2014: 52): reduced
the divergence to the number of spines of the parameres between C. idriensis and

C goricensis. However, the basic pattern of paramere spines are the result of older
divergences and may be exposed to significant fluctuating asymmetry of match-
ing type indicative of developmental instabilities caused by adverse environmen-
tal condition or by genetic challenges (Oláh et al. 2015a). In this species group
the real divergences have been evolved in the non-neutral, adaptive speciation
trait that is in the structure and shape of the aedeagus head.
Chaetopteryx idriensis Oláh et Kovács, 2012: Oláh et al. (2015b: 95): see
Chaetopteryx giuliensis Oláh et Kovács, 2012.
Diagnosis – Oláh et al. (2012: 75–77): “This new species belongs to the C.
irenae new species subgroup of the Chaetopteryx rugulosa species group. Close
to C. goricensis Malicky & Krušnik, 1986 but differs by having cerci differently
shaped; paraproct broader in apical view; gonopods longer; paramere spines tri-
pled, not single; lateral subapical processes slender, not broad both in lateral and
dorsal view. Female has ventrolateral setose processes differently shaped; inter-
nal sclerite of the anal tube protruding and producing lateral lobes blunt, not
acute triangular; supragenital plate blunt, not pointed.”
Etymology – “The new species is named after town Idrija, located nearby the
type locality.”
Chaetopteryx irenae Krušnik et Malicky, 1986

Chaetopteryx irenae Krušnik et Malicky, 1986 in Malicky et al. (1986:
14): “Untersuchtes Material: Slowenien, Artvize, Izvir Vrtice, 8. 11. 1983: 1²
(Holotypus, leg. & coll. Krušnik).”
Chaetopteryx irenae Krušnik et Malicky, 1986: Oláh et al. (2012: 77):
“Slovenia: Artviže stream Brusnica, 18.10.2000, G. Urbanič (1², reared in the
laboratory, PMS). Misliče, upper reach of Sušica stream, 617 m, N45°37’14.8”,
E14°02’16.7”, 06.12.2009, A. Déry, I. Szivák (13², OPC); 04.12.2011, A. Déry, I.
Szivák (6², 2³, OPC; 2², 1³, MM).”
Chaetopteryx marinkovicae Malicky et Krušnik, 1988

Chaetopteryx marinkovicae Malicky et Krušnik, 1988: 180: “Holotype
² and paratypes (²,³): Yugoslavia, Istria, Kompanj, 20. 10. 1988. Paratypes also
from nearby Rockopolje (²,³) and Uugrini (1³); in the collections of the authors.”
Chaetopteryx marinkovicae Malicky et Krušnik, 1986 in Oláh et al. (2012:
77): “Croatia: Istria, Kompanj, 1989, H. Malicky (2², 2³, OPC). Istria, Kompajn,
Klobasi, N45.39111°, E14.07111°, 14.11.2009, M. Kučinić (4², 9³, OPC; 1²,
1³, MM). Slovenia: Vala Zelenica, Loka, Črni kal, 17.10.1990, leg. G. Urbanič
(2²,1³; PMS).”

210 J. Oláh
Psilopteryx Stein, 1874

Psilopteryx bosniaca Marinković-Gospodnetić, 1970
Psilopteryx bosniaca Marinković-Gospodnetić, 1971a: 83–84: “Central
Bosnia, m, f, the source of the river Stavnja.”
Psilopteryx bosniaca Marinković-Gospodnetić, 1971: Marinković-Gos-
podnetić (1971b: 144): “Southeast Bosnia, 1² 1³, the source of the river Stavnja
near Vares.”
Psilopteryx curviclavatus Botoşăneanu, 1957

Psilopteryx curviclavatus Botoşăneanu 1957a: 64–65: “Ariescheni (Ari-
esch-Tal, Bihar-Gebirge); leg. Mihai Serban, 28. IX. 1954. 1² in Alkohol (Holo-
typus SMF N6).”
Psilopteryx montanus Kumanski, 1968

Psilopteryx montanus Kumanski, 1968a: 216–218: “Fundort: 10.X.1967,
Vitosa-Gebirge, Bojanski-Bach bei der Berghütte “Rodina” 1² und 1³. Holotypus
² und Paratypus ³ (mit getrenntem Abdomen) in der Sammlung des Verfassers
(in Alcohol).”
Psilopteryx schmidi Kumanski, 1970

Psilopteryx schmidi Kumanski, 1970: 277–279: “Fundort: 23.IX.1967,
Rila-Gebirge, Unterer Jakorudsko-See (2191 m Höhe), 1³; 22.X.1968, Rila-
Gebirge, Zirkus “Die sieben Seen”, Abfluss des 5. Sees (2240 m Höhe), 1² und ein
Paar in Copula. Holotypus und Paratypen (in Alcohol) in meiner Sammlung im
Holotypus ² und Paratypus ³ (mit getrenntem Abdomen) in der Sammlung des
Verfassers (in Alcohol) Zoologischen Museums der Bulgarischen Akademie der
Wissenschaften.”
Psilopteryx psorosa species group

Psilopteryx bohemosaxonica species complex
Psilopteryx bohemosaxonica Mey et Botosaneanu, 1985
Psilopteryx psorosa bohemosaxonica Mey et Botosaneanu, 1985: 120: “Holo-
typus ² und Allotypus ³: Erzgebirge, südlich von Cinovec, 12.X.19862 (in coll.
Mey); 4²- und 3³-paratypen vom selben Ort, (1², 1³ in coll. Botosaneanu). Areal:
Erzgebirge, Böhmerwald, (Zwischen Osterzgebirge und Isergebirge existiert offen-
sichtlich eine Verbreitungslücke. In diesem fast 100 km langem Bereich konnte trotz
intensiver Suche 1982 keine Psilopteryx-Population nachgewieisen werden.”

Psilopteryx bohemosaxonica Mey et Botosaneamu, 1985: Oláh et al. (2015b:

98–99): raised to species status. “Holotype and Allotype. Czech Republic,
Erzgebirge, S of Cinovec, 12.X.1982, leg. W. Mey (MFN). Austria, Superior,
Schwarzenberg, 28.X.1987, leg H. Malicky (2 males, 2 females; OPC). Czech
Republic, Southern Bohemia, Sumava Mts. inlet of Laka lake N49°06’30”
E13°19’34”, 1090m, 30. IX. 2010, leg. J. Bojkova (9 males, 4 females, NMPC).
Southern Bohemia, Sumava Mts. inlet of Cerné jezero lake, N49°10’44”
E13°10’51”, 1030m, 11. X. 2007, leg. P. Chvojka (5 males, 3 females, OPC).
Southern Bohemia, Sumava Mts. left tributary of Teplé Vltava river below source,
N48°58’44” E13°33’45”, 1160m, 16. X. 1992, leg. P. Chvojka (3 males, 1 female,
NMPC). Northern Bohemia, Jizerské hory Mts. Straceny potok brook below
Smrk Mt. N50°53’44” E15°14’49”, 680m, 11.X.1989, leg. J. Preisler & P. Vonicka (2
males, 3 females; NMPC). Northern Bohemia, Jizerské hory Mts. left tributary of
Sous reservoir. N50°47’40” E15°19’22”, 800m, 4.XI.1989, leg. J. Chvojka (23 males,
16 females; NMPC). Northern Bohemia, Krkonose (Giant Mts.), left tributary of
Labe (Elbe) river below Labsky vodopad waterfall, N50°46’03” E15°33’12”, 1100m,
11.X.1992, leg. P. Chvojka (6 males, 5 females; NMPC). Easthern Bohemia, Orlické
hory Mts., brook, Zidovsky kout SW Orlické Záhori, N50°16’16” E16°27’03”,
775–870m, 27.X.1994, leg. P. Chvojka (18 males, 12 females; NMPC). Easthern
Bohemia, Orlické hory Mts., brook, NPR Bukacka reserve NW of Serlich Mt.,
N50°20’06” E16°22’31”, 880–970m, 26.X.1994, leg. P. Chvojka (158 males, 4 fe-
males; NMPC). Easthern Bohemia, Orlické hory Mts., brook S of Serlich Mt.,
N50°19’15” E16°22’59”, 870–940m, 24.X.1994, leg. P. Chvojka (9 males, 7 females;
NMPC). Germany: Bayern, Böhmerwald, inlet of Kleiner Arbersee, N49°07’24”
E13°07’14”, 925m, 10.X.2007, leg. P. Chvojka (1 male, 1 female; NMPC). Bayern,
Böhmerwald, inlet of Rachelsee, N48°58’34” E13°24’03”, 925m, 9.X.2007, leg. P.
Chvojka (4 males, 2 females; OPC). Poland: Kudlon Mt. Gorce Mts. 11.XI.1996,
leg. B. Szczęsny (4 males, OPC). Gorce Mts. IX-X. leg. B. Szczęsny (3 males, 3
females; OPC). Babia Gora, West Beskidy Mts. 1964, leg. B. Szczęsny (6 males,
5 females; OPC). Babia Gora, West Beskidy Mts. 1997, leg. B. Szczęsny (2 males,
2 females; OPC). Dolina Pięciu Stawów Polskich (Lengyel-Öt-tó völgy) Roztoki
Spring, the Tatra Mts. 26.X.1985, leg. B. Szczęsny (3 males, 5 females, OPC).
Roztoki Valley, the Tatra Mts. 26.X.1985, leg. B. Szczęsny (5 males, 3 females,
OPC). Slovakia: Banskobystrický region, Poľana Mts, Hriňová, Bystré, spring
brook of Bystrý Stream, N48°37.671’ E19°28.655’, 1200m 8.X.2013, singled leg.
J. Oláh & L. Szél (1 male, OPC). Banskobystrický region, Poľana Mts, Hriňová,
Bystré, spring brook of Bystrý Stream, N48°37.569’ E19°29.261’, 1025m 8.X.2013,
singled leg. J. Oláh & L. Szél (1 male, 1 female; OPC).”
Diagnosis – Oláh et al. (2015b: 98–99): “This sibling incipient species is
characterized by short dorsal branch of paraproct. The short and high subtrian-

212 J. Oláh
gular paraproct is rather stable. Some populations in Gorce Mts and Babia Gora
Mts have some sign of paraproct elongation probably as a result of having contact
zone or very complex and irregular moving cline with P. psorosa. The curvature of
the paramere rod is shallow. The dental pattern on the paramere head is rather
elaborated and packed with apical and dorsosubapical teeth. P. bohemosaxonica
is distributed from the Ore Mts to the Poliana Mts.”
Psilopteryx gutinensis Mey et Botosaneanu, 1985

Psilopteryx psorosa gutinensis Mey et Botosaneanu, 1985: 120: “Holotypus
² und Allotypus ³: Gutin-Berge, Sasarului-Tal bei Baia Sprie, 14.XI.1962 (in
coll. Botosaneanu); 1²- und 1 ³-paratyp vom selben Ort, in coll Royal Ontario
Museum. Areal: Gutin-Berge.”
Psilopteryx gutinensis Mey et Botosaneanu, 1985: Oláh et al. (2015b: 99–
100): raised to species status. “Romania, Maramureş county, Muntii Ignis, Deseşti-
Staţiunea Izvoare, open brook on the Valhani Plateau, 1020m, N47°43.015’
E23°44.547’, 7.X.2010, leg. P. Barcánfalvi, D. Murányi & J. Oláh, (1 males, OPC).
Maramureş county, 2010/4, Muntii Ignis, Deseşti-Staţiunea Izvoare, brook in
bushy edge on the Valhani Plateau, 930m, N47°44.374’ E23°43.331’, 8.X.2010, leg.
P. Barcánfalvi, D. Murányi & J. Oláh, (1 female, OPC). Maramureş county, Muntii
Ignis, Deseşti-Staţiunea Izvoare, open brook on the Valhani Plateau, 1020m,
N47°43.015’ E23°44.547’, 21.X.2010, leg. Á. Ecsedi, J. Oláh & I. Szivák, (1 male, 1
female, OPC). Maramureş county, Muntii Ignis, Deseşti-Staţiunea Izvoare, open
spring brook at settlement, 920m, N47°45.167’ E23°43.013’, 22.X.2010 leg. Á.
Ecsedi, J. Oláh & I. Szivák, (3 males, 1 female, OPC). Maramureş county, Muntii
Ignis, Deseşti-Staţiunea Izvoare, side valley spring brook along the road between
Firiza and Statiunea Izvoare, 600 m, 22.X.2010 leg. Á. Ecsedi, J. Oláh & I. Szivák,
(1male, 1 female, OPC).”
Diagnosis – Oláh et al. (2015b: 99–100): “This sibling incipient species is
characterized by short dorsal branch of paraproct. The lateral profile of the short
and high subtriangular paraproct is rather variably. The curvature of the para-
mere rod is deep. The dental pattern on the paramere head is characterized by an
enlarged single leading tooth. This species is distributed in the Ignis and Gutin
Mts.”
Psilopteryx javorensis Oláh et Chvojka, 2015

Psilopteryx javorensis Oláh et Chvojka, 2015 in Oláh et al. (2015: 100):
“Holotype. Slovakia, Javorie Mts, Blyskavica, stream Tisovnik, 657m, 20.X.2005,
leg. D. Murányi (1 males, HNHM). Allotype. Same as holotype (1 female,
HNHM). Paratypes. Same as holotype (3 males, 1 female HNHM). Slovakia,

Banskobystrický region, Poľana Mts, Hriňová, sidebrook of Slatina Stream,

N48°37.210’ E19°31.582’, 514m 8.X.2013, singled leg. J. Oláh & L. Szél (1 male,
OPC). Banskobystrický region, Javorie Mts, Stará Huta, Blýskavica, Tisovník
Stream, N48°27.553’ E19°18.048’, 671m, 7–9.X.2013, singled leg. J. Oláh & L.
Szél (1 female, OPC).”
Diagnosis – Oláh et al. (2015: 100): “This new sibling incipient species is
characterized by short dorsal branch of the paraproct and belongs to the P. bohe-
mosaxonica species cluster. Most close to the nominate species P. bohemosaxoni-
ca, but differs by having the lateral profile of the short and high paraproct very
regular triangular at all the examined 5 specimens, not irregular subtriangular.
The dental pattern on the paramere head is reduced to a narrowing bifid tip, not
densely packed with teeth apicad and dorsosubapicad. This species is distributed
in the Javoros Mts.”
the type material was collected, Javoros Mts.”
Psilopteryx psorosa species complex

Psilopteryx harmas Oláh et Chvojka, 2015
Psilopteryx harmas Oláh et Chvojka, 2015 in Oláh et al. (2015b: 101):
“Holotype. Poland, East Carpathians, Bieszczady Mts. at Wolosatka brook, 900m,
28.X.2010, leg B. Szczęsny (1 male, OPC). Allotype. Same as holotype (1 female,
OPC). Paratypes. East Carpathians, Bieszczady Mts. at Wolosatka brook, 850-
1000m, 22.X.2014, leg B. Szczęsny (1 male, OPC). East Carpathians, Bieszczady
Mts. no more data, leg B. Szczesny (2 males, 1 female; OPC). Slovakia, NE
Slovakia, Bukovské Hills, Stuzica stream, left tributary Kremenny potok stream,
N49o04’23” E22o32’33”, 7. XI. 1999, leg J. Lukas (1 male, 2 females; OPC, ). NE
Slovakia, Bukovské Hills, Kremenny potok stream, N49o04’23” E22o32’33”, 8.
XI. 1999, leg J. Lukas (1 male, OPC; 1 male, NMPC). East Slovakia, Vihorlat
Mts. Cerny potok (above Zemplinske Hamre), 640m, 12.X.1990, leg P. Chvojka
(4 males, 1 female, NMPC; 1 male, 1 female; OPC). East Slovakia, Vihorlat Mts.
Zemplinske Hamre, stream below source, 700m, 15.X.1962, leg. J. Sykora (5
males, 4 females, NMPC; 2 males, 1 females,OPC; 1 male, 1 female; CSNMB).
East Slovakia, Vihorlat Mts. Strihovsky potok brook NW Strihovce, 590-600m,
12.X.1990, leg P. Chvojka (1 male, 3 fgemales; NMPC). East Slovakia, Vihorlat
Mts. Malá Bystra, 560-700m, 9.X.1990, leg P. Chvojka (2 males, OPC).”
Diagnosis – Oláh et al. (2015b: 101): “This new incipient species is char-
acterized by medium long dorsal branch of the paraproct and therefore belongs
to the P. psorosa species cluster. Close to the nominate species P. psorosa, but
differs by having the lateral profile of the paraproct higher, not as low; para-

214 J. Oláh
mere curvature deeper and the dentate apex turned mesad or upward. It is dif-
ficult to demonstrate how its incipient specific status is related to the putative P.
bohemosaxonica/P. psorosa hybrid cline effect. However its paraproct and para-
mere clearly differ from the typical paraproct and paramere structures of pure
P. psorosa populations. This species might represent a hybrid species of dubious
origin and needs further taxonomic and population genetical studies. The forma-
tion of new hybrid taxa is possible by introgression of those loci that promote
adaptive divergence or reproductuve barrier building (Abbott et al.. 2013). This
species is distributed in the Vihorlat Mts. and Beszczady Mts.”
Etymology – “Harmas from “hármas” triple in Hungarian, refers to the dis-
tributional area of the species centred around the triple borders of three coun-
tries: Poland, Slovakia and Ukraine.”
Psilopteryx psorosa (Kolenati, 1860)

Chaetopteryx psorosa Kolenati, 1860: 388–389: “Areal: Biesczady, Beski-
den, Hohe- und Niedere Tatra, Sudeten.”
Psilopreyx psorosa (Kolenati, 1860): Stein (1874: 250): Psilopteryx genus
erected and C. psorosa transferred as type species.
Psilopteryx psorosa (Kolenati, 1860): Mey & Botosaneanu (1985: 120):
nominate subspecies of the complex: Psilopteryx psorosa psorosa.
Psilopteryx psorosa carpathica Schmid, 1952: Szczęsny (1986: 537): misi-
dentification.
Psilopteryx psorosa (Kolenati, 1860): Oláh et al. (2015b: 102–103): raised
to species status. “Czech Republic, Jesenik, Ovcarna, source of Bila Opava, 110m,
24. IX. 2000, leg. Sivec & B. Horvát near type locality (10², 1³ OPC). Northern
Moravia, Králicky Snéznik Mt., Morava, Morava River below Králicky Snéznik
Mt., N50°11’59” E16°50’36”, 1100m, 19.X.2000, leg. P. Chvojka (6 males, 3 fe-
males; NMPC). Eastern Bohemia, Králicky Snéznik Mt., brook, “Strasidla”SW
Králicky Snéznik Mt., N50°11’35” E16°49’39”, 1070m, 19.X.2000, leg. P. Chvojka
(14 males, 5 females; NMPC). Poland: Right tributary of the Olczyski Potok,
(Olczyski Spring), the Tatra Mts., 12.XI.1986, leg. B. Szczęsny males, 5 females,
Koscieliska Valley, the Tatra Mts. 25.X.1985, leg. B. Szczęsny males, 5 females,
Tatra Mts., IX-X. leg. B. Szczęsny males, 2 females, The Biała Lądecka River,
Masyw Śnieżnika, The East Sudety, N 50° 14’ 17” E 17° 0’ 5.76”, 9.XI.2009, leg.
K. Majecka 1 female The Kleśnica River, Masyw Śnieżnika, The East Sudety, N
N 50° 15’ 35” E 16° 51’ 41” , 10.XI.2009, leg. K. Majecka 1 male Biały Spław, be-
ginnig of stream the Biała Lądecka River, Masyw Śnieżnika, The East Sudety, N
50° 13’ 47” E 17° 1’ 4”, 9.XI.2009, leg. K. Majecka 1 female ). Karkonowski Park
Narodowy, Hala Szrenicka, Kamienczyk stream, spring area, 11.X.2014, leg. J.

Majecki (1 male, OPC). Karkonowski Park Narodowy, Równia pod Sniezka,

spring area, 12.X.2014 leg. J. Majecki (1 male, 1 female in copula; OPC). Slovakia,
Vysoké Tatry, brooks E of Zlomiskovy potok brook, N49°08’14” E20°01’05”,
1460m, 7.X.1989, leg. P. Chvojka (3 males, NMPC). Vysoké Tatry, Furkotsky
potok brook, N49°08’014” E20°02’015”, 1480m, 7.X.1989, leg. P. Chvojka (1 fe-
male, NMPC). Vysoké Tatry, Zlomiskovy potok brook, N49°07’18” E20°00’59”,
1460m, 7.X.1989, leg. P. Chvojka (1 male, 1 female, NMPC). Podtatranská kot-
lina basin, Lieskovec, N49°07’13” E20°03’04”, 1280m, 7.X.1989, leg. P. Chvojka
(1 male, 1 female; NMPC). Podtatranská kotlina basin, Nové Strbské pleso,
N49°07’04” E20°03’56”, 1315m, 7.X.1989, leg. P. Chvojka (2 males, NMPC).
Podtatranská kotlina basin, right tributary of Bela stream, SW Hrdovo (NE
Pribylina) N49°07’04” E19°50’45”, 830m, 11.X.1989, leg. P. Chvojka (2 males,
3 females; NMPC). Oravské Beskydy Mts. left tributary of Bystrá stream below
Babia hora Mt., N49°33’39” E19°30’47”, 1300m, 15.X.1991, leg. P. Chvojka (6
males, 4 females; NMPC).”
Diagnosis – Oláh et al. (2015b: 102–103): “This incipient species is char-
acterized by medium long dorsal branch of the paraproct. The lateral profile of
the paraproct is subtriangular and very low, the lowest compared to all the other
species. The curvature of the paramere rod is shallow. The dental pattern on the
paramere head is concentrated on the very apical region. Individual variation
and intermediate cline formations might be connected with complex contact
zones with P. bohemosaxonica. This species is distributed in Jesenic, Králicky
Snéznik, Sudety, Tatra, Babia Hora Mts. However its distributional and hybrid
formation processes needs further studies as was suggested already long before
(Mey et Botosaneanu, 1985).”
Psilopteryx carpathica species complex

Psilopteryx carpathica Schmid, 1952
Psilopteryx carpathica Schmid, 1952: 142–144: “Holotype: 1² Czarnohora
17. X. 1908, Carpathes (Dziędzielewicz).”
Psilopteryx psorosa carpathica Schmid, 1952: Mey & Botosaneanu
(1985: 120): downgraded to subspecies level. “Areal: Rodna-, Maramures- und
Tschernogora-Gebirge.”
Psilopteryx carpathica Schmid, 1952: Oláh et al. (2015b: 104): species status
reinstated. “Romania, Borsa, 26.IX.1992, leg J. Oláh, (1 male, OPC). Maramures
county, Rodna Mts. Borsa-Staţiunea Borsa, stream along the road towards Prislop
Pass, 1014 m, N47° 37’ 34.0’’ E24° 49’ 13.0’’, 26.IX.2006, leg. L. Dányi-J. Kontschan
& D. Murányi (1 male, HNHM). Maramures county, Maramures Mts. Borsa-Baile
Borsa, brook over the village 1046 m, N47° 40’ 21.5’’ E24° 50’ 16.7’’, 26.ix.2006

216 J. Oláh
leg. L. Dányi, J. Kontschán & D. Murányi (1 male deformed, HNHM). Radnei

Mts. numerous spring streamlets on the spring area of Cailor waterfall, Piatra
Rea, N47°35’1.9” E24°47’49.4”, 1564m, 28. IX. 2014, leg. J. Oláh & Cs. Balogh (1
male, OPC). Radnei Mts. small spring below Lake Isvoru Bistritei, N47°34’46.4”
E24°48’49.34”, 1586m, 28. IX. 2014, leg. J. Oláh & Cs. Balogh (1 male, 1 female;
OPC). Ukraine: East Carpathians, Gorgany Mts., at Sitny brook, 1111m, 29.X.2005,
leg B. Szczęsny (1 male, 1 female; OPC). East Carpathians, Skupova Mts., Hnylec
brook, 1470 m, 26.X.2006, leg B. Szczęsny (4 males, 3 females; OPC). Czarnohora
Massif, Prut River at Forestenka, 26.X.2005, leg. B. Szczęsny (1 male, 1 female;
OPC). Czarnohora Massif, 8.X.19955, leg. B. Szczęsny (4 males, 1 female; OPC).”
Diagnosis – Oláh et al. (2015b: 104): “This incipient nominate species of
the complex is characterized by long dorsal branch of the paraproct. The lateral
profile of the paraproct is elongated subtriangular and the lowest compared to
the two other species. The curvature of the paramere rod is shallow. The deeper
curvature found at the single specimen examined from the Gorgany Mts. could
be a result of copulatory processes. The simplified dental pattern on the para-
mere head is restricted to the very apical almost bifid tip. This species populates
spring fed streams in the Gorgany, Czarnohora, Maramaros and Rodnai Mts. of
the North-East Carpathians.”
Psilopteryx retezatica Botosaneanu et Schneider, 1978

Psilopteryx psorosa retezatica Botosaneanu et Schneider, 1978: 321–
322: “Retezat-Geb. 100–2100 m, 17.IX.-14.X.1927 Diós”.
Psilopteryx psorosa retezatica Botosaneanu et Schneider, 1978: Mey & Bo-
tosaneanu (1985: 119).
Psilopteryx retezatica Botosaneanu et Schneider, 1978: Oláh et al. (2015b:
104–105): raised to species status. “Romania, Retezat Mts., Gura Apelor, N45.33
E22.88, 1500m, 20.X.2007, leg. M. Bálint, E. Magyari & M. Braun (23 males,
13 females; OPC). Retezat Mts. 24 km from Baile Herculane, spring area of a
small tributary to River Cerna, N45° 2’32.14” E22°35’3.36”, 13.XI.2010, singled
leg. Á. Ecsedi & I. Szivák (2 males, OPC). Caraş-Severin county, Ţarcu Mts. left
side brook of open stream on the N slope of Mt. Ţarcu, 1500 m, N45°17’40.7”,
E22°31’44.5”, 14.10.2011, leg. Á. Ecsedi, T. Kovács, G. Puskás (1², OPC). Caraş-
Severin County, Ţarcu Mts. spring and its outlet at Cuntu Meteorological Station,
N45°18’00.2”, E22°30’04.3”, 1465 m, 14.10.2011, leg. Á.Ecsedi, T. Kovács, G.
Puskás, (19²,8³, OPC). Vâlcea county Parâng Mts, Obrâşia Lotrului, open brook,
200 m of Transalpina (67C) road, 45°22’46.1”, 23°38’30.6”, 1765 m, 9.XI.2014,
leg. T. Kovács & G. Magos (3 males, 3 females, OPC). Vâlcea county, Parâng
Mts, Obrâşia Lotrului, open brook, 900 m of Transalpina (67C) road, 45°23’9.9”,

23°39’24.9”, 1780 m, 9.XI.2014, leg. T. Kovács & G. Magos (2 males, 1 female,

OPC). Vâlcea county, Parâng Mts, Obrâşia Lotrului, open spring area, 100 m
of Transalpina (67C) road, 45°22’27.7”, 23°39’4.0”, 1915 m, 9.XI.2014, leg. T.
Kovács & G. Magos (1 male, OPC).”
Diagnosis – Oláh et al. (2015b: 104–105): “This incipient species of the
complex is characterized by long dorsal branch of the paraproct. The lateral
profile of the paraproct is irregular subtriangular and the highest in the species
group. The curvature of the paramere rod is deeper compared to the other two
species of the complex and having constrictions and dilatations more frequently.
The dental pattern on the paramere head is most elaborated in the entire species
group. Dentate pattern moves more far into the middle direction in lateral view.
This species populate spring fed streams in the Tarcu, Retezat, Muntele Mic,
Cerna and Parang Mts. In Parang Mts. there are sign of hybrid contact zone with
P. transsylvanica in the shape or dental pattern of the speciation traits of parap-
roct and paramere.”
Psilopteryx transylvanica Mey et Botosaneanu, 1985

Psilopteryx (Psilopteryx) carpathica Schmid, 1952: Botoşăneanu (1957b:
193–194): misidentification.
Psilopteryx psorosa transylvanica Mey et Botosaneanu, 1985: 120: “Holo-
typus ² und allotypus ³: Fagaras, Bilea Cascada, 28. 10. 62, (in coll. Botosaneanu):
13²- und 6³-Paratypen vom selben Fundort, (2 ²-Paratypen in coll. Mey).”
“Areal: Cibin-Gebirge, Fagaras, Bucegi, Karpathenknie, Hargitha-, Ceahlau und
Stinisoara-Gebirge.”
Psilopteryx transylvanica Mey et Botosaneanu, 1985: Oláh et al. (2015b: 105–
107): raised to species status. “Romania. Parcul Natural Bucegi, Peles, 7.X.1954,
leg. P. Iuncu (1 male, OPC). Parcul Natural Bucegi, V.Pelesului, 4.X.1954, leg. P.
Iuncu (4 males, OPC). Parcul Natural Bucegi, Peles, 27.X.1954, leg. P. Iuncu (2
males, 1 female; OPC). Parcul Natural Bucegi, Parcul Natural Cascada Urlatoarea,
22.X.1954, leg. P. Iuncu (2 males, 1 female; OPC). Gurghiu Mts. near Bucin
Pass, Tárnava Mica springs and stream, N: 46°39’ 16,63”E: 25°16’ 42,46”, 1290,
30.X.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (13 males, 9 females;
OPC). Gurghiu Mts. near Bucin Pass, Gainasa springs and stream, N: 46°40’
11,35” E: 25°17’ 39,06”, 1400, 30.X.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés
& J. Oláh (66 males, 7 females; OPC). Gurghiu Mts. near Bucin Pass, Frasileasa
stream with side springs, N46°38’ 37,45” E: 25°17’ 35,08”, 1193, 29.X.2014, leg. Z.
Baczó, Cs. Balogh, J. Kecskés & J. Oláh (8 males, 3 females, OPC). Hargitha Mts.
Filio stream side spring, N: 46°27’ 03,90” E: 25°33’ 29,29”, 1350m, 31.X.2014
leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (9 males, 5 females; OPC).

218 J. Oláh
Hargitha Mts. Filio stream side spring, N: 46°27’ 14,53” E: 25°33’ 53,04”, 1415m,
31.X.2014 leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (12 males, 6 females;
OPC). Hargitha Mts. Filio stream side spring, N: 46°26’ 45,18” E: 25°34’ 25,73”,
1600m, 31.X.2014 leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (7 males, OPC).
Hargitha Mts. Filio stream side spring, N: 46°26’ 29,54” E: 25°34’ 48,09”, 1625m,
31.X.2014 leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (29 males, 9 females,
OPC). Dâmboviţa county, Bucegi Mts, M. Dichiu, left sidebrook of V. Oboarele,
45°19’32.8”, 25°26’05.0”, 1420 m, 6.XI.2014, leg. T. Kovács & G. Magos (2 males,
1 female; OPC). Dâmboviţa county, Bucegi Mts, Hotel Peştera, Valea Cocora,
45°23’04.1”, 25°26’37.6”, 1590 m, 6.XI.2014., leg. T. Kovács & G. Magos (1 male,
1 female; OPC). Dâmboviţa county, Bucegi Mts, Hotel Peştera, V. Şugărilor,
45°24’42.1”, 25°27’23.5”, 1850 m, 6.XI.2014, leg. T. Kovács & G. Magos (1 male,
OPC). Dâmboviţa county, Bucegi Mts, Hotel Peştera, left sidebrook of Ialomiţa,
45°24’08.8”, 25°26’35.1”, 1690 m, 6.XI.2014, leg. T. Kovács & G. Magos (9 males,
7 females, OPC). Dâmboviţa county, Bucegi Mts, Hotel Peştera, spring area be-
side the V. Şugărilor, 45°24’25.0”, 25°26’47.8”, 1760 m, 6.XI.2014, leg. T. Kovács
& G. Magos (5 males, 3 females, OPC). Dâmboviţa county, Bucegi Mts, Hotel
Peştera, Ialomiţa, 45°23’54.5”, 25°26’25.1”, 1610 m, 7.XI.2014 leg. T. Kovács & G.
Magos (1 male, 1 female, OPC). Dâmboviţa county, Bucegi Mts, Hotel Peştera,
spring area beside the Cascada Obrisia Ialomiţei, 45°25’35.2, 25°26’42.8”, 2030
m, 7.XI.2014, leg. T. Kovács & G. Magos (3 males, 1 female, OPC). Dâmboviţa,
Bucegi Mts, Lacul Bolboci, Blana Stream, 45°22’06.0”, 25°26’40.9”, 1515 m,
7.XI.2014, leg. T. Kovács & G. Magos (2 males, 3 females, OPC). Sibiu county,
Făgăraş Mts, Cârţişoara, spring beside the Bâlea Stream, 45°37’59.4”, 24°36’31.3”,
1290 m, 8.XI.2014, leg. T. Kovács & G. Magos (13 males, 11 females, OPC). Sibiu
county, Făgăraş Mts, Cârţişoara, open sidebrook of Bâlea Stream below the Bâlea
Lake, 45°36’30.4”, 24°37’14.6”, 1940 m, 8.XI.2014, leg. T. Kovács & G. Magos (1
male, 2 females, OPC). Sibiu county, Făgăraş Mts, Cârţişoara, Bâlea Stream be-
low the Bâlea Lake, 45°36’30.4”, 24°37’14.6”, 1940 m, 08.11.2014, leg. T. Kovács
& G. Magos (1 male, 2 females, OPC).”
Diagnosis – Oláh et al. (2015b: 105–107): “This incipient species of the
complex has long dorsal branch of the paraproct. The lateral profile of the para-
proct is characterized by almost regular straight dorsum and downward directed
basal region. The curvature of the paramere rod is shallow and having constric-
tions and dilatations more frequently in the most western populations in the
Fagaras Mts. The dental pattern on the paramere head is restricted to apicad
and dorsosubapicad. Dentate pattern does not move more far into the middle
direction in lateral view, except again in populations of the Fagaras Mts. The
contact zone with P. retezatica could be more complicated extending even into
the Fagaras Mts. This species is most distributed in the complex; populating

spring-fed small streams on higher elevations in mountan ranges in the eastern

and southern Carpathians. We have collected numerous specimens in Gurghiu,
Hargitha, Bucegi and Fagaras Mts.”
Stenophylacini
Acrophylax Brauer, 1867
Acrophylax vernalis Dziędzielewicz, 1912
Acrophylax vernalis Dziędzielewicz, 1912: 134–136: “Spostrzegałem ten
gatunek na górnej granicy lasów Czarnohory i w krainie kosodrzewn pod szc-
zytami Dancerza i Breskula w wysokości ad okolo 1200–1500 m. n. p. m, przy
potokach, w czasieod 15 do 20-go maja w malej ilości okazów, zgromadzonych
tylko na pewnych przestrzeniach potoków. Jawi się on między zaspami şniegu,
zalegającego od zimy. Ukriwa się na brzegach poloków w szczelinach skał i kami-
eni. Tak samce jak i samice wylatują z kryjówek, gdy słońce świeci, i w miejscach
oświetlonych przelatują szybko z biegiem potoku i w górę w wysokości okolo 15
metra nad powierzchnią wody; od czasu do czasu usiadają na gałęziach drzew
lub kosodrzewu, zwisających nad wodą, albo na krawędziach zasp śniegowych,
na których biegają a od czasu di czasu zatrzymują się i podnosząc glówke i różki,
rozgladają się po otocheniu.”
Acrophylax vernalis Dziędzielewicz, 1912: Szczęsny (1980: 464): in NHM-
ISEA: “2³³; both specimens have the same inventory number 3/25 and identical
labels “Czarnohora, Dancerz, 15.V.1911” and “Acrophylax vernalis Dz. ³ ver. C.
Tomaszewski”. Dziędzielewicz (1912) desdribed this species on the basis of speci-
mens collected from Czarnohora near the summits of Dancerz and Breskul dur-
ing the period 15–20.V.1911 thus the specimens deposited in Cracow belong to
the typical series and are syntypes. I am desinating one of the females preserved
in a better state as a lectotype”.
Acrophylax vernalis Dziędzielewicz, 1912: Szczęsny & Godunko (2007:
36): in SMNHL: “12²², 7³³; East Carpathians: Czarnohora Massif; 4 ²², 6³³
which Dziędzielewicz collected during 15–30.V.1911 (on the Czarnohora massif on
slopes of Dancerz and Breskul) are paralectotypes according to /01–10, i. e. belong
to the series of specimens on basis of which Dziędzielewicz decsribed this species.
A designated lectotype is stored in MP ISEZ (Szczęsny 1980). Chornohora Massif
above forest line is locus typicus for the species (Szczęsny & Godunko, 2008).”
Allogamus Schmid, 1955

Allogamus dacicus (Schmid, 1951)
Halesus dacicus Schmid, 1951a: 65–66: “Retyezat (Transylvanie), déposé
au musée de Budapest.”

220 J. Oláh
Allogamus dacicus (Schmid, 1951): Schmid (1955: 194–196): transferred to

the newly established genus Allogamus.
Allogamus dacicus (Schmid, 1951): Botoşăneanu (1959: 67): “Această spe-
cie a fost descrisă de Schmid (1951) din Retezat; noi nu am reuşit să o regăsim
în acest masiv si putem raporta tot materialul de Allogamus colectat in Retezat la
specia foarte îndeaproape înrudită uncatus Brau. Pe A. dacicus îl cunoaştem din
Făgăraş (lacul Podragul).”
Allogamus dacicus (Schmid, 1951): Botosaneanu (1961a: 59): “Lac
Podragul, Fagaras, VIII. 1950.”
Allogamus dacicus (Schmid, 1951): Botosaneanu (1975: 99): “A. daci-
cus Schmid ist bis jetzt durch drei Belegstücke bekannt: ein Stück vom Retezat-
Massiv, die zwei anderen vom Fagarasch-Massiv (Südkarpaten). Es ist eine
Hochgebirgsart (1900–2100 m), die im August in der Nahe von Seen und deren
Ausflüssen fl iegt.”
Allogamus dacicus (Schmid, 1951): Mey (1978: 62): collected in Romania:
“Fagaras Gebirge: Lacul Avrig: 8. X.1977., 5²; Serata: 12. X.1977., 3²; Podragu:
14. X.1977. 2².”
Allogamus dacicus (Schmid, 1951): Szczęsny & Chvojka (2008: 164):
“Ukraine, Czarnohora Massif (Chornohora), Dancerz stream at forest line,
9.X.1995, leg B. Szczęsny, 1 ².”
Allogamus dacicus (Schmid, 1951): Oláh et al. (2014: 74): new records:
“Romania. Argeş county, Făgăraş Mts, Căpătânenii Ungureni, small springlake by
the Capra Stream along road No.7C, N45°34.605’ E24°37.060’, 1405m, 29.VIII.2012
leg. T. Kovács, D. Murányi, J. Oláh (1 male, 1 female; OPC). Braşov county, Făgăraş
Mts, Dejani, right sidebrook of Dejani stream, N45°35.446’ E24°56.348’, 1755m,
30.VIII.2012 leg. T. Kovács, D. Murányi, J. Oláh (1 male, OPC).”
Allogamus starmachi Szczęsny, 1967

Allogamus starmachi Szczęsny, 1967: 479: “Holotype ²: Tatra, Vallée Gasi-
enicowa, 1680 m, 2.X.1966; paratypes 6²² capturés avec holotype, et 4²² Vallée
Panszczyca, 1600 m, 3.X.1966. leg M-me M. Kownacka. Tous les specimens dans
la collection de l’auteur.”
Allogamus starmachi Szczęsny, 1967: Botosaneanu & Malicky
(1978: 353): synonymised with A. starmachi Szczęsny, 1967: Allogamus lazarei
Szczęsny, 1967: 480–481. Allogamus tatricus Szczęsny, 1967: 481–482.
Allogamus tomor Oláh, 2012

Allogamus tomor Oláh, 2012 in Oláh & Kovács (2012: 95–96): “Holotype.
Albania: Skrapar district, Ostrovicë Mts, Backë, Krojmbret Spring and its out-

let brook NE of the village, N40°31.753’ E20°25.152’, 1965 m, 12.10.2012, leg. P.

Juhász, T. Kovács, D. Murányi, G. Puskás (1², OPC). Allotype. Same as holotype
(1³, OPC).”
Diagnosis – Oláh & Kovács (2012: 95–96): “Having mesad angled gono-
pods together with three-armed aedeagus and fused paramere this new species is
close to Allogamus uncatus (Brauer, 1857), but differs by having “apparent harp-
ago” with transversally cut trilobed apical margin, not with pointed or narrow-
ing monolobed apicoventral corner; aedeagus abbreviated and dilated, not long
and slender; on female the elongated mesal structure of the vaginal sclerite com-
plex short, not as long as at A. uncatus. This elongated sheath is connected to the
dorsum of the vaginal or spermathecal sclerite was first mentioned by Schmid
(1951 as a vestibular apparatus with equilibrating function. Later Schmid (1955)
mentioned as bursa copulatrix. We have found this long tube-like structure as
functioned to receive the long fused paramere in copulation.”
Etymology – “Tomor from “tömör” solid or concise in Hungarian, refers to
the abbreviated and dilated aedeagus of the phallic organ.”
Allogamus zugor Oláh, 2014

Allogamus zugor Oláh, 2014 in Oláh & Kovács (2014: 113–114): “Holotype.
Macedonia, Southwestern region, Jablanica Mts, 6.5 km W of Labuništa, open
brook at Labuniško Lake, N41°16.069’, E20°31.242’, 1905 m, 10.10.2014, leg. P.
Juhász, T. Kovács, G. Puskás (1², OPC). Allotype. Same as holotype (1³, OPC).
Paratypes. Same as holotype (1³, OPC; 1³, MM).”
Diagnosis – Oláh & Kovács (2014: 113–114): “Having three-armed ae-
deagus and fused paramere this new species is a member of uncatus group and
having mesad angled gonopods is close to A. uncatus, it is a sister species of A.
uncatus, and A. tomor. It differs form both sisters by having “apparent harpago”
with monolobed apical margin turned back from transversal to sagittal plane; ae-
deagus minituarized shrunk, not long slender like at A. uncatus or broad dilated
like at A. tomor; parallel with shrunk aedeagus, the vagina is very small, similarly
abbreviated in sexual coevolution processes.”
Etymology – “Zugor from “zsugor”, shrink in Hungarian, refers to the ab-
breviated and highly shrunk aedeagus, which coevaluated with the abbreviated
vaginal chamber.”
Chionophylax Schmid, 1951

Chionophylax czarnohoricus (Dziędzielewicz, 1911)
Acrophylax czarnohoricus Dziędzielewicz, 1911b: 45–46: “Czarnohora,
mons Tomnatyk, in altitudine 1791 m. s. m., 30. V. 1909. 3² zebrał P. Karol

222 J. Oláh
Huppenthal przy jeziorku pod szczytem Tomnatyka na Czarnohorze w wysocości

1791 m. n. p. m; ukrywały się tu w kosodrzewinie. 30.V.1909.”
Chionophylax czarnohoricus (Dziędzielewicz, 1911): Schmid (1951b: 55–
58): transfered to the new genus Chionophylax.
Chionophylax czarnohoricus (Dziędzielewicz, 1911): Szczęsny (1980: 464):
in NHM-ISEA: “4²², 2³³ with the inventory number 4/25 are labelled with the
inscription “Czarnohora, Dancerz 29.V.1911” the other two with the inventory
number 5/25 are labelled with the inscription “Czarnohora Breskul 1.VI.1911”.
Dziędzielewicz (1911) described the species on the basis of 3²² collected
by K. Huppenthal by a small lake near the summit of Tomnatek, Czarnohora
30.V.1909 so none of the specimens deposited in Cracow belong to the typical
series. However, they do come from the “terra typica”.
Chionophylax czarnohoricus (Dziędzielewicz, 1911): Szczęsny & Godunko
(2007: 36): in SMNHL: “16²², 1³; East Carpathians: Czarnohora Massif; 3²² col-
lected by Karol Huppenthal 30.V.1909 at the pond on slope of Tomnatyk Mt (“przy
jeziorku pod szczytem Tomnatyka”) on Czarnohora massif (1791 m a.s.l.) were the
basis for description of the species under generic name Acrophylax (Dziędzielewicz
1911). In SMNHL there are stored 2²² collected in 1909: one male “30.V.1909
Czarnohora, Gadzyna, jeziorko” – now designated as lectotype (according to the
ICZN Articles 73.2 and 74.1, No /01), second male “4–5.VI.1909, Czarnohora,
zeb. Dr. Antoni Lomnicki”. As to the former specimen, it is highly probable that
Dziędzielewicz when writing “pond on slope of the Tomnatyk Mt” wanted to point
at the pond in Gadzyna valley, and not on the Barbeniescu lake which is situated di-
rectly at Tomnatyk Mt. The latter specimen not mentioned in this paper was found
by Raciecka (1934) in Dziędzielewicz’s collection. The remaining ones (14²² –
1911 and 1³ – 1910) were collected by Dziędzielewicz in locus typicus. The paralec-
totype is stored in NMP (P. Chvojka, pers. comm.).
Chionophylax czarnohoricus (Dziędzielewicz, 1911): Szczęsny & Chvojka
(2008: 157): in NMP: Paralectotypes of 3² specimens were collected in 30.V.1909
in the East Carpathians, Czarnohora from the type locality and deposited in NMP.
Chionophylax mindszentyi Schmid, 1951

Chionophylax mindszentyi Schmid, 1951b: 59–61: “Holotype ²: Vurfu
Mare, 11. IV. 1910 (Szeben, “Cziki”, correctly: Csíki); il appartient au musée
Budapest.”
Chionophylax mindszentyi bulgaricus Kumanski, 1973

Chionophylax mindszentyi bulgaricus Kumanski, 1973b: 194–196:
“Holotype ² et paratype ³ (les deux a sec), recueillis le 26. IV. 1968 sur la neige

couvrant un torrent au-dessous du mont Botev, Stara planina, environ 2100–2200

m d’altitude (leg. P. Beron).”
Chionophylax monteryla Botoşăneanu, 1957

Chionophylax czarnohoricus monteryla Botoşăneanu, 1957c: 599–603:
“Récemment nous avons eu le plaisir de recevoir Prof. Dr. Al. Valkanov un tube
contenant 28 exemplaires (25 ²², 3³³) d’un Trichoptère capturé par lui-même le
13 avril 1955 sur les bords du “VIème lac” (“Sedemite ezera”) dans le haut massif
de Ryla, en Bulgarie.”
Chionophylax monteryla Botosaneanu, 1957: Kumanski (1973b: 196–203):
raised to species status. “Les 28 exemplaires dont Botosaneanu a disposé lors de
description de ssp. monteryla sont en réalité une petite partie d’une récolté beau-
coup plu riche comprenant 394 exemplaires. La récolte est faite le 13. IV.1955.”
Isogamus Schmid, 1955

Isogamus aequalis (Klapálek, 1907)
Anisogamus aequalis Klapálek, 1907: 24–27: “Naleziště Chomiak (Błotek)
ve Vých. Karpatech 22./IX.1906, leg. Joz. Dziędzielewicz, 3² a 2³”.
Isogamus aequalis (Klapálek, 1907): Schmid (1955: 183–184): transferred
to his newly erected genus Isogamus.
Isogamus aequalis (Klapálek, 1907): Szczęsny (1980: 466–471): differential
diagnoses of both sexes of I. aequalis and I. czarnohorensis from the Ukrainian
Carpathians were elaborated and detailed drawings prepared.
Isogamus aequalis (Klapálek, 1907): Chvojka (1993: 217–220): the first
collection and reliably records from Slovakia with excellent drawings were pre-
sented.
Isogamus aequalis (Klapálek , 1907): Oláh et al. (2015b: 41–42): “Poland,
Bieszczady Mts. X. leg. B. Szczęsny (2 males, 2 females; OPC). Slovakia, Vihorlat
Mts. Zempl Hamre, 700 m, 15. IX. 1962, leg. P. Chvojka (3 males, 4 females,
aedeagus and parameres on slide No. 13,14, NMPC; 3 males, 2 females, OPC).
Vihorlat Mts. prameniste pot.-Morska eko, 15.IX.1962, leg. P. Chvojka, (4 males,
aedeagus and parameres on slide No. 16, NMPC). Vihorlat, 16.IX.1962, leg. P.
Chvojka (2 males, aedeagus and parameres on slide No. 1,2, NMPC). Vihorlat
Mts. prm. N. Zempl. Hamre, 740 m, 12.X.1990, leg. P. Chvojka (1 male, ae-
deagus and parameres on slide No. 18, NMPC). Vihorlat Mts. prm. N. Zempl.
Hamre, 740 m, 12.X.1990, leg. P. Chvojka (1 male, aedeagus and parameres on
slide No. 18, NMPC). Vihorlat Mts. potucek severne, N Sedlice, 10.X.1990, leg.
P. Chvojka (1 male, aedeagus and parameres on slide No. 17, NMPC). Vihorlat
Mts. levostr. Pritok Bystre, 5–700 m, 9.X.1990, le. P. Chvojka (4 males, aedea-

224 J. Oláh
gus and parameres on slide No. 16, NMPC). Ukraine, Klapálek Collection: No.
21, Worochtensky, 7.IX.1908, leg. Dziędzielewicz (1 female, NMPC). Klapálek
Collection: No. 5, Chomiak (Blotek), 22.IX.1906, leg. Dziędzielewicz (1 male,
aedeagus and parameres on slide No K5, NMPC). No. 6, Chomiak, 5.IX.1908,
leg. Dziędzielewicz (1 male, aedeagus and parameres on slide No K6, NMPC).
No. 7, Chomiak, 5.IX.1908, leg. Dziędzielewicz (1 male, aedeagus and para-
meres on slide No. K7, NMPC). No. 8, Chomiak, 5.IX.1908, leg. Dziędzielewicz
(1 male, aedeagus and parameres on slide No K8, NMPC). No. 9, Chomiak,
5.IX.1908, leg. Dziędzielewicz (1 male, aedeagus and parameres on slide No K9,
NMPC). No. 10, Chomiak, 5.IX.1908, leg. Dziędzielewicz (1 male, aedeagus and
parameres on slide No K10, NMPC).” No. 20, Chomiak (Blotek), 22.IX.1906, leg.
Dziędzielewicz (1 female, NMPC). Bieszczady Mts (Besszádok), Ung National
Park, below Lubnya (Kiesvölgy), N: 49°00’ 54,81” E: 22°43’ 23,82”, 478 m, sin-
gled, 20. IX. 2013, leg. J. Oláh, Cs. Balogh, Cs. Deák & I. Meszesán (1 fe-
male, OPC).”
Isogamus balinti Oláh, 2015

Isogamus aequalis aequalis Klapálek, 1907: Botosaneanu (1961a: 60):
records from Southern Carpathians. Misidentifications.
Isogamus balinti Oláh, 2015 in Oláh et al. (2015b: 43): “Holotype. Romania,
Southern Carpathians, Parang Mts. Calcescu Lake, 45.356 23.614, 1802 m,
3.VIII.2004, leg. L. Ujvárosi (1 male, OPC). Paratypes. Same as holotype (1 male,
OPC). Southtern Carpathians, Ieser Mts, 45.45 25.02, 1050m, 3.VIII.2006, leg.
M. Bálint (2 males, OPC).”
Diagnosis – Oláh et al. (2015b: 43): “Sibling species of I. baloghi having
truncate, concave gonopod apex in lateral view; dorsally convex paraproct; par-
ameres with less developed spine pattern of the modified setae; shaft terminal
slender spine-like. Differs from its sibling I. baloghi by having high paraproct in
lateral view, not slender, paramere dorsum convex, not straight; reduced spine-
like setae located ventrad, not laterad. Female unknown.”
Etymology – “We dedicated this species to Miklós Bálint, one of the collector
who has accompanied the senior author (J.O.) in his first Romanian Carpathian
collecting trip.”
Isogamus baloghi Oláh, 2015

Isogamus aequalis aequalis (Klapálek, 1907): Botosaneanu (1961b: 60):
records from Rodnei Mts. Misidentifications.
Isogamus baloghi Oláh, 2015 in Oláh et al. (2015b: 44–45): “Holotype.
Romania, Rodnei Mts. Iza stream, side spring with sphagnum bog, N47°36’19.3”

E24°31’53.4”, 993m 27. IX. 2014, leg. J. Oláh & Cs. Balogh (1 male, OPC).
Allotype: same as holotype (1 female). Paratype: same as holotype (14 males, 3
females). Rodnei Mts. Numerous spring streamlets on the spring area of Cailor
waterfall, Piatra Rea, N47°35’1.9” E24°47’49.4”, 1564m, 28. IX. 2014, leg. J. Oláh
& Cs. Balogh (1 male, OPC).”
Diagnosis – Oláh et al. (2015b: 44–45): “Sibling species of I. balinti hav-
ing truncate, quadratic gonopod apex in lateral view; dorsally straight paraproct;
parameres with less developed spine pattern of the modified setae; paramere
shaft terminal slender spine-like. Differs from its sibling I. balinti by having slen-
der paraproct in lateral view, not broad, paramere dorsum straight, not convex;
reduced spine-like setae on the paramere located laterad, not ventrad. Female
lateral lobes of tergite IX long triangular, not short and not blunt like at I aequa-
lis; basal region of tergite IX shouldered and without subbasal dark crossline,
not without lateral shoulder and not with pronounced subbasal transversal line
present at female of I. aequalis.”
Etymology – “We dedicated this species to Csaba Balogh, one of the collec-
tor who has accompanied the senior author (J.O.) in several Carpathian collect-
ing trips.”
Isogamus czarnohorensis (Dziędzielewicz, 1912)

Anisogamus aequalis Klapálek var. czarnohorensis Dziędzielewicz, 1912:
137–138: “Odmiana ta pojawia się w malych gromadkach na Czarnohorze
wysokości od 1300–1700 m n. p. m. przy potokach i źródlach w czasie od polowy
sierpnia poza połowę września. Samec latają w świetle słonecznem i okrążają
drzewa świerkowe lub krzewy kosodrzewu, na których usiadają, samice zaś
ukrywają się w gęstwinach nizkiej róślinności lub między głazami na brzegach
wody i popszukując źródeł, przy których najchętniej składają jaja, wydostają się
do znacznej wysokości.”
Isogamus aequalis czarnohorensis (Dziędzielewicz, 1912): Schmid (1955:
183–184): transfered to his newly erected genus Isogamus.
Isogamus czarnohorensis (Dziędzielewicz, 1912): Szczęsny (1980: 466–
471): elevated to species rank. Lectotype and paralectotypes designated, label-
ling: Czarnohora, Breskul 25.IX.1910. In NHM-ISEA: “In the Cracow collection
I found 6²² and 4³³ collected by Dziędzielewicz and 1³ collected by Fudakowski
in the Eastern Carpathians; 4²² with inventory numbers 37/24, 38/24 and 39/24
were wrongly identified as “Stenophylax millenii Klap.” and 1³ with inventory
number 2/25 as “Anisogamus aequalis Klap.” Dziędzielewicz described this form
on the basis of specimens caught by the streams and springs of Czarnohora in the
sub-alpine and alpine levels between 1300 and 1700 m asl., during the months

226 J. Oláh
of August and September. It happened probably in the year 1910 as the speci-
mens dated from this time are already identified as “Anisogamus aequalis Klap.
czarnohorensis Dz.”. In the Cracow collection are 2 ²² and 2³³ which come from
this period bearing the same inventory number 1/25, I believe that these may be
called the typical series. I am choosing from these the ² lectotype; the specimen
has the labelling “Czarnohora, Breskul 25.IX.1910” and “Isogamus aequalis czar-
nohorensis (Dz.) ver. C. Tomaszewski”. The abdomen of the lectotype as well as
the paralectotypes are macerated in KOH.” Differential diagnoses of both sexes
of I. aequalis and I. czarnohorensis from the Ukrainian Carpathians were elabo-
rated and detailed drawings prepared.
Isogamus czarnohorensis (Dziędzielewicz, 1912): Szczęsny & Godunko
(2007: 36–37): in SMNHL: “7²² and 2³³; East Carpathians: Czarnohora Massif
(1200 m a.s.l.); all the specimes match the criteria for paralectotypes (No 01–09).
Several paralectotypes are stored also in NMP (Chvojka, pers. comm.). The tax-
onomical status of this taxon in opriginal description was Anisogamus aequalis
Klap. var. czarnohorensis which corrseponds to the rank of subspecies (Schmid
1955). Dziędzielewicz in the description ephasized the differences in body and
wing color and size of wings between parental forms, aequalis and czarnogoren-
sis. The specific wing color pattern as a main character of czarnohorensis is also
stressed by Raciecka (1934). Subsequent, more careful studies of genital structure
of both taxa allowed the status of latter (Szczęsny 1980)”.
Isogamus czarnohorensis (Dziędzielewicz, 1912): Szczęsny & Chvojka
(2008: 157): in NMP: Altogether 21²² and 3 ³³ specimens were collected dur-
ing 9–12.IX.1908 in the East Carpathians from the type locality and deposited
in NMP.
Isogamus czarnohorensis (Dziędzielewicz, 1912): Oláh et al. (2015b: 45–46):
“Romania. Maramures county, Maramaros Mts. Hututeanca stream, 1020 m, N47°
52’ 27’’ E24° 20’ 31’’, 7.VIII.2012, leg. J. Oláh & L. Szél (1 male, OPC). Maramures
Mts. Bistra stream valley, small spring with sphagnum bog, N47°53’1.3”
E24°16’47.7”, 609m, 27. IX. 2014, leg. J. Oláh & Cs. Balogh (1 male, OPC).
Ukraine: Klapálek Collection: K218, Forestanka, 12.IX.1908, leg. Dziędzielewicz
(1 male, NMPC). K219, Porzyszewska, 9.IX.1908, leg. Dziędzielewicz (1 male,
NMPC). K220, Pod Turkul, 10.IX.1908, leg. Dziędzielewicz (1 male, NMPC).
No. 22, Pod Turkul, 10.IX.1908, leg. Dziędzielewicz (1 female, NMPC). No. 23,
Pod Breskul, 11.IX.1908, leg. Dziędzielewicz (1 female, NMPC). No. 24, Pod
Dancerz, 10.IX.1908, leg. Dziędzielewicz (1 female, NMPC). No. 12, Pod Turkul,
10.IX.1908, leg. Dziędzielewicz (1 male, aedeagus and parameres on slide No
K12, NMPC). No. 11, Pod Turkul, 10.IX.1908, leg. Dziędzielewicz (1 male, ae-
deagus and parameres on slide No K11, NMPC).”

Remarks – Oláh et al. (2015b: 45–46): “According to the original description

(Dziędzielewicz 1912) and confirmed by Raciecka (1934) and Szczęsny (1980) this
species differs from the closely related sibling species of I. aequalis by its darker
colour. We have collected specimens in Romania in sunny environment and the
shining dark coloration of the freshly collected living specimens was remarkable.”
Isogamus lineatus (Klapálek, 1903)

Anisogamus lineatus Klapálek, 1903a: 1–2: “Ein ², Styrie, Scheibleregger
Hochalpe, leg. Strobl”.
Anisogamus lineatus Klapálek, 1903b: 1–2: “Mám před sebou jediný kus ²
ze Scheiblereger Hochalpe (3./6.98, Strobl)”.
Isogamus lineatus (Klapálek, 1901): Botosaneanu (1967b: 103): trans-
ferred to genus Isogamus. “L’exemplaire type (que nous designons ici comme lec-
totype ²; conservé à sec; abdomen en préparation au baume du Canada, attachée
à l’épingle supportant la reste de l’insecte), provient de Styrie, Scheibleregger
Hochalpe, 3. VI. 1898 (Strobl). Nous ne savons rien du milieu ou se dévelop-
pent les larves. Notre material comprend 38 ² des Mts. de Fagarash (Carpates
Méridionales de Roumanie). Ces insectes furent capturés le 7. VI. 1961, le 8. VIII.
1960, extre les 7.–9. VIII. 1962, ainsi que le 16. IX. 1964. Il s’agit donc d’une
espèce “on the wing” durant le printemps, l’été et au moins la premiére moitié
de l’automne alpine. Les stations sont des sources, des ruisseaux et des torrents,
dans la zone alpine, dans celle subalpine et à la limite supérieure de la forêt de
conifères, eaux appartenant aussi bien à des bassins du versant N. que du ver-
sant S. du massif (Podu Giurgiului – Buda, Podrag – Arpas, Bîlea – Cîrtzisoara,
Sîmbata). Les altitudes varient de 1500 à 2300 m.”
Isogamus lineatus (Klapálek, 1901): Malicky (2005b: 576): “Nach Angabe in
der Beschreibung stammt der Holotypus von der Scheibleregger Hochalpe bei Ad-
mont (Steiermark). Diese Art ist sonst nie mehr in den Alpen, sondern immer nur
in Südkarpaten gefunden worden. Ich habe mehrere Jahre hindurch wiederholt die
Scheibleregger Hochalpe besucht und dort zu allen Jahreszeiten gesammelt, auch
viele Larven aus einzigen dort fl iessenden Bach mitgenommen und im Labor zu
Adulten gezüchtet. Dabei ich niemals einen Isogamus gefunden. Offenbar ist en-
tweder Klapálek oder dem Sammler Strobl eine Etikettenverwechslung unterlaufen.
Beide haben auch in der Karpaten gesammelt. Die Art kommt in Alpen nicht vor.”
Isogamus lineatus (Klapálek, 1901): Oláh et al. (2015b: 46–47): “Romania.
Argeş county, Făgăraş Mts, Căpătânenii Ungureni, small springlake by the Capra
Stream along road No.7C, N45°34.605’ E24°37.060’, 1405m, 29.VIII.2012 leg. T.
Kovács, D. Murányi, J. Oláh (2 males, 5 females; OPC). Argeş county, Făgăraş Mts,
Căpătânenii Ungureni, sidebrook of Capra Stream along road No.7C, N45°35.185’

228 J. Oláh
E24°37.691’, 1705m, 29.VIII.2012 leg. T. Kovács, D. Murányi, J. Oláh (8 males, 7 fe-

males, 1 copula; OPC). Sibiu county, Făgăraş Mts, Cârţişoara, forest seep along road
No.7C, N45°38.742’ E24°36.464’, 1325m, 29.VIII.2012 leg. T. Kovács, D. Murányi,
J. Oláh (3 females, OPC). Braşov county, Făgăraş Mts, Dejani, forest sidebrook of
Dejani stream, N45°36.408’ E24°56.466’, 1310m, 30.VIII.2012 leg. T. Kovács, D.
Murányi, J. Oláh (8 males, 5 females, OPC). Braşov county, Făgăraş Mts, Dejani,
right sidebrook of Dejani stream, N45°35.446’ E24°56.348’, 1755m, 30.VIII.2012
leg. T. Kovács, D. Murányi, J. Oláh (1 male, 1 female, OPC). Braşov county, Făgăraş
Mts, Dejani, forest sidestream of Dejani stream, N45°36.720’ E24°56.533’, 1250m,
30.VIII.2012 leg. T. Kovács, D. Murányi, J. Oláh (5 males, 3 females, OPC).”
Remarks – Oláh et al. (2015b: 46–47): “This species was described from
Austrian Alp (Styria), but never collected there, despite systematic search, but
found very common in the South Carpathians. This confusion was probably re-
sulted by mistakes in the original labelling (Malicky, 2005).”
Melampophylax Schmid, 1955

Melampophylax austriacus Malicky, 1990
Melampophylax austriacus Malicky, 1990: 8: “Material: Wechsel oberhalb
von Mariensee (Niederösterreich), 900 m, 24.X.1985 – 1² (Holotypus), 15³³
(paratypen). Weitere Paratypen: selber platz, aber 26.X.1977 – 1³; Mönichner
Schwaig, 1100 m, 26.X.1977 – 6³; Steiermark: Vorauer Schwaig (Wechsel), 1500
m, 26.X.1977 – 5², 4³; do 24.X.1985 – 14³; Teichalpe, leg Hölzel, 2.X.1965 – 2²,
3³; do 20.X..1963 – 3²; do 2.X.1966 – 4²; do20.X.1968 – 2²; Reinischkogel,leg.
Hölzel,6.X.1968 – 2²; do 13.X.1968 – 6², 1³; Klugveitl, leg. Hölzel, 2.X.1969
– 2², 1³. Kärnten: Saualpe, Ladinger Alm, 1800 m, 28.IX.1988 – 3², 2³.
Slowenien: Pohorje, leg. Sivec, 20.X.1981 3³. Die Tiere von der Teichalpe, vom
Reinischkogel (ausser 1²) und Klugveitl (ausser 1³ in coll. Hölzel; je 1 Pärchen
vom Vorauer Schwaig in der Zoologischen Staatssammlung München und im
Forschunginstitut Senckenberg Frankfurt; alle anderen in meiner Sammlung.”
Melampophylax austriacus Malicky, 1990: Oláh et al. (2015b: 54): “Austria,
Schwarze Sulm, 20. X. 2013 leg. W. Graf (1 male, 1 female; OPC).”
Remarks – Oláh et al. (2015b: 54): “Auxilliary mesal elongation of the go-
nopods blunt broad triangular. Terminal blade on paramere sagittaly flattened.
Elongated vaginal sclerite complex broad basad and narrowing distad.”
Melampophylax banaticus Botosaneanu, 1995

Melampophylax nepos ssp. banaticus Botosaneanu, 1995: 76.
Melampophylax nepos ssp. banaticus Botosaneanu, 1995: Malicky (2005b:
577).

Melampophylax banaticus Botosaneanu, 1995: Oláh et al. (2015b: 50–

51): elevated to species status. “Romania, Caraş-Severin county, Ţarcu Mts.,
open brook on the W slope of Mt. Ţarcu, N45°17’30.9”, E22°30’59.9”, 1770 m,
14.10.2011, Á. Ecsedi, T. Kovács & G. Puskás (4 males, 2 females; OPC). Retezat
Mts. Gura Apelor, N45.33 E22.88, 1500 m, 20.X. 2007 leg. M. Bálint, E. Magyari
& M. Braun (1 male, OPC).”
Remarks – Oláh et al. (2015b: 50–51): “This is an incipient sibling species
closely related M. polonicus and M. gutinicus, but differs from both by having fe-
male with differently shaped vaginal sclerite elongation and males with pointed
mesal corner on the mesal plate of the gonopods. Also differs by having the speci-
ation trait, the paramere very stout with shallow curvature and with pronounced
mace-like terminal blade.”
Melampophylax gutinicus Botosaneanu, 1995

Melampophylax nepos ssp. gutinicus Botosaneanu, 1995: 75–76.
Melampophylax nepos ssp. gutinicus Botosaneanu, 1995: Malicky (2005b:
577).
Melampophylax banaticus Botosaneanu, 1995: Oláh et al. (2015b: 50–51):
elevated to species status. “Romania, Maramureş county, Muntii Ignis, Deseşti-
Staţiunea Izvoare, open brook on the Valhani Plateau, 1020m, N47°43.015’
E23°44.547’, 07.10.2010, leg. P. Barcánfalvi, D. Murányi & J. Oláh, (5 males, 2
females, OPC). Maramureş county, Munţii Ignis, Deseşti-Staţiunea Izvoare, open
stream on the Valhani Plateau, 940m, N47°43.945’ E23°44.661’, 08.10.2010, leg.
P. Barcánfalvi, D. Murányi & J. Oláh, (1 male, OPC). Maramureş county, Munţii
Ignis, Deseşti-Staţiunea Izvoare, spring brook on the Valhani Plateau, 955m,
N47°44.177’ E23°43.971’, 08.10.2010, leg. P. Barcánfalvi, D. Murányi & J. Oláh,
(4 males, OPC). Radnei Mts. Rodnei Mts. small tributary just below Iza Spring,
Albastru al Izei, 1020m, 27. IX. 2014, leg. J. Oláh & Cs. Balogh (1 male, OPC).”
Remarks – Oláh et al. (2015b: 50–51): “Incipient sibling species closely re-
lated M. banaticus and M. polonicus, but differs from both by having female with
differently shaped vaginal sclerite elongation, that is short and narrowing distad
and the male has the most rounded mesal corner on the mesal plate of the gono-
pods as well as very slender paramere that has minute terminal blade.”
Melampophylax polonicus Malicky, 1990

Melampophylax polonicus Malicky, 1990: 8–9: “Material: Polen,
Bieszczady-Gebirge, Polaniec, 600–1000 m, 22.X.1976 – 1² (Holotypus), 2³
(Paratypen); do. Wolosatka, 21. X. 1974 – 1³ (paratype) in coll. Malicky. 3 ² mit
den gleichen Funddaten in coll. Szczęsny (Paratypen).”

230 J. Oláh
Melampophylax polonicus Malicky, 1990: Oláh et al. (2015b: 54): “Poland,

Bieszczady Mts. X. (October, without day and year), leg. det. B. Szczęsny (2 males,
1 female; OPC).”
Remarks – Oláh et al. (2015b: 54): “Incipient sibling species closely related
M. banaticus and M. gutinicus. Differs from both by having female with differ-
ently shaped vaginal sclerite elongation with produced subapical constriction;
males with straight truncate apical margin on the mesal plate of the gonopods as
well as long and strong paramere that has strong terminal blade.”
Melampophylax triangulifera Botoşăneanu, 1957

Melampophylax nepos triangulifera Botoşăneanu, 1957d: 400–401.
Melampophylax nepos ssp. triangulifera Botosaneanu, 1957: Malicky (1990:
2, 2005b: 576).
Melampophylax nepos ssp. triangulifera Botosaneanu, 1957: Botosaneanu
(1995: 74–75): female described.
Melampophylax triangulifera Botosaneanu, 1957: Oláh et al. (2015b:
57): elevated to species status. “Romania, Eastern Carpathians, Hargitha Mts.,
springs and streamlets between Baile Hargitha and Cabana Madaras, 1650–1700
m, 14. X. 1970 leg. L. Botosaneanu (6 males, OPC). Gurghiu Mts. near Bucin
Pass, Tárnava Mica springs and stream, N: 46°39’ 16,63”E: 25°16’ 42,46”, 1290,
30.X.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés & J. Oláh. (49 males, 6 females;
OPC). Gurghiu Mts. near Bucin Pass, Gainasa springs and stream, N: 46°40’
11,35” E: 25°17’ 39,06”, 1400, 30.X.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés &
J. Oláh (1 male, 1 female; OPC). Hargitha Mts. Filio stream side spring, N: 46°27’
03,90” E: 25°33’ 29,29”, 1350m, 31.X.2014 leg. Z. Baczó, Cs. Balogh, J. Kecskés
& J. Oláh. (1 male, 3 females; OPC). Caliman Mts. Fantanele stream, N: 46°59’
04,00” E: 25°05’ 52,56”, 776m, 1.XI.2014, leg. Z. Baczó, Cs. Balogh, J. Kecskés &
J. Oláh. (3 females; OPC).”
Remarks – Oláh et al. (2015b: 57): “This is a sibling species of M. nepos and
M. szczesnyorum. Auxilliary mesal elongation of the gonopods laterad curving.
Terminal blade of the paramere robust and rounded. Elongated vaginal sclerite
complex rather variable inside populations.”
Melampophylax szczesnyorum Oláh et Chvojka, 2015

Melampophylax nepos triangulifera Botosaneanu, 1957: Szczęsny (1980:
465–466): misidentification.
Melampophylax szczesnyorum Oláh et Chvojka, 2015 in Oláh et al. (2015b:
56–57): “Holotype- Ukraine, original label: “Chomiak, (pot. podziemny),
6.X.1905, leg. Dziędzielewicz (1 male, Klapálek’s Collection in NMPC: No. 50).

Allotype: Ukraine: original label: “Chomiak. p. Weredyk, 1–10.–1907, leg Dz.”

(1 female, Dziędzielewicz’s collection in NHM-ISEA). Paratypes. Ukraine, origi-
nal label: “Tatarow (Prutec), 7.–10.1905 Dz” (1 male, Dziędzielewicz’s collection
in NHM-ISEA). Ukraine: original label: “Chomiak, p. Weredik, 1.–10.-1907
Dz” (1 male, Dziędzielewicz’s collection in NHM-ISEA). Ukraine: original la-
bel: “Chomiak (pot. podziemny), 6.–10.-1905 Dz” (1 male, Dziędzielewicz’s col-
lection in NHM-ISEA). Ukraine: original label: “Tatarov, (Blotek.) X. 1906” (1
male, Dziędzielewicz’s collection in SMNHL: No. 1385). Ukraine: original label:
“Czarnohora, Foreszczynka 4.X. 1910” (1 male, Dziędzielewicz’s collection in
SMNHL: No. 1399). Ukraine: original label: “Czarnohora, Foreszczynka 19.X.
1910” (1 male, Dziędzielewicz’s collection in SMNHL: No. 1402). Ukraine: origi-
nal label: “Czarnohora, Kozmieska 16.X. 1908” (1 male, Dziędzielewicz’s col-
lection in SMNHL: No. 1396). Ukraine: original label: “Chomiak, Barani. 9.X.
1907” (1 female, Dziędzielewicz’s collection in SMNHL: No. 1389). Ukraine:
original label: “Chomiak, Prutec. 2.X. 1912” (1 female, Dziędzielewicz’s col-
lection in SMNHL: No. 1403). Ukraine: original label: “Chomiak, Prutec. 2. X.
1912” (1 male, Dziędzielewicz’s collection in SMNHL: No. 1401). Ukraine: origi-
nal label: “Chomiak, P. Weredyk. 7. X. 1907” (1 female, Dziędzielewicz’s collec-
tion in SMNHL: No. 1386). Ukraine: original label: “Czarnohora, Foreszczynka
4.X. 1910” (1 male, Dziędzielewicz’s collection in SMNHL: No. 1397). Ukraine:
original label: “Chomiak, Gnilec. 5. X. 1907” (1 male, Dziędzielewicz’s collec-
tion in SMNHL: No. 1394). Ukraine: original label: “Chomiak, P. Weredyk. 23.
IX. 1907” (1 male, Dziędzielewicz’s collection in Dziędzielewicz’s collection in
SMNHL: No. 1388). Ukraine: original label: “Chomiak, (Blotek.) 22. IX. 1906”
(1 female, Dziędzielewicz’s collection in SMNHL: No. 1392). Ukraine: original la-
bel: “Chomiak, Potok Barani. 18. X. 1907” (1 female, Dziędzielewicz’s collection in
SMNHL: No.1384).”
Diagnosis – Oláh et al. (2015b: 56–57): “This new species differs from its
close relatives, from the sibling species of M. nepos and M. triangulifera by having
the auxilliary mesal elongation of the gonopods with straight apex, mesad turn-
ing at M. nepos and laterad turning at M. triangulifera; the terminal blade of the
paramere produced into a very flattened shape, a unique and powerful device in
copulation; the elongated vaginal sclerite complex broadening distad. Only his-
torical material was available for this study. No newly collected material!”
Etymology – “We dedicated this species to Dr. Bronislav Szczęsny and
his wife, who has produced significant contribution to the knowledge of the
Trichoptera in the North-East Carpathians covering both Poland and Ukraine.”
Potamophylax Wallengren, 1891

Potamophylax borislavi Kumanski, 1975

232 J. Oláh
Potamophylax borislavi Kumanski, 1975a: 85–87: “Holotype ²: (pinned,

abdomen in glycerine, attached to the same needle) Bulgaria, the Rhodops near
Polkovnik Serafimovo village (about 1100 m), 4. IX. 1970, S. Zagortshinov leg.
(by light).”
Further records – Bulgaria, Rhodope Region Rozsen, 13.IX.1979, 1 ² (leg. J.
Ganev) (Kumanski 1983). Rila Mts. River Slavova, above the Tchaira-Dam, ca
1350 m, 22–23.VIII.1997, 1², 1–2.X.1997, 2² leg. Kumanski at light (Kumanski
& Malicky 1999). Greece, near Dasikon Chorio, in Elatia area 1300m, 24°20’E
41°30’N, 10. X. 1991, 2 ², leg H. Malicky (Malicky 2005a).
Potamophylax carpathicus (Dziędzielewicz, 1912)

Stenophylax carpaticus Dziędzielewicz, 1912: 136–137: “In Montibus
Carpaticis Orientalibus sub cacuminibus: Chomiak ad Tatarów, Rebrowach in
Worochta et in regione alpina Czarnohorae in altitudine 800–1400 m s m. apud
fontes 12 VI– 6 VII rarus.”
Potamophylax carpathicus (Dziędzielewicz, 1912): Schmid (1955: 176):
transferred and listed in genus Potamophylax.
Potamophylax carpathicus (Dziędzielewicz, 1912): Szczęsny (1980: 463): in
NHM-ISEA: “1², 3³³; 2³³ were labelled wrongly as “Stenophylax millenii Klap.”
(now known as Potamophylax millenii (Klap.), their labels had the following notes:
nr. 16/20 – “Chomiak (Blotek) 3.VII.1905 Dz.” Nr. 43/23 – “Chomiak Potok
Barani zrodla 27. VI. 1907”. The two remaining specimens had identical inventory
numbers 6/25, they were named correctly and bore the following labelling: ² –
“Worochta 12. VI. 1911”, ³ (without the last abdominal segments) – “Worochta
24.VI.1911 pod Rechaczem”. The species was described (Dziędzielewicz, 1912) on
the basis of the specimens collected “near the summits of Chomiak and Rebrowacz
at Worochta during the period 12. VI. – 6. VII. 1911”. I beleive, the last two speci-
mens come from the typical series and so satisfy the conditions for syntypes. I
am designating as a lectotypus the male of nr. 6/25 with an adduitional labelling
“Potamophylax carpathicus (Dz.) ² ver. C. Tomaszewski.”
Potamophylax carpathicus (Dziędzielewicz, 1912): Szczęsny & Godunko
(2007: 37): in SMNHL: “5²², 6³³; West Carpathians: Beskid Wyspowy, East
Carpathians (5²², 5³³): Czarnohora Massif (Worochta), Gorgany Massif
(Chomiak) – of those collectedi n East Carpathians, 2²², 3³³ are paralectotypes
according to the ICZN Article 74.1.4 (No 24.12.23.01/01–05), but 3²², 2³³ ori-
gin from locus typicus. Dziędzielewicz did not mention the series of specimens
being the basis for species description. He informed only in which localities the
species was observed, i.e.: Chomiak Mt at Tatarow, Rebrowacz Mt at Worochta
and Czarnohora Massif, 12. VI – 6. VII. As a matter of fact, he had not recog-

nised correctly the species because all the earlier (until 1908) collected speci-
mens, both I. czarnohorensis and P. carpathicus, were labelled as “Stenophylax
millenii Klap. det Dz.” (those stored in MP ISEZ). The museum specimens stored
in Lviv were labelled “Stenophylax carpathicus Dz. Det. M.R. (Maria Raciecka)”
or “Potamophylax carpathicus Dz.” – name changed recently.”
Potamophylax carpathicus (Dziędzielewicz, 1912): Szczęsny & Godunko
(2008: 74): the Khomiak Mts. is locus typicus for the species.
Potamophylax jungi Mey, 1976

Potamophylax jungi Mey, 1976: 166–167: “Holotypus: 1² Rumanien,
Schupichbach südlich von Sibiu, 31.VII.1975; in coll Mey. Paratypus 1²:
Rumanien, Nebenbach des Dragan bei Ciucea, 26.VIII.1975; in coll Botosaneanu.”
Potamophylax millenii (Klapálek, 1898)

Stenophylax millenii Klapálek, 1898a: 488: “Mehádia, Carniareva,
Ferenczfalva im Krassó-Szörényer Comitat. 3², 1³.”
Stenophylax millenii Klapálek, 1898: Klapálek (1899a: 431–432).
Potamophylax millenii (Klapálek, 1898): Schmid (1955: 177): transferred
and listed in genus Potamophylax.
Potamophylax seprus Oláh, Lodovici et Valle, 2011

Poptamophylax seprus Oláh, Lodovici et Valle, 2011: 13: “Holotype
male. Albania: Skrapar county, Tomor Mts, Kulmak Pass, mountain grassland
near the bektashi teqe, N40°37.116’ E20°11.945’, 1485m, 23.VIII.2006, leg. Z.
Fehér, A. Hunyadi, T. Huszár & D. Murányi (1 male, HNHM).”
Potamophylax nigricornis species group

Potamophylax elegantulus (Klapálek, 1899)
Stenophylax nigricornis var. elegantulus Klapálek, 1899b: 325: “Lako ga je
razlikovati od tipićne forme po jako iztaknutim svjetlim prugama na tamnoj os-
novnoj boji krila. Ćitav niz ekzemplara sa Vrela Bosne 16 maja, i Trebevića (Win-
negth.).”
Potamophylax nigricornis elegantulus (Klapálek, 1899): Schmid (1955: 176):
transferred to genus Potamophylax.
Potamophylax elegantulus (Klapálek, 1899): Oláh et al. (2013a: 184–185):
raised to species status. “The original description in 1899 was based on 6 males
and 2 females specimens without any type labels. Here we have designated lecto-
type and selected allotype. Lectotype. Bosnia-Herzegovina: Vrelo Bosna, 16 May

234 J. Oláh
(1 male, K369, NMPC). Allotype. same as lectotype (1 female: K373, NMPC).

Paralectotypes. same as lectotype (1 male: K371, 1 male: K264, 1 male: K265,
1 male: K370, 1 male: K372; NMPC). Trebević leg. Winnegth (1 female, K374,
NMPC). New record. Bosnia-Herzegovina: Toplice, N43°35.658 E18°29.697,
995 m 3.VI.2009, leg. W. Graf, (1 female, OPC).”
Diagnosis – Oláh et al. (2013a: 184–185): “This species having dark striped
forewing was described as a variant. The original description of Potamophylax
nigricornis var. elegantulus is based upon the characteristic forewing pattern.
Klapálek has not cleared the abdomen and thus he was unable to detect the sig-
nificant modifications in the structure of the phallic organ. The ability of the
aedeagal head to extend so much laterad during erection has been never observed
at any specimens of P. nigricornis. Compared to P. nigricornis the parameres has
lost the basal quadratic basement of the shaft ; sigmoid pattern reduced; the basal
tuft of 5 strogly mesad curving setae shortened, straightened, doubled to 10 and
spread upto the end of the shaft ; apical seta fused to the end of the shaft , alveo-
lus almost indiscernible. Female vaginal sclerite complex rounded, not elongated
longitudinally and has short sclerotized opening on the spermathecal process,
not long; transversal bursal slerite long, not short.”
Potamophylax fules Oláh et Ibrahimi, 2013

Potamophylax fules Oláh et Ibrahimi, 2013 in Oláh et al. (2013a: 185–186):
“Holotype. Romania: South Carpathians, Caras-Severin county, Tarcu Mts. Poina
Marului, upper section of Sucu Stream, S of the village, N45°20.907’ E22°31.073’,
955 m, 08.VI.2011, leg. T. Kovács, D. Murányi & G. Puskás, (1 male, HNHM).
Paratypes. Kosovo: Skënderaj Municipality, entrance into the Kuçicë village, Klinë
river sidespring, N42°48.36’ E20° 46.29’, 690 m, 12.V.2011, leg. H. Ibrahimi, Ar.
Gashi & B. Deliu (1 male, DBFMNSUP). Prishtinë Municipality, Gollak region,
Keqekollë village, streamlet, N42.7237°, E21.3067°, 804 m, 15.06.2009, leg. H.
Ibrahimi & F. Asllani Ibrahimi (1 male, DBFMNSUP). Montenegro: Biogradska
Gora, N42 54 01.5 E19 35 44.7, 1093 m, 29.V.2009, leg W. Graf (1 male, OPC).”
Diagnosis – Oláh et al. (2013a: 185–186): “This species having dark striped
forewing belongs to the Potamophylax elegantulus species subgroup. Close to P.
ureges sp. n. but differs by having the ventral subapical double layered heel with
lateral plates abbreviated; the lateral plate is longer at P. fules; as a result the mesal
plate is long exposed free in lateral view; pectinate patterned setae distributed
along to the subapical region, not limited to the basal half of the paramere shaft.”
Etymology – “Fules from “füles” auriculate in Hungarian, refers to the ear-
shaped small corner produced by the abbreviated lateral plate of the double-lay-
ered ventral subapical heel on the aedeagus.”

Potamophylax hasas Oláh, 2013

Potamophylax nigricornis (Pictet, 1834): Kumanski (1971: 103): Bulgaria
(Rodope Mts). Misidentification.
Potamophylax hasas Oláh, 2013 in Oláh et al. (2013a: 186): “Holotype.
Bulgaria: Smoljan Province, Perelik Mts. Pamporovo, open brooks and alpine
grassland at the settlement, 1560 m, N41°37.540’ E24°42.411’, 31.V.2012, leg.
J. Kontschán, D. Murányi & T. Szederjesi (1 male, OPC). Allotype. Bulgaria,
Rodope Mts. above Triglad 25. VII. 1968, leg. A. Slivov (1 female, NMNHBAS)
Paratype. Same as allotype (1 male, NMNHBAS).”
Diagnosis – Oláh et al. (2013a: 186): “Close to P. kethas sp. n. but differs by
having single belly on the aedeagus without a strong constriction midway, apical
seta on paramere long, not short; basal setae four, not eight. Kumanski’s original
male and female specimens were examined designated as allotype and paratype
and compared to the holotype.”
Etymology – “Hasas from “hasas” bellied in Hungarian, refers to the subapi-
cal lateral bulging on the aedeagus.”
Potamophylax kethas Oláh, 2013

Potamophylax kethas Oláh, 2013 in Oláh et al. (2013a: 186–188): “Holo-
type. Bosnia-Harcegovina: Vucjaluka, N43.93221 E18.52135, 12.VII.2008, leg.
M. Bálint & S. Lelo (1 male, OPC).”
Diagnosis – Oláh et al. (2013a: 186–188): “Close to P. hasas sp. n. but dif-
fers by having double bellies on the aedeagus with a strong constriction midway,
apical seta on paramere short, not long; basal setae eight, not four.”
Etymology – “Kethas from “két has” double bellies in Hungarian, refers to
the strong constriction midway on the aedeagus resulting double bellies in ven-
tral view.”
Potamophylax lemezes Oláh et Graf, 2013

Potamophylax lemezes Oláh et Graf, 2013 in Oláh et al. (2013a:
188):“Holotype. Macedonia: Mavrovo, Galicnicka spring, N41.35364, E20.39523,
1407m, 2.VII.2010, leg. W. Graf (1 male, OPC). Allotype. Same as holotype (1 fe-
male, OPC). Paratypes. Same as holotype (1 males, 2 females; OPC). Vitosa, 27.
V. 2010, leg. I. Sivec (1 male, 1 female; OPC).”
Diagnosis – Oláh et al. (2013a: 188): “This species having no double layered
heels (P. ureges, P. fules), no bellies on aedeagus (P. hasas, P. kethas), no elongated
pointed heels (P. apados) and no broad paramere shaft (P. schmidi) has resem-
blance in the subgroup to P. elegantulus, but differs by having the unique lateral

236 J. Oláh
plate developed on the aedeagus subapicad, and by the differently patterned setal
structure on the paramere shaft.”
Etymology – “Lemezes” from “lemezes” laminate in Hungarian, refers to lat-
eral laminar plates present subapicad on the aedeagus.”
Potamophylax schmidi Marinković-Gospodnetić, 1971

Potamophylax schmidi Marinković-Gospodnetić, 1971a: 80–83: South-
east Bosnia.
Potamophylax schmidi Marinković-Gospodnetić, 1971: Marinković-
Gospodnetić (1971b: 144): “Southeast Bosnia, ²² ³³ in many small brooks on
the mountain Zelengora and Maglić.”
Potamophylax schmidi Marinković-Gospodnetić, 1971: Oláh et al. (2013a:
188–189): “Bosnia-Herzegovina: Sutjeska NP, Izvor uz Cestu, N43°15.902
E18°35.56716, 1109 m. 15. V. 2008, leg. W. Graf & H. Ibrahimi (7 females, OPC).
Same but 2.VI.2009, leg. W. Graf (2 females, OPC). Same, but spring by the road,
2.VI.2009, leg. A. Previsic (1 female, OPC).”
Diagnosis – Oláh et al. (2013a: 188–189): “Type specimens of this species
were lost. We have collected only females and effort to borrow males failed. Male
drawings are reproduced from the original description and drawings. Compared
to other species of this group the reconstructed lateral view of phallic organ and
the dorsal view of the paramere with setal formation is rather particular. We have
examined the availaable 9 females. Dorsal profile of the vaginal sclerite complex
as well as the spermathecal process and the bursal sclerite are, as usual, very sta-
ble indicated by the narrow range of phenotypic variation.”
Potamophylax ureges Oláh, 2013

Potamophylax ureges Oláh, 2013 in Oláh et al. (2013a: 189): “Holotype.
Montenegro: Žabljak municipality, Sinjajevina Mts, Dobrilovina, forest stream
at the monastery, N43°01.652’, E19°24.086’, 765 m, 25.05.2013, leg. P. Juhász, T.
Kovács, G. Magos, G. Puskás, (1 male, OPC). Allotype same as paratype: (1 fe-
male, OPC). Paratypes. Same as holotype (4 males, 3 females; OPC; 1 male, 2
females, NMPC). Same but 14.VI.2012 leg. Z. Fehér, T. Kovács, D. Murányi (6
females, OPC).”
Diagnosis – Oláh et al. (2013a: 189): “This beautiful species having dark
striped forewing belongs to the Potamophylax elegantulus species subgroup.
Close to P. fules sp. nov. but differs by having the ventral subapical double layered
heel with mesal and lateral plates almost equal enclosing a concavity; the lateral
plate is shorter at P. fules, pectinate patterned setae distributed only on the basal
half of the paramere shaft, not along to subapical region.”

Etymology – “Ureges from “üreges” supplied with hollow in Hungarian, re-

fers to the concavity enclosed by the double-layered heels.”
Potamophylax winneguthi species group

Potamophylax alsos Oláh, 2015
Potamophylax alsos Oláh, 2015 in Oláh & Kovács (2015: 117–120): “Holo-
type. Macedonia, Southwestern region, Jablanica Mts, 6.5 km W of Labuništa,
open brook at Labuniško Lake, N41°16.069’, E20°31.242’, 1905 m, 10.10.2014,
leg. P. Juhász, T. Kovács, G. Puskás (1 ², OPC). Allotype. Same as holotype (1³,
OPC). Paratypes. Same as holotype (7², 3³, OPC, 2², 1³, MM).”
Diagnosis – Oláh & Kovács (2015: 117–120): “The fourth member of the
Potamophylax tagas species cluster. A sister or sibling species of P. hajlos but dif-
fers by having gonopod apical margin with backward produced lower corner, not
downward directed outgrow; paramere tip with three spines; apicomesal excision
on the aedeagus wide U-shaped, not narrow; lateral margin of the dorsal vaginal
sclerite complex short with lateral hump, not long and convex.”
Etymology – “Alsos” from “alsó, alsós”, lower in Hungarian, refers to lower
apical corner of the gonopods more produced compared to its dorsoapical cor-
ner. Here we follow to name this new species of the complex according to the
lateral shape of the gonopod.”
Potamophylax haidukorum Malicky, 1999

Potamophylax haidukorum Malicky, 1999 in Kumanski & Malicky, 1999:
28–29: “Surroundings of the Motel Hajducka voda between Doboj and Banja
Luka, 28. X. 1988, holotype and many paratypes. Vares near Sarajevo 2.XI.1988,
1 female.”
Potamophylax haidukorum Malicky, 1999: Oláh & Kovács (2012: 97–98):
female description. “Bosnia & Herzegovina: Banja Luka region, Borja Planina,
between Maslovare and Klupe, Hajduk spring, N44°35’29.2”, E17°35’50.9”, 790
m, 06.11.2012, leg. T. Kovács, G. Magos (1³, OPC; 1², 1³, MM). Banja Luka
region, Kozara Mts, forest brook below the Vrbaška–Kozarac road, N45°02.480’,
E16°54.266’, 560 m, 07.11.2012, leg. T. Kovács, G. Magos (1³, OPC). Bosnien,
Hajdučka voda, Zucht 1990, leg Malicky (1², 1³, OPC).
Notes – Oláh & Kovács (2015: 117–120): “There seems to be no clear fea-
ture for separating the females of P. haidukorum and P. winneguthi wrote Malicky
(Malicky & Kumanski 1999). After a detailed examination of both the external
and internal genital structure we have found significant differences to separate
the females of these species. The fused regions of segment IX and X, the closed
anal tube is subquadrangular on P. winneguthi and triangular on P. haidukorum

238 J. Oláh
both in dorsal and ventral view; sternite IX, the setose lateral lobes differently
shaped, longer than high on P. winneguthi and higher than long on P. haiduko-
rum; vulvar scale, the lower vulvar lip very developed on P. winneguthi and less
developed on P. haidukorum; the mesal lobe of the vulvar scale is present on P.
winneguthi and almost vestigial on P. haidukorum; the internal genital structure,
the vaginal or spermathecal sclerite complex clearly differently formed in the two
species.”
Potamophylax hajlos Oláh, 2012

Potamophylax hajlos Oláh, 2012 in Oláh & Kovács (2012: 98–100):
“Holotype: Albania: Tiranë district, Gropë Mts, Vakumonë, karst spring and
brook along the road to Elbasan, N41°15.109’, E20°05.805’, 1195 m, 11.10.2012,
leg. P. Juhász, T. Kovács, D. Murányi, G. Puskás (1², OPC). Allotype. Same as
holotype (1³, OPC). Paratypes. Same as holotype (1², OPC). Bulqizë district,
Çermenikë Mts, open brook beneath Mt. Kaptinë, N41°23.212’, E20°17.506’, 1610
m, 10.10.2012, leg. P. Juhász, T. Kovács, D. Murányi, G. Puskás (3², OPC; 2²,
MM). Mat district, Gropë Mts, brook along the Klos-Elbasan road, N of Shtyllë
Pass, N41°22.455,’ E20°05.073’, 1505 m, 11.10.2012, leg. P. Juhász, T. Kovács, D.
Murányi, G. Puskás (1², OPC).”
Diagnosis – Oláh & Kovács (2012: 98–100): “A member of the Potamophy-
lax winneguthi new species group and of the Potamophylax tagas new species clus-
ter. Most close to P. kesken sp. n. but differs by having gonopod apical margin
with downward directed outgrow and paramere tip with two spines; apicomesal
excision on the aedeagus narrow, not wide U-shaped.”
Etymology – “Hajlos from “hajló, hajlós” bending in Hungarian, refers to the
downward directed ventroapical corner of the gonopods.”
Potamophylax juliani Kumanski, 1999

Potamophylax juliani Kumanski, 1999 in Kumanski & Malicky (1999:
27): “Material studied: Ossogovo Mts. (W-Bulgaria), zpper stream of the Mlachka
river, with very small brooklets nearby (mixed coniferous and deciduous forest,
1600–1700 m a.s.l.), 10. 11. 1996, Holotype male (leg. Kumanski). The type spec-
imen deposited (in alcohol, the abdomen separated and treated in KOH) in the
collection of the National Museum of Natural History, Sofia.”
Potamophylax juliani Kumanski, 1999: Oláh & Kovács (2013: 118–119):
“Here we describe the unknown female of this sexually dimorphic species. We have
collected three females associated with two males from the same habitat. Females
are brachypterous, smaller sized similarly to other species of the Potamophylax
winneguthi speciwes group.” “Bulgaria, Kyustendil province, Osogovska planina,

beech forest and open brook at Osogovo hut, 1625m, N42°11.791’, E22°37.409’,
23.10.2013, J. Kontschán, D. Murányi, T. Szederjesi, (1², HNHM). Kyustendil
province, Osogovska planina, spruce forest, forest brook below Trite buki hut,
1520m, N42°10.463’, E22°38.066’, 23.10.2013, J. Kontschán, D. Murányi, T.
Szederjesi, (2², 3³, HNHM).”
Potamophylax kesken Oláh, 2012

Potamophylax kesken Oláh, 2012 in Oláh & Kovács (2012: 100–101):
“Holotype: Albania: Dibër district, Korab Mts, open stream above Fushë
Korabit, N41°49.215’ E20°32.738’, 1945 m, 07.10.2012, leg. P. Juhász, T. Kovács,
D. Murányi, G. Puskás (1², OPC).”
Potamophylax kesken Oláh, 2012: Oláh & Kovács (2015: 122–124):
“Albania, Dibër district, Korab Mts, 3.5 km SE of Radomirë, spring area of right
tributary of Elbini Stream, N41°48’10.9”, E20°31’27.4”, 1830 m, 11.10.2014, P.
Juhász, T. Kovács, G. Puskás (4², 8³, OPC). Dibër district, Korab Mts, 4.5 km
SE of Radomirë, open brook, N41°47’44.2”, E20°31’51.7”, 2050 m, 11.10.2014,
P. Juhász, T. Kovács, G. Puskás (7², 6³, OPC; 2², 2³, MM).” “This species was
described from a single male. In 2014 we have succeded to collect 13 males and 16
females from a new spring area in the Korab Mts. Here we describe the unknown
female. Like each member in the Potamophylax winneguthi species group the fe-
male has brachyptery.”
Diagnosis – Oláh & Kovács (2012: 100–101): “A member of the Potamo-
phylax winneguthi new species group and of the Potamophylax tagas new species
cluster. Most close to P. hajlos sp. n. but differs by having gonopod apical margin
without any downward outgrow and paramere tip with a single spines accompa-
nied with three very short basal spines; apicomesal excision on the aedeagus very
wide, U-shaped, not narrow.”
Etymology – “Kesken from “keskeny” narrow in Hungarian, refers to the
narrow gonopods in lateral view compared to the gonopod of P. tagas.”
Potamophylax tagas Oláh et Kovács, 2012

Potamophylax tagas Oláh & Kovács, 2012: 101–102: “Holotype. Alba-
nia: Dibër district, Korab Mts, spring brooks of the bog beneath Mt. Korab,
N41°47.913’, E20°33.561’, 2165 m, 07.10.2012, leg. P. Juhász, T. Kovács, D.
Murányi, G. Puskás (1², OPC). Paratypes: same as holotype (2², OPC; 1²,
MM).”
Potamophylax tagas Oláh et Kovács, 2012: Oláh & Kovács (2015: 124–
127): “Albania, Dibër district, Korab Mts, 5.5 km SE of Radomirë, spring and
open brook, N41°47’20.0”, E20°32’23.0”, 2330 m, 11.10.2014, P. Juhász, T. Kovács,

240 J. Oláh
G. Puskás (14², 22³, 1 copula, OPC; 2², 2³, MM).” “This species was described
without female. In 2014 we have succeded to collect 17 males and 24 females
from new spring area in the Korab Mts. Here we describe the unknown female.
Like each member in the Potamophylax winneguthi species group the female is
brachypterious.”
Diagnosis – Oláh & Kovács (2012: 101–102): “A member of the Potamo-
phylax winneguthi New Species Group. Potamophylax tagas sp. n. forms a new
species cluster together with P. kesken sp. n. and P. hajlos sp. n. This cluster is char-
acterized by apical margin of the gonopods without any significant projections;
superanal genital complex of cerci and paraproct rather uniform; paramere forms
stout, upward arching and slightly narrowing rod. The very tip of the rod armed
with a few number of short and stout spines. P. tagas sp. n. differs from both by
the very long and high gonopods, by the tip of aedeagus and parameres.”
Etymology – “Tagas from “tágas” spacious or wide in Hungarian, refers to
the enlarged gonopods.”
Potamophylax winneguthi (Klapálek, 1902)

Stenophylax winneguthi Klapálek, 1902: 161–162: “Sarajevo 12/11 1899
(Winneguth) 2², Pale (Winneguth) 2²”.
Potamophylax winneguthi (Klapálek, 1902): Schmid (1955: 177): trans-
ferred and listed in genus Potamophylax.
Potamophylax winneguthi (Klapálek, 1902): Oláh & Kovács (2012:
102): “Serbia: Zlatibor Mts, Čigota Mts, spring area of Crni Rzav, N43°37’52.6”,
E19°46’18.0”, 1150 m, 03.11.2011, leg T. Kovács & G. Magos (2², 2³, OPC)
(Oláh 2011). Zlatibor Mts., Čigota Mts., spring area of Crni Rzav, N43°37’52.6”,
E19°46’18.0”, 1150 m, 20.11.2011, leg. T. Kovács & Cs. Oberczán, (15², 7³, OPC)
(Oláh 2011).” “There seems to be no clear feature for separating the females of
P. haidukorum and P. winneguthi wrote Malicky (Malicky & Kumanski 1999).
Differences in both the external and internal genital structure have been found
and compared. See at P. haidukorum.”
Rhadicoleptus Wallengren, 1891

Rhadicoleptus macedonicus Botoşăneanu et Riedel, 1965
Rhadicoleptus alpestris macedonicus Botoşăneanu et Riedel, 1965: 550–
551: “9. Macédoine, Yougoslave, Sar Planina près de Crno Jezero, 9.VIII.1954,
leg. M. Marincovic; 10. Bulgarie, Pirin Planina, 28.VII.1962, leg. B. Russev. Le
holotype ³ de la nouvelle ssp. est choisi parmi les exemplaires du point 9 de notre
liste de stations; il est conservé dans la coll. L. Botoşăneanu; le ² figuré provient
de la station 10, et il est raisonable de l’attribuer à cette race.”

Rhadicoleptus macedonicus Botosaneanu et Riedel, 1965: Oláh et al.

(2015b: 63): Elevated to species rank. “Albania: Erseke county, Grammos Mts,
mountaine pasture on the slope of Mt Varibob, 2.1 km NW of Mt Oukapeci,
2249 m, N40.366220° E20.770510° 19.VII.2006, leg. Z. Barina, T. Pifkó & D.
Pifkó (4 males, 3 females, HNHM). Librazhd county, Jablanica Mts, Quarishte,
brook 6.1km E of the village, 1899 m, N41.24569° E20.51238° 4.VII.2008, leg.
Z. Barina, T. Pifkó & D. Pifkó (3 males, HNHM). Bulqizë district, Çermenikë
Mts, brooks in open forest beneath Mt. Kaptinë, N41°23.199’, E20°17.338’, 1600
m, 21.06.2012, leg. Z. Fehér, T. Kovács, D. Murányi (3 males, 3 females; OPC).
Bulgaria: Rila Mts, Jazovir Belmeken, springbrook, N42°10’22.7”, E23°48’04.4”,
1332m, 23. VI. 2011, P. Juhász, T. Kovács, & L. Urbán, (6 males, 1 female, OPC).
Macedonia: Korab Mts., Malá Korab Vrata, 20.VII.1930 leg. Komarek (3 males,
NMPC; 2 males, OPC).”
Diagnosis – Oláh et al. (2015b: 63): “The subspecies was established only
by female character having supragenital plate very short similar to R. spinifer, but
paramere and gonopod apical tip is different. The lateral profile of parameres is
characterized by low (shallow) curvature, by narrowing gradually apicad without
subbasal constriction and subapical dilatation. This profile is very stable in the
examined populations of Bulgaria, Macedonia and Albania. Ventral profile of the
gonopod apical tip is without highly varying pattern of small lobes, a single blunt
but varying lobe is present usually mesad. Female supragenital plate very short
triangular, but as rounded as the supragenital plate of R. spinifer.”
Rhadicoleptus meridiocarpaticus Botoşăneanu et Riedel, 1965

Rhadicoleptus alpestris meridiocarpaticus Botoşăneanu et Riedel, 1965:
549–550: distingishable only by female. “Nous avons néanmoins considéré néces-
saire de séparer les populations de Bihar et de Banat de celles des Carpates sep-
tentrionales, c’est que le gonopode des ²² de Banat et de Bihar n’est jamais type si
caractéristique pour R. a. sylvanocarpaticus ssp. Locus Typicus: sources réocrene
à Stina de Vale, Mts. de Bihar ou Apuseni, Roumanie, juin 1956. Holotype dans la
coll. L. Botoşăneanu.”
Rhadicoleptus meridiocarpaticus Botoşăneanu et Riedel, 1965: Oláh et al.
(2015b: 63): elevated to species rank. “Romania, Apuseni Mts. Vartop, spring
streams, N46°31.045’ E22°39.821’, 1209m, 29.V.2013, singled leg. J. Oláh, E.
Bajka, Cs. Balogh, & G. Borics (1 female, OPC). Apuseni Mts. Vartop, spring
stream (Flescula), N46° 31’07,23” E22° 39’41,69”1209 m, 14.V.2014, leg. Cs.
Balogh & B.V. Béres (1 male, OPC).”
Diagnosis – Oláh et al. (2015b: 63): “The lateral profile of parameres is
characterized by high (deep) curvature, even, uniform shape without narrowing,

242 J. Oláh
constriction and dilatation. Ventral profile of the gonopod apical tip is slightly
dilated with subtle tiny irregular pattern. Female supragenital plate is long trian-
gular, but more robust than at R. alpestris.”
Rhadicoleptus sylvanocarpaticus Botoşăneanu et Riedel, 1965

Rhadicoleptus alpestris sylvanocarpaticus Botoşăneanu et Riedel,
1965: 547–549: distingishable by both sexes. “3. Region sous-carpatique
d’Ukraine, plusieurs localités: Yanov ( Janow), voisinage Lvov, Ivano-Frankovsk
(Stanislawow), Mlodiatin (Mlodzyatyn) entre Delatin et Kolomyia, (diverses
dates entre 1902–1906), leg. J. Dziędzielewicz. 5. Beskides Orientales, affluents
de la Wetlinka, bassin du San, 22.V.1962, leg. W. Riedel; 6. Maramouresch (di-
verses localités dans les Mts. de Gutin, 20.V. et 23.VI.1964, leg. L. Botoşăneanu.
La ssp. sylvanocarpaticus ssp. n. décrite d’après les spécimens des points 3,5 et
6 de notre liste de stations. Locus Typicus: complexe de sources à Tau la Gutii,
Mts. de Gutin, Maramouresch, Roumanie, 23.VI.1964. Holotype dans la coll. L.
Botoşăneanu.”
Rhadicoleptus sylvanocarpaticus Botoşăneanu et Riedel, 1965: Oláh et
al. (2015b: 65): elevated to species rank. “Romania, Muntii Lapusului (Lápos),
Valeni (Mikolapatak), peat bog, its inflow and outflow brooks in a beech for-
est, N47°42’43.2” E24°01’48.7”, 987m, 23.V.2006 leg. L. Dányi, M. Földvári, J.
Kontschán & D. Murányi (1 male NHMB). Ukraine, Zakarpattia province,
Mizhhirya raion, Kolochava (Alsókalocsa), right sidebrook of Tereblja River by
the village, N48°25.41’ E23°41.56’, 565m, 16.05.2002, leg. D. Murányi (2 males,
4 females, HNHM). Tiachiv raion, Krasna Mts, beech forest edge in the upper
valley of Luzanka River, N48°22.564’ E23°45.081’, 1295 m, 19.05.2002, leg. D.
Murányi (1 female, HNHM).”
Diagnosis – Oláh et al. (2015b: 65): “Paramere is very robust: The lateral
profile of parameres is characterized by very short (very shallow) curvature, by
uniform, even shape without narrowing, and constriction, but with some apical
dilatation. Ventral profile of the gonopod apical tip is with a single blunt narrow-
ing. Female supragenital plate is very long triangular, the dorsal profile of the
vaginal sclerite complex with slight middle constriction.”
BRACHYCENTRIDAE
Micrasema McLachlan, 1876
Micrasema sericeum Klapálek, 1902
Micrasema sericeum Klapálek, 1902: 164–165: “Pazaric, Krupthal 3 ²
(Winneguth), Stolac (Winneguth) 2³.”

HELICOPSYCHIDAE
Helicopsyche Siebold, 1856
Helicopsyche bacescui Orghidan et Botoşăneanu, 1953
Helicopsyche bacescui Orghidan et Botoşăneanu, 1953: 425–431:
“Greaca (Reg. Bucureşti, Olteniţa) în izvoare reocrene ce ies din malul nordic al
bălţii, in apropiere de punctul pescăresc. Comana (Reg. Bucureşti, Olteniţa) în
“izvorul cu nuc”. Comuna Prahova (Reg. Bucureşti). Nenumărate larve şi nimfe,
precum şi 4 ²² şi 1 ³ (toţi adulţii au fost găsiţi la Greaca de către M. Băcescu)”.
BERAEIDAE
Beraea Stephens, 1833
Beraea gurba Oláh, 2014
Beraea gurba Oláh, 2014 in Oláh & Kovács (2014: 127–129): “Holotype.
Albania, Pukë District, rocky stream above Blinisht, 1010m, N42.08290
E19.96340, 13.05.2014 leg. Z. Barina, D. Pifkó & G. Puskás (1², OPC). Allotype.
Same as holotype (1³, OPC). Paratypes. Same as holotype (6², 1³, OPC).”
Diagnosis – Oláh & Kovács (2014: 127–129): “Similar to Beraea zawadil
Malicky described from Greece, but differs by the shorter segment IX, by the
differently shaped gonopods and by the highly bent and four tipped phallic
organ.”
Etymology – “Gurba, from “girbe-gurba”, menadering in Hungarian, refers
to the mesomarginal profile of gonopod ventral branch as visible in ventral view.”
Beraeamyia Mosely, 1930

Beraeamyia schmidi Botoşăneanu, 1960
Beraeamyia schmidi Botoşăneanu, 1960a: 282–286: “Un ² et 1³ de Trnovo
23.VII.1955, désignés comme holotype and allotype (F. Schmid).”
Ernodes Wallengren, 1891

Ernodes skipetarum Malicky, 1986
Ernodes skipetarum Malicky, 1986: 6: “Holotypus ² und 2 ³ Paratypen:
Jugoslawien: Kosovo, östlich des Cakor-Passes, 1500 m, 19.VII.1985. – In meiner
Sammlung”.
SERICOSTOMATIDAE
Notidobia Stephens, 1829
Notidobia bizensis Malicky, 1993

244 J. Oláh
Notidobia bizensis Malicky, 1993: 472–473: “Holotypus ²: Albanien,

Bize bei Shengjergji, 1400–1500 m, 10–15.VII.1961. Paratypus ²: (ehem. Jugos-
lavisch) Makedonien, Galicica-Vojetina, 4.VIII.1930, leg. Komarek, coll. Národni
Muzeum, Prag.”
Notidobia melanoptera Stein, 1863

Notidobia melanoptera Stein, 1863: 415: “Nur diese eine griechische Serico-
stomide in der K. Sammlung wurde von Dr Krüper eingesandt. In Dalmatien
fand ich an einer für diese Thiere ungünstigen Localität während etwas zu später
Jahreszeit 9 Phryganeiden-Arten, darunter das interessante Sericostoma collare
Burm. in Mehrzahl in beiden Geschlechtern.”
Notidobia nekibe Klapálek, 1903

Notidobia nekibe Klapálek, 1903a: 3–4, 1903b: 4–5: “1 ² Janina (Apfel-
beck).”
Notidobia nogradorum Oláh, 2010

Notidobia nogradorum Oláh, 2010: 114–115: “Holotype, male, HNHM.
Albania: Korcë district, Zvirine, Trifti spring N of the village, 835 m, N40°47.644’,
E20°44.128’, 24.V.2007, leg. Z. Barina, Cs. Németh, D. Pifkó.”
Diagnosis – Oláh (2010: 114–115): “This new species belongs to the ho-
mogene group of species: Notidobia melanoptera Stein, 1863 (Greece), N. nekibe
Klapálek, 1903 (Greece), N. sagarrai Navas, 1917 (Sardinia), N. bizensis Malicky
and Sipahiler, 1993 (Albania) and N. salihli Malicky et Sipahiler, 1993 (Turkey).
It is closest to N. bizensis, but differs by having (1) more robust and curve-shaped
groove pattern on the IXth dorsum, not slender and straight; (2) the heavily scle-
rotized pair of paraproctal processes with extremely enlarged dorsal and down-
curving hook-formation, not with small hook; (3) the two spine-shaped proc-
esses on the basomesal surface of the gonopod with separated individual bases,
not with long joint basal plate.”
Etymology – “This species was dedicated to Sára Nógrádi and her husband
Ákos Uherkovich, who have made the Hungarian caddisfl ies one of the best
studied national fauna of the planet.”
ODONTOCERIDAE
Odontocerum Leach, 1815
Odontocerum hellenicum Malicky, 1972

Odontocerum hellenicum Malicky, 1972: 41: “Holotypus ²: Griechenland,

Pertuli (Trikalon), Pindos 1250 m, 1.6.68, leg. Roesler, in coll. Museum Alexander
König, Bonn, Paratypoide: 4 ²² mit den selben Daten, i davon in meiner
Sammlung, die anderen im Museum Bonn.”
LEPTOCERIDAE
Adicella McLachlan, 1877
Adicella altandroconia Botosaneanu et Novák, 1965
Adicella altandroconia Botosaneanu et Novák, 1965: 475–477: “Jusqu’à
present, A. altandroconia est connue des stations suivantes: a. Carpates du Banat
de Roumanie (source dans le Massif de Domogled, près Herkulesbad, leg. L.
Botosaneanu, 1.VII.1956, 1² paratype et 13.VII.1964 3³ paratypes, en alcool);
b. Bulgarie (source dans la vallée du Isker près Svoger, leg. K. Novák, 1.VII.1962.
Holotype ², Allotype ³ et 16 ² paratypes, en alcool).”
Adicella balcanica Botosaneanu et Novák, 1965

Adicella balcanica Botosaneanu et Novák, 1965: 472–476: “Adicella bal-
canica sp. n. est actuellement connue de: A. Bosnie (Ilidze, ruisseau Rakitnica,
leg F. Klapálek, juillet-aôut 1897), 2² 3³ (collection: Musée National, Section
entomologique, Prague). B. Macédoine Yougoslave (Capari, ruisseau moussu,
leg. F. Schmid, 11.VIII.1955). Holotype ² et allotype ³ (Coll. F. Schmid, 1 para-
type ² (Coll. L. Botosaneanu). Les 3 exemplaires conservés à sec, ailes déployées,
les génitalia en préparations microscopiques. C. Bulgarie (Rhodopes Centrales,
petit ruisseau à Smoljan, leg. K. Novák, 7.VII.1962: 3 paratypes ², 1 paratype ³,
tous dans coll. K. Novák, à l’exception d’un ² dans coll. L. Botosaneanu; Stara
Planina, petit ruisseau près de la rivière Zaselja à Bov, leg. K. Novák, 1.VII.1962,
coll. K. Novák, 1 ³). Le materiel est conservé en alcool.”
Setodes Rambur, 1872

Setodes bulgaricus Kumanski, 1976
Setodes viridis bulgaricus Kumanski, 1976 in Kumanski & Malicky
(1976: 120–123): “Holotypus ², Allotypus ³ und Paratypen 3², 1³: Strandscha-
Gebirge, Fluss Weleka bei der Ortschaft Kowatch, 20.VII.1973, leg. A. Sliwow. 1²
und 1³ in coll. Malicky, die anderen in coll. Kumanski. Weitere paratypen in coll.
Kumansky: Fluss Weleka bei der Ortschaft Katschul, 13.VII.1974, 3²; Sakar-
Berge bei Topolowgrad, 11.VII.1974, 1², beide leg. S. Zagortschinow.” Elevated
to species rank by Schmid (1987: 48, 135).

246 J. Oláh
*
Acknowledgements – Important type materials as well as historical and recently collected
specimens have been provided to our revision from several museums with national and region-
al collections and our studies were supported in various ways by Ronald Bellstedt (Museum der
Natur, Gotha); Douglas C. Currie and Mateus Pepinelli (Royal Ontario Museum, Toronto, Cana-
da); Jean-Luc Gattolliat, Anne Freitag and Marion Podolak, (Cantonal Museum of Zoology Laus-
sane, Switzerland); Marcos A. González (Department of Zoology and Physical Anthropology,
Faculty of Biology, University of Santiago de Compostela, Spain); Lars Hendich and Katja Neven,
(Naturwissenschaftliche Sammlungen Bayerns, Zoologisches Staatssammlung, München, Ger-
many); Owen Lonsdale and Serge Laplante (Canadian National Collectionof Insects, Arachnids
and Nematodes, Ottawa, Canada); Martin Lödl and Susanne Randolf (Naturhistorisches Museum
Wien, Austria); Hans Malicky (Lunz-am-See, Austria); Wolfram Mey (Museum für Naturkunde
der Humboldt Universität zu Berlin, Germany); Benjamin Price (The Natural History Museum,
London, England); Peter J. Schwendinger (Museum of Natural History Geneva, Switzerland). We
are thankful to Miklós Bálint, Csaba Balogh, Péter Juhász, Mihály Máté, Dávid Murányi, László
Szél, Ákos Uherkovich for their materials as well as for their valuable helps during our field studies
and in our collecting efforts. We are grateful for their ideas, information and advices giving us dur-
ing the entire five years period of this research.
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Malicky H. 1976: Beschreibung von 22 neuen westpaläarktischen Köcherfl iegen (Trichoptera).
– Arbeitsgemeinschaft Österreichischer Entomologen 27(3–4): 89–104.
Malicky H. 1977: Weitere neue und wenig bekannte mediterrane Köcherfl iegen (Trichoptera). –
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Malicky H. 1984: Vier neue mediterrane Köcherfl iegen (Trichoptera). – Entomologische Zeit-
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Malicky H. 1986: Beschreibung von vier neuen Köcherfl iegen-Arten aus der Türkei und aus
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Malicky H. 1990: Revision der Gattung Melampophylax Schmid, 1955 (Trichoptera, Limnephili-
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FOLIA ENTOMOLOGICA HUNGARICA

Trichoptera endemic in the Carpathian Basin and the adjacent areas

H-4032 Debrecen, Tarján u. 28, Hungary. E-mail: [email protected]

DOI: 10.17112/FoliaEntHung.2017.78.111 Folia ent. hung. 78, 2017

Folia ent. hung. 78, 2017

Thinking of population concept in the new taxonomy requires more elabo-

BDOL = Biologiezentrum des Oberösterreichischen Landesmuseums, Linz, Austria;

Folia ent. hung. 78, 2017

NMW = Naturhistorisches Museum Wien, Austria;

Wormaldia albanica Oláh, 2010: 68–69: “Holotype, male, HNHM. Albania:

Wormaldia balcanica Kumanski, 1979

Folia ent. hung. 78, 2017

12 ²² (all. leg. Malicky); Turkey, Prov. Samsun, 10 km NW Havza (500 m.a.s.l.),

Wormaldia bosniaca Botoşăneanu, 1960

Wormaldia bulgarica Novák, 1971

Wormaldia daga Oláh, 2015

Folia ent. hung. 78, 2017

Wormaldia hellenica Jacquemart, 1962

Wormaldia homora Oláh, 2015

Folia ent. hung. 78, 2017

Wormaldia juliani Kumanski, 1979

Wormaldia kimminsi Botoşăneanu, 1960

Folia ent. hung. 78, 2017

paratypes ² et ³; holot. ² + allot. ³: F. Schmid; 1 ² parat.+ 1³ parat.: Deutsches

Wormaldia subterranea Radovanović, 1932

Folia ent. hung. 78, 2017

Tinodes Curtis, 1834

Tinodes polifurculatus Botoşăneanu, 1956

Tinodes popovi Kumanski, 1975

Folia ent. hung. 78, 2017

Tinodes raina Botoşăneanu, 1960

Tinodes unidentatus Klapálek, 1894

Tinodes urdhva Oláh, 2010

Plectrocnemia minima Klapálek, 1899

Plectrocnemia mojkovacensis Malicky, 1982

Plectrocnemia smiljae Marinković-Gospodnetić, 1966

Polycentropus Curtis, 1835

Polycentropus ierapetra septentrionalis Kumanski, 1986

Polycentropus ierapetra slovenica Malicky, 1998

Polycentropus schmidi Novák et Botoşăneanu, 1965

Folia ent. hung. 78, 2017

Hydropsyche botosaneanui Marinković-Gospodnetić, 1966a: 112.

Hydropsyche dinarica Marinković-Gospodnetić, 1979

Hydropsyche emarginata Navas, 1923

Hydropsyche mostarensis Klapálek, 1898

Hydropsyche peristerica Botosaneanu et Marinković-Gospodnetić, 1966

Hydropsyche sarnas Oláh, 2015

Folia ent. hung. 78, 2017

river Drino, 170m, N40.17854° E20.07981°, 10.05.2014, leg. Z. Barina, D. Pifkó

Hydropsyche sinuata Botosaneanu et Marinković-Gospodnetić, 1966

Hydropsyche smiljae Marinković-Gospodnetić, 1979

Hydropsyche tabacarui Botoşăneanu, 1960

Folia ent. hung. 78, 2017

Rhyacophila akutila Oláh, 2010: 82–83: “Holotype male, HNHM. Bulgaria:

Rhyacophila balcanica Radovanović, 1953

Rhyacophila biegelmeieri Malicky, 1984

Folia ent. hung. 78, 2017

Rhyacophila bosnica Schmid, 1970