Opusc. Zool. Budapes , 2025, 56(3): 13–96
u n: lsid:zoobank.o g:pub:9AA9E894-1687-4DEB-B120-26C07CD98213 published: 30 Sep embe 2025
HU ISSN 2063-1588 (online) h ps://doi.o g/10.18348/opzool.2025.3.13
On he T ichop e a o he Caucasus, wi h emphasis on he
Plec ocnemia la issima and P. zeka ia species complexes,
and he gene a Ce asma and No idobia
J. OLÁH1, G. VINÇON2, H. IBRAHIMI3, A. MEMISHISHI4 & K. DZULIASHVILI4
1János Oláh, Residence pos al add ess: Ta ján u. 28, H-4032 Deb ecen, Hunga y.
E-mail: [email p o ec ed] h ps://o cid.o g/0000-0002-6137-0553
2Gilles Vinçon, Residence pos al add ess: 55 Bd Joseph Vallie , F-38100 G enoble, F ance.
E-mail: [email p o ec ed] h ps://o cid.o g/0000-0003-4702-5136
3Halil Ib ahimi, Uni e si y o P ish ina, Facul y o Ma hema ics and Na u al Sciences, Depa men o Biology,
Mo he Te esa p.n., 10000 P ish ina, Koso o. E-mail: halil.ib ahimi@uni-p .edu
h ps://o cid.o g/0000-0002-4301-4387
4Aleksi Memishishi & K is ina Dzuliash ili, Ins i u e o Zoology, Ilia S a e Uni e si y, Cholokash ili A e. 3/5
Tbilisi, 0162, Geo gia. E-mail: al[email p o ec ed] h ps://o cid.o g/0000-0002-5862-5105
E-mail: [email p o ec ed] h ps://o cid.o g/0009-0000-2994-6586
Abs ac . In ou hi d pape on he Caucasian T ichop e a we ha e desc ibed 56 new caddis ly species mos ly om Geo gia:
Wo maldia elko a Oláh & Vinçon sp. no ., W. gode dza Oláh & Vinçon sp. no ., W. halchala, Oláh & Vinçon sp. no ., W.
kin isha Oláh & Vinçon sp. no ., W. omala Oláh & Vinçon sp. no ., Tinodes dobe ad Oláh & Memishishi sp. no ., T.
adzen Oláh & Memishishi sp. no ., Plec ocnemia a asba anica Ib ahimi & Oláh sp. no ., P. ka lia Oláh & Vinçon sp.
no ., P. kope dagha Oláh & Ib ahimi sp. no ., P. mashukana Oláh sp. no ., P. ming elica Oláh & Vinçon sp. no ., P.
s ane ica Oláh & Vinçon sp. no ., P. ucnobi Oláh sp. no ., P. gomis a Oláh & Vinçon sp. no ., P. kin isha Oláh &
Memishishi sp. no ., Cheuma opsyche bushe ica Oláh sp. no ., C. e zincana Oláh sp. no ., C. io ica Oláh & Memishishi sp.
no ., Hyd opsyche dashbasha Oláh & Vinçon sp. no ., H. io ica Oláh & Memishishi sp. no ., H. gaghma Oláh &
Memishishi sp. no ., H. akhalchala Oláh & Vinçon sp. no ., H. gode dza Oláh & Vinçon sp. no ., H. mes ia Oláh & Vinçon
sp. no ., H. sabna a Oláh & Memishishi sp. no ., Agape us akhalchal Oláh & Vinçon, sp. no ., A. bakhma Oláh & Vinçon,
sp. no ., A. lashijal Memishishi & Oláh sp. no ., Mic asema kaskadian Oláh & Memishishi sp. no ., Th emma kin icum
Oláh & Memishishi sp. no ., T. adja icum Oláh & Memishishi sp. no ., Ma ynomyia bakhma a Oláh & Vinçon sp. no ., M.
kin isha Oláh & Memishishi sp. no ., M. omala Oláh & Vinçon sp. no ., D usus akhalchala Oláh & Vinçon, sp. no ., D.
gudau Oláh & Vinçon, sp. no ., D. kheled Oláh & Vinçon, sp. no ., D. zeka Oláh & Vinçon, sp. no ., Sakala gou ica Oláh
& Vinçon, sp. no ., S. ja ake ica Oláh & Vinçon, sp. no ., Rizeiella bakhma a Oláh & Vinçon sp. no ., S enophylax bo os
Oláh & Memishishi sp. no ., S. chokhiensis Oláh & Vinçon sp. no ., E nodes gou iensis Oláh & Vinçon sp. no ., Ce asma
a ina Oláh sp. no ., C. bakhma a Oláh & Vinçon sp. no ., C. gode da Oláh & Vinçon sp. no ., C. goma Oláh & Vinçon
sp. no ., C. kin isha Memishishi & Oláh sp. no ., C. za zma Oláh & Vinçon sp. no ., No idobia kin isha Oláh &
Memishishi sp. no ., N. k a ia i Oláh & Memishishi sp. no ., N. ma ielucae Oláh & Vinçon sp. no ., N. ekona Oláh &
Memishishi sp. no ., Se odes io icus Oláh & Memishishi sp. no . The ollowing 13 species a e eco ded as new o he
Geo gian Fauna: Wo maldia daga Oláh, 2014, Lype phaeopa S ephens, 1836, Plec ocnemia izeiensis Sipahile , 1987,
Cheuma opsyche la ella a Mey, 2004, Hyd opsyche scilig a Malicky, 1977, Agape us caucasicus, Ma yno , 1913,
Hyd op ila a ma hai Schmid, 1959, H. au ica Ma yno , 1934, S ac obia wimme i Malicky, 1988, Limnephilus au icula
Cu is, 1834, S enophylax me idio ien alis Malicky, 1980, S. solo a ewi (Ma yno , 1913), T iaenodes kaw aiskii Ma yno ,
1909.
Keywo ds. Caucasus, Geo gia, T ichop e a, species complexes, new species.
Oláh e al.: On he T ichop e a o he Caucasus III
14
INTRODUCTION
his is ou hi d pape on he T ichop e a o
he Caucasus (Oláh e al. 2020, Oláh & Vin-
çon 2024). Ou cu en sampling and s udy
ac i i ies a e mos ly es ic ed o Geo gia and a
small pa o he Lesse Caucasus (also known as
he Caucasus Mino o An icaucasus) and he
G ea e Caucasus moun ain anges. The wo
anges un pa allel o a ound 600 kilome es
h ough six coun ies: Russia, Tu key, Geo gia,
A menia, Aze baijan and I an. Only a e y small
numbe o limi ed habi a s we e sampled. To
compensa e he limi s o sample size we applied
he p inciple and p ac ices o “pu oge he ” p o-
cedu e in o de o ha e a li le mo e han no hing,
a eally e y small s udy ma e ial a ou disposal
o axonomic esea ch. Ne e heless, e en om
his small p i a e ial we ha e desc ibed many
new caddis ly species. This clea ly e lec s he
almos p is ine s a e o he en i e Caucasus, which
has been sampled spo adically and s udied by
only a ew expe s, such as he Russian And ei
Vasilie ich Ma yno , he Ge man Wol am Mey,
and he Tu kish Füsun Sipahile . The disco e y
and desc ip ion o 173 new caddis ly species om
such small, agmen ed samples highligh s he in-
adequacy o suppo o he ins i u ional scien i ic
ne wo k o axonomy – he na u al his o y muse-
ums, uni e si ies, and aqua ic and ecological
esea ch ins i u es – and demons a es how his
hinde s hei abili y o ca y ou i al biodi e si y
esea ch. Fu he mo e, i clea ly shows how li le
we know abou he s uc u e o li ing aqua ic
communi ies in such a as , moun ainous egion
wi h i s highly di e se and isola ed habi a s. Un-
o una ely his a he sad scena io o he con em-
po a y ins i u ional axonomic esea ch is no con-
ined o T ichop e a and o he Caucasus, bu i is
also e iden in o he g oups and h oughou
Eu ope (Páll-Ge gely e al. 2024).
MATERIAL AND METHODS
Ou p esen s udy is based on caddis ly spe-
cimens sampled by he second au ho Gilles Vin-
çon in Geo gia du ing his ea ly sp ing (20.IV–
4.V) and midsumme (2–18.VIII) collec ing ips
in he yea o 2024 co e ing he basic aqua ic
habi a s o all he basic egions mapped al eady
be o e (Oláh & Vinçon 2024). Halil Ib ahimi has
con ibu ed o he knowledge o Plec ocnemia
la issima new species complex by his specimens
collec ed in I an, Eas Aze baijan, and he Elbu z
Moun ains. Aleksi Memishishi and K is ina Dzui-
lash ili ealised some collec ing ips in unex-
plo ed habi a s o Geo gia, as well as iden i ied a
pa o a ma e ial om a big Malaise ap esea ch
p ojec o Illia S a e Uni e si y wi h aps in-
s alled in 18 loca ions up o he sp ing a ea on he
Kin ishi Ri e in he Adja a egion. Some spe-
cimens we e collec ed and p esen ed o Aleksi
Memishishi by he Finnish lepidop e ologis , Ja i
Junnilainen as byp oduc o his Lepidop e a col-
lec ions using ligh aps.
Pho og aphs (Appendix 1) illus a ing he
ecology and dis ibu ion o each new species a e
included a he end o he ex .
The abb e ia ions o he deposi o y ins i u ions
a e as ollows:
Depa men o Biology, Facul y o Ma hema ics and
Na u al Sciences, Uni e si y o P ish ina „Hasan
P ish ina”, P is inë, Koso o (UPK).
Ins i u e o Zoology, Ilia S a e Uni e si y, Tbilisi,
Geo gia (IZISUT)
Oláh P i a e Collec ion, Deb ecen, Hunga y; unde
na ional p o ec ion by he Hunga ian Na u al His-
o y Museum, Budapes (OPC).
TAXONOMY
Philopo amidae S ephens, 1829
Wo maldia McLachlan, 1865
Wo maldia daga Oláh, 2014
(Pho os 7a–7b)
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 11, 1637m, 41.7476°N, 42.095°E, 29.VI.–13.
VII.2018, leg. GGBC membe s (5 males, OPC).
T
Oláh e al.: On he T ichop e a o he Caucasus III
15
Rema ks. This species is desc ibed and known
only om Bulga ia (Bosna M s., S andzha M s.).
A e i s desc ip ion i is i s eco ded he e in he
Kin ishi s eam o Adja a, Geo gia a om he
locus ypicus. The e is a sibling specie, Wo mal-
dia di a Sipahile , 2020 desc ibed om No heas
Tu key, A in, Bo cka wi h almos iden ical
endo hecal spine pa e n, li le di e ged head o
segmen X, and wi h mo e di e ged pe iphallic
o gan o unca ed ce ci. Su p isingly, he i e
Malaise- apped specimens in Kin ishi s eam
ha ing e en he ce ci ounded, no unca ed;
du ing a ca e ul compa ison wi h he pa a ype
om S andzha M s we we e unable o ind any
di e gences. We do no know which kind o
mechanisms e ained i s iden i y he e in Kin ishi
alley and al e ed sligh ly in A in. Also we do
no know how con inuous is i s dis ibu ion in
be ween o S andzha and Kin ishi moun ains
and how a sp ead u he eas wa d.
Wo maldia da idi Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, High S ane i,
abo e Nak a, b ook and sp ing, 1520m, 43.0781°
N, 42.3695°E, 16.VIII.2024, leg. Gilles Vinçon (1
male, OPC).
Rema k. This species was known only by he
holo ype.
Wo maldia elko a Oláh & Vinçon, sp. no .
(Figu es 1–3, Pho os 1–2)
Wo maldia kime a Oláh & Vinçon, 2020:10–11 (pa .).
“Pa a ype: Geo gia, Kakhe ia egion, abo e Le-
chu i, in di ec ion o Omalo, big o en abo e he
b idge, ibu a y o S o i A ag i Ri e , 880m,
42°12’19”N, 45°27’45”E, 3.X.2019, leg. G. Vinçon
(1 male, OPC).” Misiden i ica ion.
Wo maldia holaga Oláh & Manko, 2020 in Oláh e al.
2020: Oláh & Vinçon 2024:3–4. Geo gia, Sam-
skhe-Ja akhe i, K sia-Taba sku i Managed Re-
se e, sp ing and b ook, 2380m, 41.6555°N,
43.49°E, 23.IX.2022, leg. Gilles Vinçon (2 males, 6
emales; OPC). Geo gia, Sam skhe-Ja akhe i, SE
Tsagh e i, nice b ook, 1210m, 41.791°N, 43.489°
E, 23.IX.2022, leg. Gilles Vinçon (6 males, 2
emales; OPC). Geo gia, Kakhe ie, abo e Lapan-
ku i, Lopo a alley, o en and nice la e al sp ing
and b ookle , 1580m - 1620m, 42.15°N, 45.6668°E,
21.IX.2022, leg. Gilles Vinçon (2 males, OPC).
Misiden i ica ion.
Wo maldia obola Oláh, 2020:13–14: Oláh e al.
2024:8. “Geo gia, Kakhe ia, Lagodekhi Na ional
Pa k, abo e Khizaba a, Ninoskhe i alley, sp ing
and b ook, 630m, 41.8778N, 46.2425E, 13.V.2023,
leg. Gilles Vinçon (1 male, OPC). Geo gia, Ka-
khe ia, abo e Khizaba a, Ninoskhe i alley,
b ook, 930m, 41.8826N, 46.2561E, 13.V.2023, leg.
Gilles Vinçon (5 males, OPC).” Misiden i ica ion.
Ma e ial examined. Holo ype: Geo gia,
K emo Ka li, Dashbashi cascades, 1410m,
41.595°N, 44.124°E, 24.IV.2024, leg. Gilles
Vinçon (1 male, OPC). Pa a ypes: same as
holo ype (2 males, OPC). Geo gia, K emo Ka li,
Kh ami Ri e a Dashbashi cascades, 1400m,
41.5945°N, 44.1238°E, 24.IV.2024 leg. Gilles
Vinçon (2 males, OPC). Geo gia, Kakhe ia
egion, abo e Lechu i, in di ec ion o Omalo, big
o en abo e he b idge, ibu a y o S o i A ag i
Ri e , 880m, 42°12’19”N, 45°27’45”E, 3.X.2019,
leg. G. Vinçon (1 male OPC). Geo gia, Sam skhe-
Ja akhe i, K sia-Taba sku i Managed Rese e,
sp ing and b ook, 2380m, 41.6555°N, 43.49°E,
23.IX.2022, leg. Gilles Vinçon (2 males, 6 e-
males; OPC). Geo gia, Sam skhe-Ja akhe i, SE
Tsagh e i, nice b ook, 1210m, 41.791°N, 43.489°
E, 23.IX.2022, leg. Gilles Vinçon (6 males, 2
emales; OPC). Geo gia, Kakhe ie, abo e La-
panku i, Lopo a alley, o en and nice la e al
sp ing and b ookle , 1580m – 1620m, 42.15°N,
45.6668°E, 21.IX.2022, leg. Gilles Vinçon (2
males, OPC). Geo gia, Kakhe ia, Lagodekhi Na-
ional Pa k, abo e Khizaba a, Ninoskhe i alley,
sp ing and b ook, 630m, 41.8778°N, 46.2425°E,
13.V.2023, leg. Gilles Vinçon (1 male, OPC).
Geo gia, Kakhe ia, abo e Khizaba a, Ninoskhe i
alley, b ook, 930m, 41.8826°N, 46.2561°E, 13.
V.2023, leg. Gilles Vinçon (5 males, OPC).
Geo gia, Sam skhe-Ja akhe i, abo e Pa a a Pa-
maji, sp ing, 1950m, 41.560°N, 42.927°E, 7.VIII.
2024, leg. Gilles Vinçon (3 males, OPC).
Diagnosis. I s ape ing ha pago ela es i o he
Wo maldia ianguli e a species g oup, and ha -
ing h ee endo hecal s ou spines i belongs he W.
Oláh e al.: On he T ichop e a o he Caucasus III
16
Figu es 1–3. Wo maldia elko a sp. no . Holo ype male: 1 = geni alia in le la e al iew; 2 = e gi e VIII
in do sal iew;3 = phallic o gan in le la e al iew.
khou mai species complex. I s ela i ely sho
head o segmen X esembles i o W. holaga Oláh
& Manko, 2020; W. khou mai Schmid, 1959; W.
obola Oláh, 2020 and W. sima Oláh & Ch ojka,
2019. This was he eason why we ha e egula ly
misiden i ied p e iously his abundan species
(Oláh e al. 2020, Oláh & Vinçon 2024). A
de ailed and compa a i e e-examina ion e ealed
he s abili y o he semi-ci cula la e al p o ile o
he head o segmen X compa ed o ull ci cula
(holaga), an e ad u ning (khou mai), elonga ed
hal -ci cula (obola) and ounded iangula (si-
ma) head. These sub le di e gences in his sen-
si i e i illa o y o gan seem s able, bu could be
exposed o a ious de o ma ions du ing copula-
ion and p epa a ion.
Desc ip ion. Male (in alcohol). La ge cas a-
nean b own animal. Scle i es medium b own,
se al wa s bo h on head and ho ax and legs
b own. Maxilla y palp o mula is I-II-IV-III-V.
Fo ewing leng h 7 mm. Spu o mula is 244.
Male geni alia. Te gi e VIII wi h e y deep
and wide mesal excision on he apical ma gin
wi h p onounced la e al lobes. La e al p o ile o
segmen X cha ac e ized by semi-ci cula head,
p onounced subapical conca i y and by s aigh
basal egion. Ce ci wi h downwa d oblique un-
ca ed ounded apex. Ha pago s aigh ape ing and
sligh ly sho e han coxopodi e. Phallic o gan
wi h one la ge s aigh s ou and wo almos
equally sized slende cu ing s ou spines accom-
panied by a small clus e o small and sho mi o
spines (spines a anged in wo lines) as well as by
some longe and slende spines.
E ymology. Species name e lec s he cha ac e
s a e o he specia ion ai , he la e al p o ile o
he head o segmen X. I is being semi-ci cula ;
he e o e “ elko a” epi he was coined om “ él-
kö ” hal ci cula in Hunga ian.
Rema ks. W. obola Oláh, 2020 is no longe e-
co ded o Geo gia since he specimens epo ed
in Oláh e al. 2024 a e assigned he e o W. elko a
Oláh & Vinçon, sp. no .
Wo maldia gode dza Oláh & Vinçon, sp. no .
(Figu es 4–6, Pho os 3–4)
Ma e ial examined. Holo ype: Geo gia, Ad-
ja a, Gode dzi Pass, below G een Lake, la ge
b ook below he swamp, 2050m, 41.682°N,
42.500°E, 8.VIII.2024, leg. Gilles Vinçon (1
male, OPC).
Diagnosis. Ha ing ape ing ha pago belongs o
he Wo maldia ianguli e a species g oup.
Oláh e al.: On he T ichop e a o he Caucasus III
17
Figu es 4–6. Wo maldia gode dza sp. no . Holo ype male: 4 = geni alia in le la e al iew; 5 = e gi e VIII
in do sal iew; 6 = phallic o gan in le la e al iew.
Ha ing single s ou spine belongs o Wo maldia
bulga ica species complex. In his apidly in-
c easing complex, he e is an inc easing species
uni wi h sho head o segmen X: Wo maldia
bulga ica No ák, 1971; W. daga Oláh, 2014; W.
di a Sipahile , 2020; W. el es a Oláh, 2020; W.
e zincanica Sipahile , 2012; W. ha ma Oláh &
Vinçon, 2020; W. kin isha Oláh & Vinçon sp.
no ., W. kumanskii Oláh & Ch ojka, 2019.
I is no easy o dis inguish he nin h species
W. gode dza Oláh & Vinçon sp. no . o he ain-
guli e a species g oup om i s ela i es. The
single male was collec ed om he dis ibu ional
a ea o W. ha ma and i is mos close o i , bu
di e s by sho e and b oade and dis inc ly un-
ca ed ce ci, by sho e and sligh ly upwa d u ning
head and by he nine, no h ee long slende spines
in endo heca o phallic o gan.
Desc ip ion. Male (in alcohol). Small cas a-
nean b own animal. Scle i es medium b own,
se al wa s bo h on head and ho ax and legs
b own. Maxilla y palp o mula is I-II-IV-III-V.
Fo ewing leng h 7 mm. Spu o mula is 244.
Male geni alia. Te gi e VIII wi h deep and
wide mesal excision on he apical ma gin o med
by p onounced la e al lobes. Segmen X cha ac-
e ized by a do sal g adually b oadening, sligh ly
upwa d di ec ed ounded head, dis inc ly, exac ly
and p ecisely isible in la e al iew; he head
obliquely and smoo hly sloping an e ad wi hou
any poin o any angle. Ce ci a he sho and
b oad wi h unca e apex. Ha pago ape ing long,
wi h sligh ly downwa d u ning apical egion,
li le sho e han coxopodi . Phallic o gan wi h a
single small s ou spine accompanied by a small
clus e o small and sho double clus e ed spines
as well as by nine longe and slende spines.
E ymology. Species name was coined om he
ype locali y, a noun in apposi ion.
Wo maldia halchala Oláh & Vinçon, sp. no .
(Figu es 7–10, Pho os 5–6)
Ma e ial examined. Holo ype: Geo gia, Ra-
cha-Lechkhumi and Lowe -S ane i, below Akhal-
chala, nice sp ings, 1870m, 42.6768°N, 42.6482°
E, 13.VIII.2024, leg. Gilles Vinçon (1 male,
OPC). Pa a ype: Geo gia, Racha-Lechkhumi and
Lowe -S ane i, Akhalchala, nice b ook and
sp ing, 2120m, 42.6928°N, 42.6447°E, 13.VIII.
2024, leg. Gilles Vinçon (1 male, OPC).
Diagnosis. Ha ing ape ing ha pago i belongs
o he Wo maldia ianguli e a species g oup, and
ha ing wo s ou spines i is a membe o he W.
khou mai species complex. I is a ecen ly di-
e ged sibling species o Wo maldia kulbaka
Oláh e al.: On he T ichop e a o he Caucasus III
18
Figu es 7–10. Wo maldia halchala sp. no . Holo ype male: 7 = geni alia in le la e al iew; 8 = geni alia in do sal iew;
9 = e gi e VIII in do sal iew; 10 = phallic o gan in le la e al iew.
Oláh & Vinçon, 2024 desc ibed om abo e
Kulbaki, downwa ds Akhalchala, bu di e s by
he deep and b oade excision on e gi e VIII, by
he do sally middle angled and la e ally sligh ly
S-shaped ce ci; by he sho e head o segmen X
wi h sligh ly conca e, no s aigh head do sum as
well as by he endo hecal spine pa e n a med
wi h an addi ional clus e o small spines.
Desc ip ion. Male (in alcohol). Small cas a-
nean b own animal. Scle i es medium b own,
se al wa s bo h on head and ho ax and legs
b own. Maxilla y palp o mula is I-II-IV-III-V.
Fo ewing leng h 6 mm. Spu o mula is 244.
Male geni alia. Te gi e VIII wi h e y deep
and na ow mesal excision on he apical ma gin
wi h p onounced iangula la e al lobes. Segmen
X cha ac e ized by elonga ed ounded head wi h
conca e do sum, shallow subapical conca i y and
by s aigh basal egion. Ce ci S-shaped in la e al
and middle angled in do sal iews. Ha pago
s aigh ape ing, as long as coxopodi e. Phallic
o gan wi h wo almos equally la ge s ou spines
accompanied by wo clus e s o small and sho
spines as well as by some longe and slende
spines.
E ymology. Species name was coined om he
ype locali y, a noun in apposi ion.
Wo maldia ha ma Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, High S ane ie,
abo e Mes ia, b ook and sp ings abo e he ish
a m in cons uc ion, 1550m, 43.0477°N,
42.8091°E, 3.V.2024, leg. Gilles Vinçon (1 male,
OPC).
Wo maldia holaga Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, High S ane ie,
abo e Mes ia, b ook and sp ings abo e he ish
a m, 1550m, 43.0477°N, 42.8091°E, 3.V.2024,
leg. Gilles Vinçon (4 males, OPC). Geo gia,
M skhe a-M iane i, Ghelis ake, 42.5289°N,
44.921°E, 9.X.2024, leg. A. Memishishi (1 male,
OPC). Geo gia, M skhe a-M iane i, Be isch dili,
42.41133°N, 44.92827°E, 10.X. 2024, leg. A.
Memishishi (3 males, OPC).
Wo maldia ke a Oláh, 2019
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe S ane i, So i, sp ing, 710m,
Oláh e al.: On he T ichop e a o he Caucasus III
19
Figu es 11–14. Wo maldia kin isha sp. no . Holo ype male: 11 = geni alia in le la e al iew; 12 = geni alia in do sal iew;
13 = e gi e VIII in do sal iew; 14 = phallic o gan in le la e al iew.
Figu es 15–16. Wo maldia kin isha sp. no . Pa a ype male: 15 = geni alia in le la e al iew;
16 = phallic o gan in le la e al iew.
42.58117°N, 43.28706°E, 26.IV.2024, leg. A.
Memishishi (1 male, 3 emales; IZISUT).
Wo maldia kime a Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, K emo Ka li,
Dashbashi cascades, 1410m, 41.595°N, 44.124°E,
24.IV.2024, leg. Gilles Vinçon (1 male, OPC).
Rema k. This species was only known om
he holo ype collec ed in he High Caucasus. He e
we epo i o he i s ime om he Lesse Cau-
casus.
Wo maldia khizaba a Oláh & Vinçon, 2024
Ma e ial examined. Geo gia, Kakhe ia, abo e
Khizaba a, b ook, 1000m, 41.8829°N, 46.2585°
E, 21.IV.2024, leg. Gilles Vinçon (4 males, OPC).
Wo maldia kin isha Oláh & Vinçon, sp. no .
(Figu es 11–16, Pho os 7–9)
Ma e ial examined. Holo ype: Geo gia, Ad-
ja a, Kin ishi, o en and b ookle s, 1040m,
41.733°N, 42.082°E, 29.IV.2024, leg. Gilles Vin
Oláh e al.: On he T ichop e a o he Caucasus III
20
çon (1 male, OPC). Pa a ypes: Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 11, 1637m, 41.7476°N, 42.0951°E,
29.VI.–13.VII.2018, leg. GGBC membe s (6
males, 6 emales, IZISUT; 6 males, 2 emales;
OPC).
Diagnosis. This new species ha ing ape ing
ha pago and a single s ou spine in he endo heca
belongs o he Wo maldia bulga ica species
complex o he Wo maldia ianguli e a species
g oup. Acco ding o he Wo maldia specia ion
ai , ha is he la e al p o ile o he head o
segmen X his new species is simila o Wo -
maldia a janulia Oláh & Vinçon, 2024, bu
di e s by he unique cla a e head o ce ci, he
e y deep and wide subapical do sal excision in
he la e al p o ile o segmen X, he poin ed
an e ad u ning an e io co ne o head o segmen
X as well as by he single, no double s ou
endo hecal spine. We ha e eco ded some sligh
a ia ion in he la e al p o ile o segmen X head
o he examined wel e males.
Desc ip ion. Male (in alcohol). Unusually
la ge cas anean b own animal. Scle i es medium
b own, se al wa s bo h on head and ho ax and
legs b own. Maxilla y palp o mula is I-II-IV-III-
V. Fo ewing leng h 10 mm. Spu o mula is 244.
Male geni alia. Te gi e VIII wi h e y deep
and wide mesal excision on he apical ma gin
wi h p onounced quad a ic la e al lobes. La e al
p o ile o segmen X cha ac e ized by sho an-
e ad poin ed head, e y deep subapical conca i y
and by s aigh basal egion. Ce ci wi h unique
cla a e apex in do sal iew. Ha pago s aigh
ape ing wi h downwa d u ning apex, sligh ly
longe han coxopodi e. Phallic o gan wi h a
single la ge s aigh s ou accompanied by a small
clus e o small and sho mi o spines as well as
by some longe and slende spines.
E ymology. Species name was coined om he
ype locali y, a noun in apposi ion.
Wo maldia kumanskii Oláh & Ch ojka, 2019
Ma e ial examined. Geo gia, Sam skhe-Ja a-
khe i, Zeda Va dzia, nice b ook and sp ing,
1410m, 41.383°N, 43.258°E, 25.IV.2024, leg.
Gilles Vinçon (3 males, OPC). Geo gia, Adja a,
Kin ishi, nice o en wi h mosses, 700m,
41.7385°N, 42.01°E, 29.IV.2024, leg. Gilles Vin-
çon (1 male, 1 emale, in copula; OPC). Geo gia,
Adja a, M i ala Na ional Pa k, nice o en , 330m,
41.6762°N, 41.8707°E, 30.IV.2024, leg. Gilles
Vinçon (5 males, OPC). Geo gia, Adja a, K a ia,
sp ing, 41.57335°N, 42.41838°E, 1.V.2024, leg.
A. Memishishi (2 males, 1 emale; IZISUT).
Geo gia, Adja a, Va janauli, sp ing, 41.788°N,
41.9629°E, 29.IV.2024, leg. A. Memishishi (2
males, IZISUT). Geo gia, Adja a, Dunga, wa e -
all, 41.47544°N, 41.85622°E, 29.IV.2024, leg. A.
Memishishi (2 males, 1 emale; IZISUT). Geo -
gia, Adja a, Mu ala, sp ing, 41.6831°N, 41.8442°
E, 29.IV.2024, leg. A. Memishishi (1 male,
IZISUT). Geo gia, Gou ia, oad o Bakhma o,
nice b ook and sp ing, 1780m, 41.8818°N,
42.3567°E, 10.VIII.2024, leg. Gilles Vinçon (3
males, OPC). Geo gia, Adja a, Gode dzi Pass,
a e Beshumi Bo anic Ga den, b ook and sp ing,
1970m, 41.6216°N, 42.5383°E, 7.VIII.2024, leg.
Gilles Vinçon (3 males, OPC). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 11, 1637m, 41.74769°N, 42.0951°E,
29.VI.–13.VII.2018, leg. GGBC membe s e al. (3
males, 6 emales; OPC).
Wo maldia omala Oláh & Vinçon, sp. no .
(Figu es 17–19, Pho os 9–10)
Ma e ial examined. Holo ype: Geo gia, Ka-
khe i, oad o Omalo, nice b ook unde cascade,
1500m, 42.2373°N, 45.4921°E, 5.VIII.2024, leg.
Gilles Vinçon (1 male, OPC).
Diagnosis. I s ape ing ha pago ela es i o he
Wo maldia ianguli e a species g oup, and ha -
ing h ee endo hecal s ou spines i belongs he W.
khou mai species complex. Mos close o Wo -
maldia elko a sp. no . bu di e s by he longe
and oundedhead, sho e and deepe subapical
do sal excision on segmen X as well as by he
shallow, wide no deep and na ow do soapical
excision o segmen VIII.
Desc ip ion. Male (in alcohol). La ge cas-
anean b own animal. Scle i es medium b own, se-
Oláh e al.: On he T ichop e a o he Caucasus III
21
Figu es 17–19. Wo maldia omala sp. no . Holo ype male: 17 = geni alia in le la e al iew; 18 = e gi e VIII
in do sal iew; 19 = phallic o gan in le la e al iew.
se al wa s bo h on head and ho ax and legs
b own. Maxilla y palp o mula is I-II-IV-III-V.
Fo ewing leng h 7 mm. Spu o mula is 244.
Male geni alia. Te gi e VIII wi h e y deep
and wide mesal excision on he apical ma gin
wi h p onounced la e al lobes. La e al p o ile o
segmen X cha ac e ized by semi-ci cula head,
p onounced subapical conca i y and by s aigh
basal egion. Ce ci wi h downwa d oblique un-
ca ed ounded apex. Ha pago s aigh ape ing and
sligh ly sho e han coxopodi e. Phallic o gan
wi h one la ge s aigh s ou and wo almos
equally sized slende cu ing s ou spines ac-
companied by a small clus e o small and sho
mi o spines (spines a anged in wo lines) as
well as by some longe and slende spines.
E ymology. Species name was coined om he
ype locali y, a noun in apposi ion.
Wo maldia sima Oláh & Ch ojka, 2019
Wo maldia holaga Oláh & Manko, 2020: Oláh &
Vinçon 2024:3. Geo gia, Ming elia - High S ane-
ia, Ma khi Valley, b ookle , 1980m, 43.0796°N,
42.231°E, and sp ing, 2030m, 43.081°N, 42.231°E,
29.IX.2022, leg. Gilles Vinçon (3 males, OPC).
Misiden i ica ion.
Ma e ial examined. Geo gia, Ming elia-High
S ane ia, Ma khi Valley, b ookle , 1980m,
43.079°N, 42.231°E, and sp ing, 2030m, 43.081°
N, 42.231°E, 29.IX.2022, leg. Gilles Vinçon (3
males, OPC). Geo gia, High S ane ie, abo e
Mes ia, b ook and sp ings abo e he ish a m in
cons uc ion, 1550m, 43.0477°N, 42.8091°E, 3.V.
2024, leg. Gilles Vinçon (11 males, OPC). Geo -
gia, M skhe a-M iane i, bellow Tsinamkha i, nice
sp ing and b ook, 1200m, 42.3267°N, 44.6469°E,
22.IV.2024, leg. Gilles Vinçon (4 males, OPC).
Geo gia, Racha-Lechkhumi and Lowe -S ane i,
Akhalchala, o en abo e cascade, 2400m,
42.7°N, 42.6455°E, 13.VIII.2024, leg. Gilles Vin-
çon (1 male, OPC).
Rema ks. The endo hecal spine pa e n o W.
sima Oláh & Ch ojka, 2019 and W. holaga Oláh
& Manko, 2020 is e y simila , almos iden ical.
Howe e , he la e al p o ile o he head o
segmen X is dis inc ly di e gen i i is compa ed
ca e ully.
Psychomyiidae Walke , 1852
Lype phaeopa S ephens, 1836
Ma e ial examined. Geo gia, Adja a, Mi e i,
wa e all, 41.53056°N, 41.71035°E, 30.IV.2024,
leg. A. Memishishi (1 male, IZISUT).
Rema k. This is he i s epo om Geo gia.
Oláh e al.: On he T ichop e a o he Caucasus III
28
Figu es 34–37. Plec ocnemia kope dagha sp. no . Holo ype male: 34 = geni alia in le la e al iew; 35 = geni alia
in do sal iew; 36 = gonopods in en al iew; 37 = phallic o gan in le la e al iew.
Figu es 38–41. Plec ocnemia kope dagha sp. no . Holo ype male, d awn in he s yle o P. a asba anica: 38 = geni alia in le
la e al iew; 39 = geni alia in do sal iew; 40 = gonopods in en al iew; 41 = phallic o gan in le la e al iew.
Plec ocnemia kope dagha Oláh & Ib ahimi,
sp. no .
(Figu es 34–41, Map 1)
Plec ocnemia ab a acou cix Malicky, 1979: Oláh
1994:282. “Tu kmenis an, Kope -Dagh M s., 10 km
Aide e, 27.VI.1992 leg. Z. Va ga (1 male, 2 e-
males; OPC). Misiden i ica ion.
Ma e ial examined. Holo ype: Tu kmenis an,
Kope -Dagh M s., Alde e, 27.VI.1992, ligh leg.
Z. Va ga (1 male, OPC). The axonomic s a us o
he 2 associa ed emales is unce ain.
Diagnosis. This is a sibling o Plec ocnemia
la issima complex, he mos eas e n membe o
he complex. Mos simila o P. ucnobi sp. no .,
Oláh e al.: On he T ichop e a o he Caucasus III
29
a Malicky d awn species o unknown o igin and
misiden i ied as P. la issima. Howe e , P. cope -
dagha sp. no . di e s om P. ucnobi sp. no . by
he di e en la e al and do sal shape o he do sal
a m. Do sal a m o pa ap oc upwa d a ching in
la e al iew wi h isible bi id apex, no downwa d
a ching and wi h dis inc isibili y o i s bi id
apex; in do sal iew he smalle spine o he bi id
apex loca ed mesad, subapicad, no la e ad and
middle.
Desc ip ion. La ge-sized Plec ocnemia spe-
cies o ligh b own body and wing colou , wi h
o ewing leng h o 12 mm. Fo ewing densely
co e ed wi h ecumben se ae.
Male geni alia. In la e al iew s e ni e IX
sub iangula , domina ing o e he en i e geni alia
a icula ing and holding he pe iphallic o gans o
used ce ci-pa ap oc complex and he gonopods.
Te gi e IX small e ically elonga ed gi ing
suppo s o he memb anous segmen X ha is
elonga ed, as long as he do sal a m o pa ap oc ,
subquad angula bo h in la e al and do sal
p o iles. Ce ci a e small se ose iangula s uc-
u es; almos igh -angled. Pa ap oc bipa i e; i s
do sal a ms sagi ally la ened, high (b oad) in
la e al iew, cha ac e ized by i s bi id e minal
well isible bo h in la e al and do sal iews; i s
en al usion o ming pa o he phalloc yp
gi ing memb anous en al suppo s o he
phallic o gan wi h a digi a e b oad-based se ose
p ocess and a hea ily scle o ized en al a m
di ec ed downwa d deep. Ven al a m o gono-
pods sho , na owing apicad in en al iew;
basodo sal pa o gonopod wi h ounded do sal
and s aigh , sligh ly conca e, apical ma gins.
Phallic o gan supplied wi h one pai o pa ame es
co uga ed on i s e y end, bo h pa ame es sligh -
ly downwa d cu ing; aedeagus wi h a mo e
pigmen ed used mesal od, possibly he pa ially
scle o ized duc us ejacula o ius wi h upwa d
b oadened apical head ep esen ing he phallo-
emmal scle i es; bo h he scle o ized mesal od
o aedeagus and he co uga ed pa ame es a e
embedded in memb anous ma ix.
E ymology. Coined om he name o locus
ypicus, a noun in apposi ion
Plec ocnemia la issima Ma yno , 1913
(Map 1)
Plec ocnemia la issima Ma yno , 1913:36–37: “1♂.
Geo gia S eam a Saka aulo, 24.VI.1911. 1♂.
Lagodekhi, 10.X.1896. Mlokose ich.”
Ma e ial examined. Geo gia, Kakhe ia, Lago-
dekhi Na ional Pa k, abo e Khizaba a, Ninos-
khe i alley, sp ing, 1250m, 41.8852°N, 46.2652°
E, 13.V.2023, leg. Gilles Vinçon (1 male, OPC).
Geo gia, M skhe a-M iane i, abo e Ukanamkha i,
nice sp ing and b ook, 1680–1730m, 42.324°N,
44.607°E, 31.VIII.2023, leg. Gilles Vinçon (1
male, OPC).
Rema k. The nomina e species o he Plec -
ocnemia la issima complex. This Caucasian spe-
cies was desc ibed by wo males, bo h collec ed in
Kakhe ia. One specimen was collec ed by Ma -
yno himsel in 1911 om Saka aulo M s. The
o he specimen was collec ed in 1896 by Mloko-
se ich om Lagodekhi. Ma yno , as usual, sup-
plied his species desc ip ion wi h d awings o
e y high quali y. Acco ding o his desc ip ion
and d awings he la e al p o ile o do sal a m o
pa ap oc is e y slende , almos spine-like.
Recen ly we ha e collec ed genuine P. la is-
sima specimens in Kakhe ia, om he egion o
he locus ypicus as well as om M skhe a-
M iane i ha ing exac ly he same geni al s uc-
u e, including he slende , spine-like la e al
p o ile o pa ap oc as p esen ed by Ma yno ’s
o iginal d awings (Oláh & Vinçon 2024). In
Malicky’s a lases (1983, 2004) he e a e se e al
d awings o a specimen wi hou any locali y
in o ma ion gi en, unde he name o P. la issima
howe e , wi h comple ely di e en la e al and
do sal p o ile o do sal a m o pa ap oc . On his
d awings he la e al p o ile o he do sal a m o
he pa ap oc is depic ed as being e y b oad and
spoon-shaped, a he han slende spine-like so
clea ly cha ac e ising he geni alia o genuine P.
la issima. The e o e, Malicky’s specimen is mis-
iden i ied and is no P. la issima. I is desc ibed
he e as a new species, Plec ocnemia ucnobi
Oláh, sp. no .
Oláh e al.: On he T ichop e a o he Caucasus III
30
Figu es 42–45. Plec ocnemia mashukana sp. no . Holo ype male: 42 = geni alia in le la e al iew; 43 = geni alia
in do sal iew; 44 = le gonopod in en al iew; 45 = phallic o gan in le la e al iew.
Plec ocnemia mashukana Oláh, sp. no .
(Figu es 42–45, Map 1)
Ma e ial examined. Holo ype: Russia, S a -
opol K ai, Pya igo sk, Besh au M s. (Fi e peaks),
a Mashuk M ., 44.05°N, 43.09°E, 21.VI.1924,
leg. Rjabo (1 male, OPC).
Diagnosis. This is a sibling in he Plec o-
cnemia la issima complex, he mos no h wes e n
membe known o he species complex. Mos
simila o he nomina e species o he complex, P.
la issima Ma yno , 1913. Pa icula ly i s speci-
a ion ai , he hea ily scle o ized do sal a m o
pa ap oc is almos iden ical, bu di e s in a
close examina ion by he i egula ly co uga ed
and obliquely no ched, no smoo h e minal ip o
his i illa ing o gan. I ce ainly has impo ance
o emale choice whe he he su ace o he
e minal egion on he i illa ing o gan is smoo h
o ha sh, scab ous due o he co uga ed no ched
su ace ex u e.
Desc ip ion. Medium-sized Plec ocnemia
species o ligh b own body and wing colou , wi h
o ewing leng h o 11 mm. Fo ewing densely
co e ed wi h ecumben se ae.
Male geni alia. In la e al iew s e ni e IX
ounded sub iangula , domina ing o e he en i e
geni alia a icula ing and holding he pe iphallic
o gans o used ce ci-pa ap oc complex and he
gonopods. Te gi e IX is small e ically elonga ed
gi ing suppo s o he memb anous segmen X
ha is sho e han he do sal a m o pa ap oc .
Ce ci a e small se ose ounded iangula s uc-
u es. Pa ap oc bipa i e; i s do sal a ms s ou
spine-like in la e al iew, sligh ly S-shaped in
do sal iew; i s en al usion o ming pa o he
phalloc yp gi ing memb anous en al suppo s
o he phallic o gan wi h a digi a e se ose p ocess
and a hea ily scle o ized en al a m di ec ed
downwa d deep. Ven al a m o gonopods medi-
um long, elonga ed o oid in en al iew wi h
ounded apical ma gin; basodo sal pa o gono-
pod wi h ounded do sal and sligh ly conca e
apical ma gin. Phallic o gan supplied wi h one
pai o pa ame es co uga ed om i s middle, bo h
pa ame es sligh ly downwa d cu ing; aedeagus
wi h a mo e pigmen ed used mesal od, possibly
he pa ially scle o ized duc us ejacula o ius wi h
upwa d b oadened apical head ep esen ing he
phallo emmal scle i es; bo h he scle o ized me-
sal od o aedeagus and he co uga ed pa ame es
a e embedded in memb anous ma ix.
Oláh e al.: On he T ichop e a o he Caucasus III
31
Figu es 46–49. Plec ocnemia ming elica sp. no . Holo ype male: 46 = geni alia in le la e al iew; 47 = geni alia
in do sal iew; 48 = le gonopod in en al iew; 49 = phallic o gan in le la e al iew.
E ymology. Coined om he name o he locus
ypicus, a noun in apposi ion. Mashuk Moun ain
is nea Pya igo sk (Fi e Peaks) no a om he
Besh au Moun ains whe e Mikhail Le mon o
had his amous duel wi h Nikolai Ma yno .
Plec ocnemia ming elica Oláh & Vinçon sp.
no .
(Figu es 46–49, Map 1, Pho os 12–13)
Ma e ial examined. Holo ype: Geo gia, Ming-
elie, abo e Koko, nice b ookle s and sp ings nea
he wa e ca chmen , on limes one subs a um,
310m, 42.6467°N, 42.2028°E, 3.V.2024, leg.
Gilles Vinçon (1 male, OPC).
Diagnosis. Mos simila o Plec ocnemia s a-
ne ica sp. no ., bu di e s by he p esence o
digi a e se ose p ocess on he do sal a m o pa a-
p oc as well as by he ounded, no s aigh
unca ed apical ma gin o he basodo sal a m o
gonopod. The sligh ly scle o ized mesal s uc u e
in he head o memb anous aedeagus disce nible
as a spine-like dis inc ly doubled, no a single
used mesal elonga ion.
Desc ip ion. La ge-sized Plec ocnemia spe-
cies o ligh b own body and wing colou wi h
o ewing leng h o 12 mm. Fo ewing densely co-
e ed wi h ecumben se ae.
Male geni alia. In la e al iew s e ni e IX
sub iangula , domina ing o e he en i e geni alia
a icula ing and holding he pe iphallic o gans o
used ce ci-pa ap oc complex and he gonopods.
Te gi e IX small e ically elonga ed gi ing sup-
po o he memb anous segmen X wi h elonga -
ed quad angula la e al and do sal p o ile. Ce ci
a e la ge ounded iangula se ose s uc u e. Pa a-
p oc bipa i e; i s do sal a ms sagi ally la ened,
e y high (b oad) in la e al iew i s en al usion
o ming pa o he phalloc yp gi ing bo h do sal
and en al suppo s o he phallic o gan, i s
en al a m di ec ed downwa d deep; do sal a m
supplied wi h a small, bu de ined digi a e and
se ose p ocess subapicad, isible bo h in la e al
and do sal iews. Ven al a m o gonopods sho ,
elonga ed o oid in en al iew wi h small api-
comesal lobe; basodo sal pa o gonopod wi h
ounded apical ma gin. Phallic o gan supplied
wi h one pai o pa ame es co uga ed on i s
apical egion, bo h pa ame es sligh ly downwa d
Oláh e al.: On he T ichop e a o he Caucasus III
32
Figu es 50–53. Plec ocnemia s ane ica sp. no . Holo ype male: 50 = geni alia in le la e al iew; 51 = geni alia
in do sal iew; 52 = le gonopod in en al iew; 53 = phallic o gan in le la e al iew.
cu ing; aedeagus less pigmen ed wi h an em-
bedded pai o spine-like od jus disce nible; bo h
he less scle o ized mesal pa o aedeagus and he
co uga ed pa ame es a e hidden in memb anous
ma ix.
E ymology. Coined om he name o he locus
ypicus, a noun in apposi ion.
Plec ocnemia s ane ica Oláh & Vinçon,
sp. no .
(Figu es 50–53, Map 1, Pho os 14–15)
Ma e ial examined. Holo ype: Geo gia, High
S ane ie, abo e Mes ia, b ook and sp ings abo e
he ish a m in cons uc ion, 1550m, 43.0477°N,
42.8091°E, 3.V.2024, leg. Gilles Vinçon (1 male,
OPC).
Diagnosis. This is a sibling o Plec ocnemia
la issima Ma yno , 1913. Has esemblance o P.
ming elica sp. no . bu di e s by he absence, no
p esence o digi a e se ose p ocess on he do sal
a m o pa ap oc as well as by he s aigh un-
ca ed, no ounded apical ma gin o he basodo sal
a m o gonopod. The sligh ly scle o ized mesal
s uc u e in he head o memb anous aedeagus
disce nible as a single used mesal elonga ion no
a dis inc pai o doubled spine-like s uc u e.
Desc ip ion. La ge-sized Plec ocnemia spe-
cies o ligh b own body and wing colou wi h
o ewing leng h o 13 mm. Fo ewing densely
co e ed wi h ecumben se ae.
Male geni alia. In la e al iew s e ni e IX
sub iangula , domina ing o e he en i e geni alia
a icula ing and holding he pe iphallic o gans o
used ce ci-pa ap oc complex and he gonopods.
Te gi e IX small e ically elonga ed gi ing sup-
po o he memb anous segmen X wi h quad -
angula la e al and do sal p o ile. Ce ci a e la ge
iangula se ose s uc u es wi h a long digi a e
highly se ose en al p ocess. Pa ap oc bipa i e;
i s do sal a ms sagi ally la ened, e y high
(b oad) in la e al iew i s en al usion o ming
pa o he phalloc yp gi ing memb anous en al
suppo s o he phallic o gan wi h a digi a e
se ose p ocess, he hea ily scle o ized en al a m
di ec ed downwa d deep. Ven al a m o gono-
pods sho , elonga ed o oid in en al iew wi h
ounded apical ma gin; basodo sal pa o gono-
pod wi h s aigh unca ed apical ma gin. Phallic
o gan supplied wi h one pai o pa ame es co u-
ga ed om middle, bo h pa ame es sligh ly down-
wa d cu ing; aedeagus wi h a mo e pigmen ed
Oláh e al.: On he T ichop e a o he Caucasus III
33
used mesal od, possibly he pa ially scle o ized
duc us ejacula o ius wi h upwa d b oadened api-
cal head; bo h he scle o ized mesal od o aede-
agus and he co uga ed pa ame es a e embedded
in memb anous ma ix.
E ymology. Coined om he name o locus
ypicus, a noun in apposi ion.
Plec ocnemia ucnobi Oláh, sp. no .
Plec ocnemia la issima Ma yno , 1913: Malicky
1983:85. D awings, o iginal igu es wi hou any
locali y in o ma ion abou he specimen d awn.
Misiden i ica ion.
Plec ocnemia la issima Ma yno , 1913: Malicky
2004:102. D awings, o iginal igu es wi hou any
locali y in o ma ion abou he specimen d awn.
Misiden i ica ion.
Ma e ial. Holo ype. Malicky 1983: 85, 2004:
102. D awings om a Caucasian specimen o
unknown locali y, deposi ed in Malicky’s P i a e
Collec ion (1 male, MPC).
Diagnosis. This species p esen ed by Malicky
(1983, 2004) as Plec ocnemia la issima Ma -
yno , 1913 wi hou any collec ing da a was al-
eady ques ioned and ema ked (Oláh & Vinçon
2024). Because he do sal a m o he pa ap oc is
spine-like, ha is slende in Ma yno ’s o iginal
species desc ip ion and d awing, and on a eal P.
la issima specimen collec ed newly nea by he
locus ypicus, no b oad and no wi h an ad-
di ional subapical spine isible only in do sal
iew on he d awing. This addi ional subapical
spine on he do sal a m o pa ap oc is unique, no
p esen on any o he o he membe s o he
species complex.
Desc ip ion. S e ni e IX sub iangula , e gi e
IX e y sho (na ow) long digi i o m and less
scle o ized. Segmen X memb anous elonga ed.
Ce ci almos sha p iangula in la e al iew. Pa-
ap oc bipa i e; i s do sal a ms sagi ally la -
ened, e y high (b oad) in la e al iew i s en al
usion o ming pa o he phalloc yp gi ing
memb anous en al suppo s o he phallic o gan
wi h a digi a e se ose p ocess, he hea ily scle-
o ized en al a m di ec ed downwa d deep. In
la e al iew basodo sal p ocess o gonopod wi h
unca ed apical ma gin; en al a m o gonopod
sho and b oad. Phallic o gan wi h a pai o long
pa ame es; i s apical egion co uga ed no ched.
E ymology. “ucnobi” unknown in Geo gian
language e e ing o he d awn specimen wi hou
collec ing de ails.
Plec ocnemia zeka ia new species complex
(Map 2)
Ha ing sho e ce ci, longe gonopod and
hooded basodo sal lobe on gonopods his new
species complex belongs o he Plec ocnemia
conspe sa species g oup. They ha e wo pai s o
pa ame es wi h co uga ed, no ched e minal e-
gion. Incipien , sibling, o sis e species o his
new species complex is dis inguished easily and
dis inc ly by hei adap i e specia ion ai , by he
sub ly, bu s ably di e ged less scle o ized abb e-
ia ed a m o pa ap oc .
Plec ocnemia gomis a Oláh & Vinçon, sp. no .
(Figu es 54–57, Map 2, Pho os 16–17)
Ma e ial examined. Geo gia, Adja a, abo e
Gomism a, nice b ook, 2120–2150m, 41.8108°N,
42.1821°E, 11.VIII.2024, leg. Gilles Vinçon (1
male, OPC).
Diagnosis. Close bo h o Plec ocnemia izei-
ensis Sipahile , 1987 desc ibed om Tu key (nea
Rize) and o Plec ocnemia zeka ia Oláh &
Vinçon, 2024 desc ibed om Geo gia, Ime e i.
Howe e di e s om bo h ha ing do sal a m o
pa ap oc bilobed as well as wo pai s o spines o
endo heca di e en ly shaped and co uga ed
pa ame es p esen on phallic o gan no lacking.
Desc ip ion. Small-sized Plec ocnemia spe-
cies o ligh b own, almos yellowish body and
wing colou wi h o ewing leng h o 7 mm.
Fo ewing densely co e ed wi h ecumben se ae.
Male geni alia. In la e al iew s e ni e IX
ounded sub iangula , domina ing o e he en i e
Oláh e al.: On he T ichop e a o he Caucasus III
34
Figu es 54–57. Plec ocnemia gomis a sp. no . Holo ype male: 54 = geni alia in le la e al iew; 55 = geni alia
in do sal iew; 56 = le gonopod in en al iew; 57 = phallic o gan in le la e al iew.
geni alia a icula ing and holding he pe iphallic
o gans o ce ci-pa ap oc used complex and he
gonopods. Te gi e IX small e ically elonga ed
gi ing suppo o he memb anous segmen X.
Ce ci la ge oli o m. Pa ap oc bipa i e e ically
elonga ed i s do sal a ms highly educed o ming
a small bilobed s uc u e, i s en al a m di ec ed
downwa d. Gonopods e y sho , subquad an-
gula in en al iew wi h ounded apical ma gin;
i s do sobasal pa wi h excised apical ma gin.
Phallic o gan supplied wi h wo pai s o spine-like
s uc u es embedded in endo heca and a pai o
ee co uga ed pa ame es jus disce nible, less
pigmen ed and less scle o ized.
E ymology. Coined om he name o locus
ypicus, a noun in apposi ion.
Plec ocnemia kin isha Oláh & Memishishi,
sp. no .
(Figu es 58–61, Map 2, Pho os 7–8)
Ma e ial examined. Holo ype: Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 11, 1637m, 41.74769°N, 42.0951°E,
29.VI.–13.VII. 2018, leg. GGBC membe s (1
male, OPC). Pa a ype: Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
13.VII–27.VII. 2018, leg. GGBC membe s (1
male, IZISUT). Geo gia, Lesse Caucasus, Ad-
ja a, Kin ishi, abo e he wa e all, malaise ap
11, 1637m, 41.74769°N, 42.0951°E, 24.VIII.–
7.IX. 2018, leg. GGBC membe s (3 males, 1
emale; OPC).
Diagnosis. Close o bo h Plec ocnemia izei-
ensis Sipahile , 1987 desc ibed om Tu key (nea
Rize) and o Plec ocnemia zeka ia Oláh &
Vinçon, 2024 desc ibed om Geo gia, Ime e i.
Howe e , di e s om bo h by ha ing he educed
and less scle o ized do sal a m o pa ap oc jus a
ounded, supe icially subdi ided s uc u e wi h-
ou any lobes jus wi h only a small upwa d
u ning lap is p esen which di icul o disce n.
Desc ip ion. Small-sized Plec ocnemia spe-
cies o ligh b own, almos yellowish body and
wing colou , wi h o ewing leng h o 8 mm.
Fo ewing densely co e ed wi h ecumben se ae.
Oláh e al.: On he T ichop e a o he Caucasus III
35
Figu es 58–61. Plec ocnemia kin isha sp. no . Holo ype male: 58 = geni alia in le la e al iew; 59 = geni alia
in do sal iew; 60 = le gonopod in en al iew; 61 = phallic o gan in le la e al iew.
Map 2. Plec ocnemia zeka ia species complex (4 species): P. gomis a sp. no ., P. kin isha sp. no .
P. izeiensis Sipahile , 1987, P. zeka ia 2024.
Oláh e al.: On he T ichop e a o he Caucasus III
36
Male geni alia. In la e al iew s e ni e IX
ounded subquad angula , domina ing o e he
en i e geni alia a icula ing and holding he pe i-
phallic o gans o ce ci-pa ap oc used complex
and he gonopods. Te gi e IX small indis inc and
e ically elonga ed gi ing suppo o he mem-
b anous segmen X, e gal and pa ap oc al com-
plex. Ce ci la ge oli o m. Pa ap oc bipa i e e -
ically elonga ed, i s do sal a ms highly educed
o ming a small ounded s uc u e subdi ided
supe icially, i s en al a m di ec ed downwa d.
Gonopods e y sho , subquad angula in en al
iew wi h ounded apical ma gin; i s do sobasal
pa wi h downwa d di ec ed and excised apical
ma gins. Phallic o gan supplied wi h wo pai s o
spine-like co uga ed pa ame es jus disce nible,
less pigmen ed and less scle o ized.
E ymology. Coined om he name o locus
ypicus, a noun in apposi ion.
Plec ocnemia izeiensis Sipahile , 1987
(Map 2)
Plec ocnemia izeiensis Sipahile , 1987: “Holo ype ♂:
Tü kiye, Rize, Ikizde e, Aşaği Vice Köyü, 19.VII.
1984. Pa a ype: Rize, Ikizde e, Çamhhemşin? Ça ,
1250 m, (in ligh ) 13.VII.1984 leg. and coll. F.
Sipahile .” “This new specie is well cha ac e ized
by he shape o he in e io and p eanal appenda-
ges. I is somewha ela ed o P. b e is McL. and P.
kydon Mal. (Malicky 1983:83), bu he di e ences
o he geni alia, especially he in e io appendages
and he aedeagus dis inguish his new species.”
Ma e ial examined. Geo gia, Sam skhe-Ja a-
khe i, abo e U kisubani, nice sp ing, 1920m,
41.6748°N, 42.5691°E, 8.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Rema k. Fi s epo om Geo gia.
Plec ocnemia zeka ia Oláh & Vinçon, 2024
(Map 2, Pho os 26–29)
Ma e ial examined. Geo gia, Sam skhe-Ja a-
khe i, abo e U kisubani, nice sp ing, 1920m,
41.6748°N, 42.5691°E, 8.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Hyd opsychidae Cu is, 1835
Diplec oninae Ulme , 1951
Diplec ona geo gica Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, Adja a, Cha-
kha i, sp ing, 90m, 41.8791 41.91601, 29.IV.
2024, leg. A. Memishishi (8 males, 2 emales;
IZISUT). Geo gia, Adja a, Dunga, wa e all,
880m, 41.47544°N, 41.85622°E, 30.IV.2024, leg.
A. Memishishi (1 male, IZISUT). Geo gia, Ad-
ja a, Kin ishi, sp ing, 270m, 41.77359°N,
41.96808°E, 29.IV.2024, leg. A. Memishishi (1
male, IZISUT). Geo gia, Adja a, abo e Keda,
b ook, 430m, 41.5844°N, 41.964°E, 1.V.2024,
leg. Gilles Vinçon (1 male, 9 emales; OPC).
Geo gia, Adja a, M i ala Na ional Pa c, nice
b ook, 160m, 41.697°N, 41.823°E, 30.IV.2024,
leg. Gilles Vinçon (1 male, OPC). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 11, 1637m, 41.74769°N, 42.0951°E,
15.VI. – 29.VI. 2018, leg. GGBC membe (38
males, IZISUT; 9 males, OPC).
Diplec ona obus a Ma yno , 1934
Ma e ial examined. Geo gia, Ming elia, abo e
Koko, nice b ookle and sp ings nea he wa e
ca chmen , on limes one subs a um, 330m,
42.648°N, 42.2033°E, 3.V.2024, leg. Gilles Vin-
çon (1 male, OPC). Geo gia, Ming elie-High S a-
ne ia, d ipping ocks, abo e Ingu i lake, 590m,
42.8585°N, 42.0336°E, 3.V.2024, leg. Gilles
Vinçon (1 male, OPC). Geo gia, Sameg elo – Ze-
mo S ane i, Deba adzeni, sp ing, 375m,
42.59264°N, 42.32199°E, 28.IV.2024, leg. A.
Memishishi (1 male, IZISUT). Geo gia, Ime e i,
Kh e e i, sp ing, 530m, 42.3740°N, 43.4378°E,
25.IV.2024, leg. A. Memishishi (1 male,
IZISUT). Geo gia, Racha-Lechkhumi and Lowe
S ane i, Babili, sp ing, 790m, 42.79156°N,
42.74598°E, 27.IV.2024, leg. A. Memishishi (5
males, 1 emale; IZISUT). Geo gia, Ming elie-
High S ane ia, Sou h Toba i, b ookle and cas
Oláh e al.: On he T ichop e a o he Caucasus III
37
Figu es 62–66. Cheuma opsyche bushe ica sp. no . Holo ype male: 62 = geni alia in le la e al iew; 63 = geni alia in do sal
iew; 64 = le gonopod in en al iew; 65 = phallic o gan in le la e al iew; 66 = phallic o gan in en al iew.
cade, 650m, 42.9571°N, 42.1952°E, 3.V.2024,
leg. Gilles Vinçon (6 males, 8 emales; OPC).
Hyd opsychinae Cu is, 1835
Cheuma opsyche Walleng en, 1891
Cheuma opsyche bushe ica Oláh, sp. no .
(Figu es 62–66)
Cheuma opsyche la ella a Mey, 2004: Oláh 2010:79.
Misiden i ica ion.
Ma e ial examined. I an, P o ince Busheh , 5
km S o Del az, 5−6.V.2001, leg. GY. Fábián (1
male, OPC). Pa a ypes: same as holo ype (5
males, OPC, 5 males, HNHM).
Diagnosis. Cheuma opsyche bushe ica sp.
no ., a species o Cheuma opsyche lepida species
g oup was de e mined ea lie by he i s au ho
(Oláh, 2004) as C. la ella a Mey, bu ha ing
examined genuine compa a i e specimens om
Geo gia and Eas Tu key by ine phenomics, he
specimens om Bushe P o ince, I an ep esen a
new species wi h i s ex emely elonga ed
apico en al se ose lobe as well as wi h he
di e en la e al p o ile o phallic o gan.
Desc ip ion. B own colou ed small-sized spe-
cies wi h o ewing leng h o 7 mm; o ewing
wi hou any disce nible pa e n.
Male geni alia. Abdominal segmen IX used
annula , a he long; bo h an e io and pos e io
ma gin a ci o m, esul ed in an ex emely sho
do sum and longe en um; pos e io lobe abo e
coxopodi e less p oduced ounded and high;
in e segmen al dep ession be ween he nin h and
en h segmen s p esen , bu low. Body o segmen
X longe han high; ci cula la e al se ose a ea
(ce ci) in apical posi ion; apico en al se ose lobe
disce nible bo h in la e al and do sal iews as a
slende a he elonga ed obliquely upwa d u ning
s uc u e; in do sal iew exhibi s peculia pa e n
o long digi a e od wi h e y wide do sal in e -
lobula gap; unse osed, smoo h mesocaudal lobe
e y p oduced ounded in do sal iew. The basal
segmen o he gonopods almos equally wide and
s aigh od wi h only sligh ly dila ed apical end-
ing; ha pago, he e minal segmen b oade basad
and na owing apicad bo h in la e al and en al
iews. Phallic appa a us wi h a he obus phallo-
heca wi h en o subapical keel; pai o endo-
hecal scle i es well de eloped elonga ed o oid.
E ymology. Coined om he name o he locus
ypicus o he holo ype, a noun in apposi ion.
Oláh e al.: On he T ichop e a o he Caucasus III
44
Figu es 92–96. Hyd opsyche akhalchala sp. no . Holo ype male: 92 = geni alia in le la e al iew; 93 = geni alia in do sal iew;
94 = le gonopod in en al iew; 95 = phallic o gan in le la e al iew; 96 = phallic o gan in en al iew.
Hyd opsyche akhalchala Oláh & Vinçon,
sp. no .
(Figu es 92–96, Pho o 6)
Ma e ial examined. Holo ype: Geo gia, Racha
-Lechkhumi and Lowe -S ane i, Akhalchala, nice
b ook and sp ing, 2120m, 42.6928°N, 42.6447°E,
13.VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Diagnosis. This da k new species has a geni al
s uc u e simila o Hyd opsyche ejsaka Oláh,
2020, bu di e s by ha ing he median keel on
e gi e IX wi h a long iangula clea window
om basad o midway, lacking in H. ejsaka; by
he in e segmen al p o ile be ween segmen s IX
and X simple igh angled, no a deep gap; by less
downwa d di ec ed apical egion o phallic o gan
as well as by he sligh ly na owing, no pa allel-
sided en al p o ile o phallic head.
Desc ip ion. Male. Body da k, do sal ho acic
scle i es da ke . Wings da k b own, wi h dense
ecumben hai s. Maxilla y palp o mula I-III-II-
IV-V. Spu o mula 244. Fo ewing leng h 9 mm
Male geni alia. Segmen IX used annula and
sho ; i s median keel b oad, V-shaped clea
window om basad almos o midway, ounded
apicad wi h g anulose do sal su ace; apical lobe
on pos e ola e al ma gin sho and ounded,
an e io ma gin con ex. In e segmen al p o ile
be ween he nin h and en h segmen s low, igh
angled. Segmen X sho , subquad angula in
do sal iew; la e al se ose a ea, he ce ci used
wi h en oapical se ose lobe, loca ed in pos e ad
posi ion; do soapical se ose lobes educed o he
se ose an e io a ea o he unse ose do sola e al
im o segmen X in la e al iew. The coxopodi
o he gonopod longe han he apex o segmen
X, sligh ly S-shaped; ha pago spa ula e in en al
iew and wi h na owing apex in la e al iew.
Phallic o gan wi h sligh ly downwa d di ec ed
apical egion and u he na owed head in la e al
iew; head sligh ly na owing and p onounced
iangula subapical la e al p ojec ion in en al
iew.
E ymology. Coined om he name o he ype
locali y, a noun in apposi ion.
Hyd opsyche ejsaka Oláh, 2020
Ma e ial examined. Geo gia, Sameg elo -
Zemo S ane i, Deba adzeni, sp ing, 375m,
42.59264°N, 42.32199°E, 28.IV.2024, leg. A.
Oláh e al.: On he T ichop e a o he Caucasus III
45
Figu es 97–101. Hyd opsyche gode dza sp. no . Holo ype male: 97 = geni alia in le la e al iew; 98 = geni alia in do sal iew;
99 = le gonopod in en al iew; 100 = phallic o gan in le la e al iew; 101 = phallic o gan in en al iew.
Memishishi (2 males, IZISUT; 1 male, OPC).
Geo gia, Racha-Lechkhumi and Lowe S ane i,
U a i, sp ing, 1000m, 42.65771°N, 43.29736°E,
26.IV.2024, leg. A. Memishishi (1 male, IZI
SUT).
Rema k. This species was only known om
he Lesse Caucasus (K emo Ka li). We epo i
o he i s ime om he High Caucasus.
Hyd opsyche gode dza Oláh & Vinçon, sp. no .
(Figu es 97–101, Pho os 3–4)
Ma e ial examined. Holo ype: Geo gia, Ad-
ja a, Gode dzi Pass, below G een Lake, la ge
b ook below he swamp, 2050m, 41.682°N,
42.5°E, 8.VIII.2024, leg. Gilles Vinçon (1 male,
OPC). Pa a ypes: same as holo ype (8 males,
OPC).
Diagnosis. This da k, new species has a geni al
s uc u e simila o Hyd opsyche adja ia Oláh &
Vinçon, 2024, bu di e s by ha ing he median
keel on e gi e IX b oadening apicad, no pa allel-
sided in do sal iew; he apicomesal lobe o
segmen IX ounded no iangula ; he unse ose
do sola e al im o segmen X a iously la , no
iangula in la e al iew. Cu a u e o he phallic
o gan di e en ly pa e ned. Howe e , hese di-
e ged cha ac e s a es ha e some ange o a i-
a ion a Hyd opsyche gode dza sp. no . and hese
a ia ion anges a e no known a Hyd opsyche
adja ia ep esen ed only by holo ype. New col-
lec ions a e equi ed o compa e and o con i m
hei exac axonomic s a us.
Desc ip ion. Male. Body da k, do sal ho acic
scle i es da ke , almos black. Wings da k b own,
wi hou p onounced pa e n. Maxilla y palp o -
mula I-III-II-IV-V. Spu o mula 244. Fo ewing
leng h 8 mm.
Male geni alia. Segmen IX used annula and
sho ; i s median keel b oadening apicad, mag-
ni ude o his b oadening cha ac e a ies among
pa a ypes; apical lobe on pos e ola e al ma gin
la ounded, an e io ma gin con ex. In e seg-
men al p o ile be ween he nin h and en h seg-
men s modi ied in o a iny deep and sho gap.
Segmen X sho , ounded quad angula in do sal
iew; la e al se ose a ea, he ce ci used wi h
en oapical se ose lobe, loca ed in pos e ad po-
si ion; sub iangula in la e al iew; do soapical
se ose lobes educed o he se ose an e io a ea o
he unse ose do sola e al im o segmen X in
la e al iew. The coxopodi o he gonopod li le
longe han he apex o segmen X; ha pago
Oláh e al.: On he T ichop e a o he Caucasus III
46
Figu es 102–106. Hyd opsyche mes ia sp. no . Holo ype male: 102 = geni alia in le la e al iew; 103 = geni alia in do sal
iew; 104 = le gonopod in en al iew; 105 = phallic o gan in le la e al iew; 106 = phallic o gan in en al iew.
spa ula e in en al iew and na owing en ad in
la e al iew. Phallic o gan wi h downwa d di ec -
ed apical egion and upwa d di ec ed blun head
in la e al iew; p onounced semici cula subapical
la e al p ojec ion in en al iew.
E ymology. Coined om he name o he ype
locali y, a noun in apposi ion.
Hyd opsyche mes ia Oláh & Vinçon, sp. no .
(Figu es 102–106, Pho os 14–15)
Ma e ial examined. Holo ype: Geo gia, High
S ane i, abo e Mes ia, b ook and sp ing abo e
he ish a m in cons uc ion, 1550m, 43.0477°N,
42.8091°E, 16.VIII.2024, leg. Gilles Vinçon (1
male, OPC). Pa a ype: same as holo ype (1 male,
OPC).
Diagnosis. This da k new species has a geni al
s uc u e simila o Hyd opsyche akhalchala sp.
no ., bu di e s by ha ing he median keel on
e gi e IX na owe and wi hou long iangula
basal window; by he mo e obus , b oad phallic
o gan wi h pa allel-sided, no na owing head o
phallic o gan and by ab up ly, no g adually
p oduced iangula subapical la e al p ojec ion on
he phallic o gan in en al iew.
Desc ip ion. Male. Small species. Body da k,
do sal ho acic scle i es da ke . Wings da k
b own, wi h dense ecumben hai s. Maxilla y
palp o mula I-III-II-IV-V. Spu o mula 244.
Fo ewing leng h 7 mm.
Male geni alia. Segmen IX used annula and
sho ; i s median keel sligh ly na owe basad
wi h g anulose do sal su ace; apical lobe on
pos e ola e al ma gin sho and ounded, an e io
ma gin con ex. In e segmen al p o ile be ween
he nin h and en h segmen s deep and sho . Seg-
men X sho , ounded quad angula in do sal
iew; la e al se ose a ea, he ce ci used wi h
en oapical se ose lobe, loca ed in pos e ad posi-
ion; do soapical se ose lobes educed o he se-
ose an e io a ea o he unse ose do sola e al im
o segmen X in la e al iew. The coxopodi o he
gonopod longe han he apex o segmen X,
sligh ly widened apical hal ; ha pago spa ula e in
en al iew and wi h sligh ly na owing apex in
la e al iew. Phallic o gan wi h downwa d di-
ec ed apical egion and li le na owed head in
la e al iew; head almos pa allel-sided wi h small
iangula subapical la e al p ojec ion in en al
iew.
E ymology. Coined om he name o he ype
locali y, a noun in apposi ion.
Oláh e al.: On he T ichop e a o he Caucasus III
47
Figu es 107–111. Hyd opsyche sabna a sp. no . Holo ype male: 107 = geni alia in le la e al iew; 108 = geni alia in do sal
iew; 109 = le gonopod in en al iew; 110 = phallic o gan in le la e al iew; 111 = phallic o gan in en al iew.
Hyd opsyche sabna a Oláh & Memishishi,
sp. no .
(Figu es 107–111, Pho os 7–8)
Ma e ial examined. Holo ype: Geo gia, Ad-
ja a, Tsablna i, wa e all, 480m, 41.67055°N,
41.87795°E, 29.IV.2024, leg. A. Memishishi (1
male, OPC). Pa a ype: Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 11, 1637m, 41.74769°N, 42.0951°E, 15.VI.–
28.VII. 2018, leg. GGBC membe s (1 male,
OPC).
Diagnosis. This new species belongs o he
Hyd opsyche ins abilis subg oup o he H. angus-
ipennis species g oup and is cha ac e ized by
used scle o ized endo hecal p ocesses, wi h digi-
i o m apico en al se ose lobe and wi h angula
subapical la e al p ojec ions be o e he cle apex
o he phallo heca (Oláh & Johanson, 2008). The
angula subapical p ojec ion is an e ad di ec ed.
Hyd opsyche sabna a sp. no . is cha ac e ized by
he e y p onounced, bu e y low basal cu e o
he phallobase. Such a low and p onounced basal
cu e on he phallic o gan is p esen only in a ew
species o he Hyd opsyche ins abilis subg oup:
he nomina e species H. ins abilis (Cu is, 1834);
H. jo danensis Tjede , 1951; H. o duensis Sipa-
hile , 1987; H. pe is e ica Bo osaneanu & Ma in-
ko ic, 1966; H. pe seus Malicky, 2001; H.
pic e o um Bo osaneanu & Schmid, 1973. How-
e e , H. sabna a sp. no . di e s om each by
he elonga ed iangula do sal shape o he do sal
keel o segmen IX as well as by he an e ad
di ec ed la e al p ojec ions on he subapical egion
o he phallic o gan.
Desc ip ion. Male (in alcohol). Medium sized
species. Body b own, bu do sal ho acic scle i es
da ke . Fo ewing b own, wi hou any disce nible
pa e n. Leng h o o ewing is 12 mm.
Male geni alia. Segmen IX used annula and
sho , pa icula ly on i s en um; i s median keel
b oad-based and g adually na owing apicad
o ming an elonga ed iangula shape in do sal
iew wi h g anulose do sal su ace; apical lobe on
pos e ola e al ma gin well p oduced, ounded, al-
mos hal ci cula . In e segmen al p o ile be ween
he nin h and en h segmen s sho , o ming a
shallow conca i y. Segmen X as long as he do -
sal keel o segmen IX, wi h p oduced elonga ed
se aless wingle s in la e al iew ha ing se ose
p oximal egion and elonga ed ough in do sal
iew; la e al se ose a ea, he ce ci ci cula and
loca ed in apical posi ion; en oapical se ose
lobes indis inc ; digi i o m p ocess sho and s ou
wi h apical se ae. The coxopodi e o he gonopod
Oláh e al.: On he T ichop e a o he Caucasus III
48
Figu es 112–114. Agape us akhalchal sp. no . Holo ype male: 112 = geni alia in le la e al iew; 113 = le gonopod
in en al iew; 114 = phallic o gan in le la e al iew.
as long as he apex o segmen X, ha pago
pa allel-sided in la e al iew wi h sligh apical
dila a ion and mesad cu ing cla a e in en al
iew. Phallic o gan almos a pa allel-sided ube,
basal bending e y p onounced, bu e y low in
la e al iew; he subapical la e al p ojec ion on
he phallic head an e ad di ec ed, apical egion
pa allel-sided.
E ymology. Coined om he name o he locus
ypicus, as a noun in apposi ion.
Glossosoma idae Walleng en, 1891
Agape us Cu is, 1834
Agape us caucasicus Ma yno , 1913
Ma e ial examined. Geo gia, Sam skhe-Ja a-
khe i, Zeda Va dzia, nice b ook and sp ing,
1410m, 41.383°N, 43.258°E, 25.IV.2024, leg.
Gilles Vinçon (13 males, OPC). Geo gia, K emo
Ka li, Kh ami Ri e a Dashbashi cascades,
1400m, 41.5945°N, 44.1238°E, 24.IV.2024, leg.
Gilles Vinçon (2 males, OPC).
Rema k. This species desc ibed om Tu key
and also epo ed om A menia (Oláh 1985). I is
newly epo ed he e om Geo gia.
Agape us gou iensis species complex
(Map 3)
This species complex is cha ac e ized by
elonga ed quad a ic gonopods wi h ine and
de ini e di e gences. Un o una ely he p obable
specia ion ai in he Agape us gou iensis com-
plex, he in e nal aedeagal scle i e is e y badly
isible e en a highe magni ica ion. I is di icul
o examine. These in e nal phallic scle i es o
spines a e no e iden immedia ely, isible mos ly
by labo ious sea ching. Mo eo e aedeagal
scle i e is some imes accompanied by da k
pigmen ed long ili o m do sal b ace s uc u e,
p obably he a iously en o ced, almos sepa a ely
embodied suppo ing do sum o aedeagus and by
ibe -composed a iously shaped s uc u es,
en ad o do sad o he aedeagal scle i e, depend-
ing p obably on he copula o y s a e o he male.
Some imes he do sal band is ligh ly scle o ized,
indis inc . The aedeagus is deeply loca ed inside
he co e age o segmen s IX and X no ans-
pa en enough, and only seldom pullable ou om
he co e o da k pigmen ed segmen IX. The e-
o e i s exac s uc u e is p ac ically indisce nible,
jus i s ou line isible by mul iply mace a ion,
mic osu gical manipula ion unde binocula and
Oláh e al.: On he T ichop e a o he Caucasus III
49
Map 3. Agape us gou iensis species complex (5 species): A. akhalchal sp. no ., A. bakhma no ., A. go gi ensis Sipahile , 1996,
A. gou iensis Oláh & Vinçon, 2020; A. lashijal sp. no .
examina ion by compound mic oscopes. I you
ha e enough specimens and pa ience o epea
many imes and a e lucky enough you can see and
d aw he h ee s uc u es p ope ly. The combi-
na ion o badly isible in e nal aedeagal s uc u es
and he well isible bu sub le di e gence o go-
nopods oge he help o ecognise and o desc ibe
siblings in he Agape us gou iensis species com-
plex. Fi e sibling species comp ise he complex:
Agape us akhalchal Oláh & Vinçon, sp. no ., A.
bakhma Oláh & Vinçon sp. no ., A. go gi ensis
Sipahile , 1996, A. gou iensis Oláh & Vinçon,
2020; A. lashijal Memishishi & Oláh sp. no .
Agape us akhalchal Oláh & Vinçon, sp. no .
(Figu es 112–114, Map 3, Pho o 6)
Ma e ial examined. Holo ype: Geo gia, Racha
-Lechkumi and Lowe -S ane i, Akhalchala, b ook
and sp ing, 2170m, 42.6947°N, 42.6463°E, 13.
VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Diagnosis. This new species is close o Aga-
pe us gou iensis Oláh & Vinçon, 2020 desc ibed
om Gou ia egion o Geo gia, bu di e s by
bo h he la e al and en al iews o he gono-
pods; unca ed apical ma gin egula ly s aigh
e ical wi h igh angles, no s aigh oblique and
no wi h ounded angles; di e s also by he shape
o he scle i e o he aedeagus being poin ed, no
i id.
Desc ip ion. Male (in alcohol). Da k b own
animal, wi h legs and en e sligh ly ligh e . Max-
illa y palp o mula: I-II-IV-V-III, second segmen
wi h globula mesola e al p ojec ion. Wing mem-
b ane b own; o ewing leng h 4 mm; Fo k I on
hindwing los . Blis e -like p o ube ance on he
do sal ma gin o s e ni e V absen , upwa d di ec -
ed ilamen p esen ; en al p ocess on s e ni e VI
sho .
Male geni alia. Segmen IX synscle o ized
wi h sligh ly conca e an e io and con ex pos
Oláh e al.: On he T ichop e a o he Caucasus III
50
Figu es 115–118. Agape us bakhma sp. no . Holo ype male: 115 = geni alia in le la e al iew; 116 = le gonopod in en al
iew; 117 = phallic o gan in le la e al iew; 118 = phallic o gan in do sal iew.
e io ma gins in la e al iew; i s do sum hal as
long as en um. Segmen X less pigmen ed,
deeply di ided up o he basemen in do sal iew.
Ce ci elonga ed. Gonopods quad angula , elon-
ga ed cla a e, s aigh e ically unca ed in la-
e al iew. Aedeagus indis inc ube wi h dis inc
poin ed scle i e inside.
E ymology. Named a e he egion o ype he
locali y, a noun in apposi ion.
Agape us bakhma Oláh & Vinçon, sp. no .
(Figu es 115–118, Map 3, Pho os 18–19)
Agape us gou iensis Oláh & Vinçon, in Oláh e al.
2020:43–45: Oláh & Vinçon 2024:25. Geo gia,
Gou ia, NE Bakhma o, nice b ook ibu a y o
Sashualo Ri e , 1990–2000m, 41.859°N, 42.357°E,
27.IX.2022, leg. Gilles Vinçon (1 male, OPC).
Misiden i ica ion.
Ma e ial examined. Holo ype: Geo gia, Gou-
ia, NE Bakhma o, nice b ook ibu a y o
Sashualo Ri e , 1990–2000m, 41.859°N, 42.357°
E, 27.IX.2022, leg. Gilles Vinçon (1 male, OPC).
Diagnosis. This new species is close o Aga-
pe us gou iensis Oláh & Vinçon, 2020 desc ibed
om Gou ia egion o Geo gia, bu di e s by
bo h he la e al and en al iews o he
gonopods; di e s also by he shape o he scle i e
o he aedeagus.
Desc ip ion. Male (in alcohol). Da k b own
animal, wi h legs and en e sligh ly ligh e .
Maxilla y palp o mula: I-II-IV-V-III, second
segmen wi h globula mesola e al p ojec ion.
Wing memb ane b own; o ewing leng h 7 mm;
Fo k I on hindwing los . Blis e -like p o ube ance
on he do sal ma gin o s e ni e V absen , upwa d
di ec ed ilamen p esen ; en al p ocess on
s e ni e VI sho .
Male geni alia. Segmen IX synscle o ized
wi h sligh ly con ex an e io and mo e obliquely
con ex pos e io ma gins in la e al iew; i s do -
sum one hi d as long as en um. Segmen X less
pigmen ed, deeply di ided up o he basemen in
do sal iew. Ce ci elonga ed. Gonopods quad an-
gula , obliquely unca ed in la e al iew. Aedea-
gus indis inc ube wi h jus disce nible pa e n o
scle i e head inside; do sum wi h se ose pla e.
Oláh e al.: On he T ichop e a o he Caucasus III
51
Figu es 119–120. Agape us gou iensis Oláh & Vinçon, 2020. Holo ype male ed awn: 119 = geni alia in le la e al iew;
120 = phallic o gan in le la e al iew.
E ymology. Named a e he egion o he ype
locali y, noun in apposi ion.
Agape us gou iensis Oláh & Vinçon, 2020
(Figu es 119–120, Map 3)
Ma e ial examined. Geo gia, Gou ia, oad o
Gomism a, sp ings, 1900m, 41.8317°N, 42.1556°
E, 11.VIII.2024, leg. Gilles Vinçon (12 males,
OPC).
Agape us lashijal Memishishi & Oláh, sp. no .
(Figu es 121–125, Map 3, Pho os 7–8)
Ma e ial examined. Holo ype: Geo gia, High
Caucasus, Ime e i, Lashijala, sp ing, 590m,
42.64795°N, 42.84235°E, 26.IV.2024, leg. A.
Memishishi (1 male, IZISUT). Pa a ypes: same as
holo ype (7 males, 3 males IZISUT). Geo gia,
Lesse Caucasus, Adja a, Kin ishi, abo e he
wa e all, malaise ap 5, 41.743426°N, 42.0
81814°E, 1252m, 19.V.–1.VI.2018, leg. GGBC
membe s e al. (1 male, IZISUT).
Diagnosis. This new species is close o
Agape us gou iensis Oláh & Vinçon, 2020 de-
sc ibed om Gou ia egion o Geo gia, bu di e s
by bo h he la e al iew o he gonopods; do -
soapical ma gin o gonopod p oduced in o a
oundly lobe, absen a A. gou iensis; in en al
iew dis inc ly p onounced single spine p esen a
he e minal end in he apex o gonopod like
Agape us go gi ensis (Sipahile , 1996), absen a
A. gou iensis; A. lashijal sp. no . di e s om A.
gou iensis also by he shape o aedeagal scle i e
wi h ipoin ed, no wi h i id apical ending.
Desc ip ion. Male (in alcohol). Da k b own
animal, wi h legs and en e sligh ly ligh e .
Maxilla y palp o mula: I-II-IV-V-III, second seg-
men wi h globula mesola e al p ojec ion. Wing
memb ane b own; o ewing leng h 6 mm; Fo k I
on hindwing los . Blis e - like p o ube ance on he
do sal ma gin o s e ni e V absen , upwa d di-
ec ed ilamen p esen ; en al p ocess on s e ni e
VI sho .
Male geni alia. Segmen IX synscle o ized
wi h sligh ly conca e an e io and obliquely wa -
ing pos e io ma gins in la e al iew; i s do sum
hal as long as en um. Segmen X less pig-
men ed, deeply di ided nea o he basemen in
do sal iew. Ce ci elonga ed whi h a sha p poin
do sobasad. Gonopods quad angula , elonga ed
cla a e, sligh ly obliquely unca ed in la e al
iew. Aedeagus indis inc ube wi h dis inc i-
poin ed aedeagal scle i e inside.
E ymology. Named a e he egion o ype
locali y, noun in apposi ion.
Rema ks. Final axonomic s a us o he single
pa a ypes om Adja a egion, Kin ishi has o be
con i med a e ha ing mo e specimens o a
Oláh e al.: On he T ichop e a o he Caucasus III
52
Figu es 121–125. Agape us lashijal sp. no . Holo ype male: 121 = geni alia in le la e al iew; 122 = geni alia in do sal iew;
123 = gonopods in en al iew; 124 = phallic o gan in le la e al iew; 125 = phallic o gan in do sal iew.
de ailed s udy. I could ep esen an independen
sibling.
Agape us kin isha Oláh & Vinçon, 2024
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe S ane i, Rachka, wa e all,
510m, 42.54357°N, 42.64466°E, 28.IV.2024, leg.
A. Memishishi (1 male, IZISUT).
Rema k. This species close o Agape us
sa ayensis (Sipahile , 1996) desc ibed om
Tu key was only known om he ype locali y in
he Lesse Caucasus (Adja a). I is he i s epo
om he High Caucasus.
Glossosoma capi a um Ma yno , 1913
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe S ane i, So i, sp ing, 710m,
42.58117°N, 43.28706°E, 26.IV.2024, leg. A.
Memishishi (1 male, IZISUT). Geo gia, Ime e i,
Kh e e i, sp ing, 530m, 42.3740942°N, 43.4378
85°E, 25.IV.2024, leg. A. Memishishi (1 male,
IZISUT). Geo gia, Zemo a ani, 42.23669°N,
45.00394°E, 10.X.2024, leg. A. Memishishi (1
male, IZISUT). Geo gia, Shuapkho, 42.41558°N,
44.99855°E, 7.X.2024, leg. A. Memishishi (1
male, 8 emales; IZISUT).
Hyd op ilidae S ephens, 1836
Hyd op ila a ma hai Schmid, 1959
Ma e ial examined. Geo gia, High Caucasus,
Ime e i, Lashijala, i e , 580m, 42.64795°N,
42.84235°E, 25.IV.2024, leg. A. Memishishi (3
males, 1 emale; IZISUT). Geo gia, Sameg elo -
Zemo S ane i, Dobe adzeni, sp ing, 370m,
42.59264°N, 42.32199°E, 28.IV.2024, leg. A.
Memishishi (1 male, IZISUT).
Rema ks. This species, desc ibed om I an,
was also epo ed om Aze baijan (Oláh e al.
2020). We epo i o he i s ime in Geo gia.
Hyd op ila au ica Ma yno , 1934
Ma e ial examined. Geo gia, Sameg elo -
Zemo S ane i, Dobe adzeni, sp ing, 370m,
42.59264°N, 42.32199°E, 28.IV.2024, leg. A.
Memishishi (1 male, IZISUT).
Rema k. This species desc ibed om Tu key is
epo ed o he i s ime om Geo gia.
S ac obia wimme i Malicky, 1988
Ma e ial examined. Geo gia, K emo Ka li,
Dashbashi cascades, 1410m, 41.595°N, 44.124°E,
6.VIII.2024, leg. Gilles Vinçon (3 males, OPC).
Oláh e al.: On he T ichop e a o he Caucasus III
53
Rema k. This species desc ibed om Tu key is
epo ed o he i s ime in Geo gia.
P ilocolepidae Ma yno , 1913
P ilocolepus colchicus Ma yno , 1913
Ma e ial examined. Geo gia, Kakhe ia, oad
o Omalo, small b ook wi h mosses, ibu a y o
S o i Ri e , 1230m, 42.2192°N, 45.4768°E,
22.IV.2024, leg. Gilles Vinçon (4 males, 1
emale; OPC). Geo gia, Ime e i, abo e Sai me,
b ook om 1270m, 41.883°N, 42.756°E, o i s
sp ing 1370m, 41.885°N, 42.758°E, 27.IV.2024,
leg. Gilles Vinçon (1 emale, OPC). Geo gia,
M skhe a-M iane i, abo e Pakh iji, 2 nice sp ings,
1450m, 42.4479°N, 44.7896°E, 22.IV.2024, leg.
Gilles Vinçon (2 males, 2 emales; OPC). Geo -
gia, K emo Ka li, Dashbashi cascades, 1410m,
41.595°N, 44.124°E, 24.IV.2024, leg. Gilles Vin-
çon (1 male, OPC). Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
10.VIII.–24.VIII.2018, leg. GGBC membe s (1
male, IZISUT). Geo gia, Lesse Caucasus, Ad-
ja a, Kin ishi, abo e he wa e all, malaise ap 5,
N41o44’36.3336, E42o4’54.4296, 1252m, 5.VI. -
20.V. 2018, leg. GGBC membe s e al. (2 males,
2 emales, IZISUT). Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
13.VII–27.VII. 2018, leg. GGBC membe s (2
males, 9 emales, IZISUT). Geo gia, Racha-
Lechkhumi and Lowe S ane i, Rachka, wa e all,
510m, 42.54357°N, 42.64466°E, 28.IV.2024, leg.
A. Memishishi (1 male, IZISUT). Geo gia, Sakh-
da i, sp ing, 42.82056°N, 42.79773°E, 27.IV.
2024, leg. A. Memishishi (1 male, IZISUT).
Geo gia, Gou ia, oad o Gomism a, s eep b ook,
1430m, 41.8373°N, 42.144°E, up o i s sp ing,
1460m, 41.8376°N, 42.1437°E, 28.IV.2024, leg.
Gilles Vinçon (1 male, 1 emale; OPC). Geo gia,
Ming elie, abo e Koko, nice b ookle s and
sp ings nea he wa e ca chmen , on limes one
subs a um, 310m, 42.6467°N, 42.2028°E, 3.V.
2024, leg. Gilles Vinçon (1 male, OPC). Geo gia,
Gou ia, oad o Bakhma o, nice sp ing, ‘Tskh a s-
qa o’, 1380m, 41.8911°N, 42.3718°E, 28.IV.
2024, leg. Gilles Vinçon (2 males, 1 emale;
OPC). Geo gia, Adja a, M i ala Na ional Pa c,
nice b ook, cascade and la e al sp ing, 1180m,
41.647°N, 41.883°E, 30.IV.2024, leg. Gilles
Vinçon (1 male, 1 emale; OPC). Geo gia, Adja a,
oad o Gomism a, nice b ook, 2150-2200m,
41.8098°N, 42.1827°E, 11.VIII.2024, leg. Gilles
Vinçon (2 males, 1 emale; OPC).
P ilocolepus dila a us Ma yno , 1913
Ma e ial examined. Geo gia, Adja a, Kin-
ishi, nice o en wi h mosses, 700m, 41.7385°N,
42.01°E, 29.IV.2024, leg. Gilles Vinçon (1 male,
1 emale; OPC). Geo gia, Adja a, abo e
Namonas e i, nice b ook and sp ing, 1200m,
41.568°1N, 42.0682°E, 1.V.2024, leg. Gilles
Vinçon (2 males, 3 emales; OPC). Geo gia,
Gou ia, oad o Bakhma o, nice b ook and sp ing,
1240m, 41.9°N, 42.37°E, 28.IV.2024, leg. Gilles
Vinçon (1 emale, OPC). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 4, 1264m, N41°44’38.8824,
E42°5’0.2904, 5.V–20.V.2018, leg. GGBC
membe s (1 male, IZISUT). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 4, 1264m, N41°44’38.8824, E42°5’
0.2904, 13.VII.–27.VII. 2018, leg. GGBC
membe s (1 emale, IZISUT). Geo gia, Racha-
Lechkhumi and Lowe S ane i, So i, sp ing,
710m, 42.58117°N, 43.28706°E, 26.IV.2024, leg.
A. Memishishi (3 males, IZISUT). Geo gia,
Chkhu une i, sp ing, 41.58117°N, 41.85786°E,
30.IV.2024, leg. A. Memishishi (9 males, 7
emales; IZISUT). Geo gia, Adja a, Chakha i,
sp ing, 90m, 41.8791°N, 41.91601°E, 29.IV.2024,
leg. A. Memishishi (1 male, IZISUT). Geo gia,
Ime e i, abo e Sai me, Zeka i Pass, below Didi-
maghala Pic, nice b ook wi h mosses and sp ing,
2055m, 41.85°N, 42.8°E, 9.VIII.2024, leg. Gilles
Vinçon (4 males, 2 emales; OPC).
Rhyacophilidae S ephens, 1836
Philoc ena iale ica Lepne a, 1956
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
Oláh e al.: On he T ichop e a o he Caucasus III
60
hal ; he phallo heca is wi hou p oximal en al
lip and wi h sho en oapical lip.
E ymology. Named a e he egion o he
locus ypicus.
Rema k. The e a e p onounced di e ences
be ween he geni al s uc u e o he holo ype and
pa a ypes. Pa icula ly he apicomesal p ocess
di e ged signi ican ly ha is well isible bo h in
la e al and en al iews. In he holo ype i is less
p oduced. We need o ha e o he specimens o
s udy he eal na u e o his di e gence.
Lepidos oma idae Ulme , 1903
Lepidos oma inae Ulme , 1903
Lepidos oma chaldy ense (Ma yno , 1909)
Ma e ial examined. Geo gia, K emo Ka li,
Kh ami Ri e a Dashbashi cascades, 1400m,
41.5945°N, 44.1238°E, 24.IV.2024 leg. Gilles
Vinçon (1 male, OPC). Geo gia, Sam skhe-Ja a-
khe i, Zeda Va dzia, nice b ook and sp ing,
1410m, 41.383°N, 43.258°E, 25.IV.2024, leg.
Gilles Vinçon (12 males, OPC). Geo gia, K emo
Ka li, Dashbashi cascades, 1410m, 41.595°N,
44.124°E, 24.IV.2024, leg. Gilles Vinçon (1 male,
OPC).
Lepidos oma i anicum Schmid, 1959
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe S ane i, Len ekhi, sp ing,
760m, 42.79081 42.73402, 27.IV.2024, leg. A.
Memishishi (1 male, OPC). Geo gia, Adja a,
Chkhu une i, sp ing, 700m, 41.4817°N, 41.857
86°E, 30.IV.2024, leg. A. Memishishi (1 male,
OPC). Geo gia, Racha-Lechkhumi and Lowe
S ane i, U a i, sp ing, 1000m, 42.65771°N,
43.29736°E, 26.IV.2024, leg. A. Memishishi (1
male, OPC). Geo gia, Lesse Caucasus, Adja a,
Kin ishi, abo e he wa e all, malaise ap 4,
1264m, N41°44’38.8824, E42°5’0.2904, 10.VII. -
27.VII.2018, leg. GGBC membe s e al. (1 male,
IZISUT). Geo gia, M skhe a-M iane i, S. Tsdo,
nice b ook, 1700m, 42.6824°N, 44.6328°E,
2.VIII.2024, leg. Gilles Vinçon (11 males, 8
emales; OPC). Geo gia, Khakhma i, 42.52761°N,
45.01662°E, 8.X.2024, leg. A. Memishishi (1
male, OPC).
Rema ks. This species, desc ibed om I an,
was la e collec ed in he Geo gian High Caucasus
(Oláh & Vinçon 2024). He e we epo i o he
i s ime in he Geo gian Lesse Caucasus.
Lepidos oma longiplica um Ma yno , 1913
Ma e ial examined. Geo gia, Kakhe ia, oad o
Omalo, small b ook wi h mosses, ibu a y o
S o i Ri e , 1230m, 42.2192°N, 45.4768°E, 22.
IV.2024, leg. Gilles Vinçon (1 male, OPC).
Geo gia, Kakhe ia, abo e Lagodekhi, Sh omis-
Khe i alley, big o en , below I on B idge,
740m, 41.8585°N, 46.3076°E, 20.IV.2024, leg.
Gilles Vinçon (2 males, OPC).
Theliopsychinae Wea e , 1992
Ma ynomyia bakhma a Oláh & Vinçon,
sp. no .
(Figu es 140–143, Map 5, Pho os 20–21)
Ma e ial examined. Holo ype: Geo gia, Gou-
ia, abo e Bakhma o, b ook and sp ing, 1980m,
41.8386°N, 42.331°E, 10.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Diagnosis. Mos close o Ma ynomyia zazai
Oláh & Vinçon, 2024 bu easily dis inguished
om all he known species o he complex by he
used ce ci and pa ap oc complex ha ing simple
od shape supplied wi h la e al ow o ine se ae,
by e y long and e y slende digi i o m middle
p ocess on gonopod as well as by he s aigh e y
long do sal and s aigh e y sho en al spines
in he endo heca o phallic o gan.
Desc ip ion. Small, da k b own animal wi h
o ewing leng h o 6 mm. Apicomesal lobe on
s e ni e VII na ow elonga ed.
Male geni alia. Segmen IX has sho do sum,
en um longe and an e io ma gin con ex
Oláh e al.: On he T ichop e a o he Caucasus III
61
Figu es 140–143. Ma ynomyia bakhma a sp. no . Holo ype male: 140 = geni alia in le la e al iew; 141 = geni alia
in do sal iew; 142 = le gonopod in en al iew; 143 = phallic o gan in le la e al iew.
Map 5. Ma ynomyia genus (8 species): M. ayde ensis Sipahile , 1989, M. bakhma a sp. no ., M. kin isha sp. no .,
M. ma yno i Sipahile , 1995, M. omala sp. no ., M. s ane ia Oláh & Vinçon, 2024,
M. ipa i a (Ma yno , 1913), M. zazai Oláh & Vinçon, 2024.
ounded iangula . Segmen X memb anous wi h
upwa d u ning apex. The used complex o ce ci
and pa ap oc is no den a e, a med only wi h
sho se ae; pa icula ly concen a ed along i s
la e al ma gin; in la e al iew b oadening basad.
The ipa i e gonopods composed o o oid highly
se ose do sal, less se osed long and slende
digi i o m middle and se ose lowe lobes dis-
ce nible in la e al iew; he digi i o m middle
p ocess se osed only on i s apical egion wi h a
ew long se ae; he highly se osed do sal o oid
p ocess ep esen s p obably he ha pago, he
second segmen o he gonopods bu used wi h
a icula ion on he e y basal egion o he
coxopodi e; he apical egion o he lowe lobes
cu ing mesad ending in a i o m head in en al
iew, basal pa b oadening. Phallic o gan wi h
wo pai s o pa ame es embedded inside he
phallo heca o endo heca; do sal spine e y long,
en al spine sho , bo h s aigh .
Oláh e al.: On he T ichop e a o he Caucasus III
62
Figu es 144–147. Ma ynomyia kin isha sp. no . Holo ype male: 144 = geni alia in le la e al iew; 145 = geni alia
in do sal iew; 146 = le gonopod in en al iew; 147 = phallic o gan in le la e al iew.
Figu es 148–150. Ma ynomyia kin isha sp. no . Allo ype emale: 148 = geni alia in le la e al iew; 149 = geni alia
in do sal iew; 150 = geni alia in en al iew.
E ymology. Coined om he name o holo-
ype’s locali y, a noun in apposi ion.
Ma ynomyia kin isha Oláh & Memishishi,
sp. no .
(Figu es 144–150, Map 5, Pho os 7–8)
Ma e ial examined. Holo ype: Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 11, 1264m, N41°44’38.8824”, E42°
5’0.2904”, 24.VII–7.IX.2018, leg. GGBC mem-
be s (1 male, OPC). Allo ype: same as holo ype (1
emale, OPC). Pa a ypes: same as holo ype (2
males, IZISUT; 4 males, OPC).
Diagnosis. Fused complex o ce ci and pa a-
p oc is ba e spine-like, no den a e, no ba e od-
like and no sho pla e-like. This new species is a
sibling o Ma ynomyia ayde ensis Sipahile ,
1989, bu di e s by he highly di e ged spine
pa e n o he used ce ci and pa ap oc as
desc ibed below. T ipa i e gonopod is also di-
e ged as isible bo h in la e al and en al iews;
middle digi i o m ba e p ocess almos as long as
he o he s, no much longe ; lowe mesad cu ing
p ocess di e en ly shaped; b oad based, no
slende along i s en i e leng h. The e a e di e -
gences also in emale geni alia; used e gi e IX
and X longe as isible in la e al iew; i s lobes
Oláh e al.: On he T ichop e a o he Caucasus III
63
wi h ounded apices, no poin ed and no mesad
u ning, he la e al shape o s e ni e X also di e s.
Desc ip ion. Small, da k b own animal wi h
o ewing leng h o 5 mm, bo h holo ype and
allo ype. Apicomesal lobe on s e ni e VII sho
ounded iangula .
Male geni alia. Segmen IX has do sum,
sho e han en um; an e io ma gin con ex
ounded iangula ; pos e io ma gin s aigh
e ical. Segmen X memb anous, jus disce nible
wi h indis inc apex. The used complex o ce ci
and pa ap oc is no den a e as M. s ane ia and M.
ipa i a, no ba e od-like as M. omala and M.
zazai and no sho pla e-like as M. ma yno i; i
is spine-like as M. ayde ensis; spine-like used
ce ci and pa ap oc has wo basomesal and wo
basola e al small spines, he e is no spines along
he s alk and he e is no addi ional apical spine.
The ipa i e gonopods composed o o oid highly
se ose do sal, less se ose digi i o m middle and
se ose lowe lobes disce nible in la e al iew; he
digi i o m middle p ocess se ose only on i s apical
egion wi h a ew long se ae; he highly se osed
do sal mo e b oad p ocess ep esen s p obably he
ha pago, he second segmen o he gonopods bu
used wi h a icula ion on he e y basal egion o
he coxopodi e; he apical digi i o m egion o he
lowe lobes sligh ly cu ing mesad wi h ape ing
poin ed head in en al iew, middle and basal
pa b oadening. Phallic o gan wi h wo pai s o
pa ame es embedded inside endo heca dis al pai
s aigh , basal cu ing.
Female geni alia. Basic a chi ec u e o emale
geni alia simila o Ma ynomyia ayde ensis Sipa-
hile , 1989, bu he memb anous p o uded s uc-
u e isible in uded; his less isible, jus disce -
nible s uc u e is a he a pa o he aginal scle-
i e complex somehow in uded-p o uded du ing
copula ion; i s shape is a he lexible ha ing no
s able diagnos ic alue. In he in uded s a e o he
memb anous s uc u e he e is a memb anous
connec ion be ween e gi e VIII and he used
e gi IX and X. I seems his memb anous con-
nec ion p essed e y sho when he memb anous
s uc u e is p o uded possibly as a esul o
copula ion. Scle o ized used e gi IX and X
sho , bu longe compa ed o M. ayde ensis. In
do sal iew bilobed, apical head ounded. S e ni e
X egula ec angula in en al iew.
E ymology. Coined om he name o holo-
ype’s locali y, a noun in apposi ion.
Ma ynomyia omala Oláh & Vinçon, sp. no .
(Figu es 151–157, Map 5, Pho os 10–11)
Ma e ial examined. Holo ype: Geo gia, Kakh-
e ia, oad o Omalo, nice b ook and la e al sp ing,
ibu a y o S o i Ri e , 650m, 42.177°N, 45.431°
E, 22.IV.2024, leg. Gilles Vinçon (1 male, OPC).
Allo ype: same as holo ype (1 emale, OPC).
Pa a ypes: same as holo ype (4 males, 1 emale;
OPC). Geo gia, Kakhe i, oad o Omalo, nice
b ook unde cascade, 1500m, 42.2373°N,
45.4921°E, 5.VIII.2024, leg. Gilles Vinçon (4
males, OPC).
Diagnosis. Fused complex o ce ci and pa-
ap oc is ba e, no den a e. This new species is a
sibling o Ma ynomyia zazai Oláh & Vinçon,
2024, bu di e s by he s aigh , no S-shaped
do sal p o ile o he used ce ci and pa ap oc as
well as by he clea ly excised, no ounded head
o used ce ci and pa ap oc . The beak o a i o m
head o he en al a m o coxopodi e is long, no
sho in en al iew.
Desc ip ion. Small, da k b own animal wi h
o ewing leng h o 5 mm. Apicomesal lobe on
s e ni e VII sho ounded iangula .
Male geni alia. Segmen IX has do sum, and
en um o almos equal leng h; an e io ma gin
con ex ounded iangula ; pos e io ma gin
s aigh e ical. Segmen X memb anous wi h
indis inc apex. The used complex o ce ci and
pa ap oc is no den a e, a med only wi h a ew
long se ae; wo a apex and h ee- ou a he basal
egion; in la e al iew he basal pa is b oad; i s
apex sligh ly excised as isible bo h in la e al and
do sal iews. The ipa i e gonopods composed
o o oid highly se ose do sal, less se ose digi i-
o m middle and se ose lowe lobes disce nible in
la e al iew; he digi i o m middle p ocess se ose
only on i s apical egion wi h a ew long se ae; he
highly se osed do sal o oid p ocess ep esen s
Oláh e al.: On he T ichop e a o he Caucasus III
64
Figu es 151–154. Ma ynomyia omala sp. no . Holo ype male: 151 = geni alia in le la e al iew; 152 = geni alia
in do sal iew; 153 = le gonopod in en al iew; 154 = phallic o gan in le la e al iew.
Figu es 155–157. Ma ynomyia omala sp. no . Allo ype emale: 155 = geni alia in le la e al iew; 156 = geni alia
in do sal iew; 157 = geni alia in en al iew.
p obably he ha pago, he second segmen o he
gonopods bu used wi h a icula ion on he e y
basal egion o he coxopodi e; he apical digi i-
o m egion o he lowe lobes sligh ly cu ing
mesad wi h ape ing poin ed head in en al iew,
middle and basal pa b oadening. Phallic o gan
wi h wo pai s o pa ame es embedded inside
endo heca dis al pai s aigh , basal cu ing.
Female geni alia. Basic a chi ec u e o emale
geni alia simila o Ma ynomyia zazai Oláh &
Vinçon, 2024, bu he la e al shape o he highly
scale o ized s uc u e o aginal scle i e as well as
bo h he la e al and pa icula ly en al shape o
s e ni e X di e en .
E ymology. Coined om he name o holo-
ype’s locali y, a noun in apposi ion.
Ma ynomyia s ane ia Oláh & Vinçon, 2024
(Map 5, Pho os 12–13)
Ma e ial examined. Geo gia, Ming elia, abo e
Koko, nice b ookle and sp ings nea he wa e
ca chmen , on limes one subs a um, 330m,
42.648°N, 42.2033°E, 3.V.2024, leg. Gilles Vin-
çon (1 male, OPC).
Ma ynomyia ipa i a (Ma yno , 1913)
(Map 5)
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe S ane i, Rachka, wa e all,
42.54357°N, 42.64466°E, 510m, 28.IV.2024, leg.
A. Memishishi (1 male, IZISUT).
Oláh e al.: On he T ichop e a o he Caucasus III
65
Figu es 158–160. Ma ynomyia zazai Oláh & Vinçon, 2024. Female: 158 = geni alia in le la e al iew;
159 = geni alia in do sal iew; 160 = geni alia in en al iew.
Ma ynomyia zazai Oláh & Vinçon, 2024
(Figu es 158–160, Map 5)
Ma e ial examined. Geo gia, Kakhe ie, oad
o Omalo, abo e Lechu i, nice b ook, 650m,
42.1643°N, 45.419°E, 24.V.2023, leg. Gilles Vin-
çon (in copula: 1 male, 1 emale; OPC).
Female desc ip ion. We ha e a g ea luck
ha ing ou emale in copula wi h a pa a ype male;
he e o e we ha e d awn he copula posi ion
wi hou de aching hem. Simila ly o he com-
pa ison o M. ayde ensis Sipahile and M. kin-
isha sp. no . i seems du ing copula ion he long
memb anous egion be ween e gi e VIII and
used e gi e IX and X p essed o almos disap-
pea ance. Female o M. zazai is simila o M.
omala, al hough in copula posi ion compa ison o
M. zazai and M. omala no easy. Fo una ely e en
in copula posi ion di e ences in he la e al and
en al shape o bilobed ending o used e gi e IX
and X, s e ni e X as well as he la e al p o ile o
he highly scle o ized aginals scle i e complex is
well isible and compa able on he d awings.
Apa aniidae Walleng en, 1886
Apa ania sub ilis Ma yno , 1909
Ma e ial examined. Geo gia, K emo Ka li,
Kh ami Ri e a Dashbashi cascades, 1400m,
41.5945°N, 44.1238°E, 24.IV.2024 leg. Gilles
Vinçon (12 males, 3 emales; OPC). Geo gia,
M skhe a-M iane i, abo e Dzeg i, sp ing, 1420m,
41.782°N, 44.512°E, 24.IV.2024, leg. Gilles Vin-
çon (2 males, OPC). Geo gia, M skhe a-M iane i,
abo e K emo Mle a, sp ings and cascades,
1560m, 42.4364°N, 44.49 46°E, 23. IV.2024, leg.
Gilles Vinçon (6 males, 3 emales; OPC). Geo -
gia, M skhe a-M iane i, abo e K emo Mle a,
sp ing and b ook, 1480m, 42.432°N, 44.50 75°E,
23.IV.2024, leg. Gilles Vinçon (1 male, OPC).
Geo gia, K emo Ka li, Dashbashi cascades,
1410m, 41.595°N, 44.124°E, 24.IV.2024, leg.
Gilles Vinçon (1 male, OPC). Geo gia, Tsana,
sp ing, 42.85197°N, 43.16296°E, 1510m, 27.IV.
2024, leg. A. Memishishi (4 males, IZISUT).
Geo gia, Ime e i, Lashijala, i e , 42.64 795°N,
42.84235°E, 26.IV.2024, leg. A. Memishishi (4
males, IZISUT). Geo gia, High S ane ie, abo e
Mes ia, b ook and sp ings abo e he ish a m,
1550m, 43.0477°N, 42.80 91°E, 3.V.2024, leg.
Gilles Vinçon (4 males, 3 emales; OPC). Geo -
gia, Kakhe ia, abo e Lagodekhi, Sh omis-Khe i
alley, b ook, 580m, 41.84056° N, 46.2844°E,
20.IV.2024, leg. Gilles Vinçon (2 males, 1 e-
male; OPC). Geo gia, Racha-Lechkhumi and
Lowe -S ane ie, abo e Kulbaki, nice b ook, 980
m, 42.6385°N, 42.611°E, 2.V. 2024, leg. Gilles
Vinçon (2 males, OPC). Geo gia, High S ane i,
nea Zaga i Pass, la e al sp ing, 2320m, 42.9
29°N, 43.133°E, 14.VIII.2024, leg. Gilles Vinçon
(4 males, 11 emales; OPC). Geo gia, M skhe a-
M iane i, Gudani 42.53277°N, 44.95968°E, 8.X.
2024, leg. A. Memishishi, (1 male, OPC).
Oláh e al.: On he T ichop e a o he Caucasus III
66
Limnephilidae Kolena i, 1848
D usinae Banks, 1916
D usus akhalchala Oláh & Vinçon, sp. no .
(Figu es 161–168, Pho o 6)
Ma e ial examined. Holo ype: Geo gia, Ra-
cha-Lechkumi and Lowe -S ane i, Akhalchala,
b ook and sp ing, 2170m, 42.6947°N, 42.6463°E,
13.VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Allo ype: same as holo ype (1 emale, OPC).
Pa a ype: same as holo ype (1 emale, OPC).
Diagnosis. Ha ing gigan ic pa ame es, dis ally
dila ed aedeagus and e ical plan o gonopods
his in e es ing species belongs o he small D u-
sus bud zi species g oup. D usus akhalchala Oláh
& Vinçon, sp. no . is he i h membe o his
highly modi ied apomo phic species g oup o he
D usus genus. This unique g oup ep esen s an
independen lineage in he genus wi h g ea
cha ac e s a e ans o ma ions ini ia ed by d as ic
s ochas ic impac s and c ea ed du ing in eg a i e
o ganisa ion. The enla gemen o pa ame e sha
is coupled wi h he shi o spine o ganising cen e
o he pa ame e e minal. The basic a chi ec u al
plan o he geni alia o D. akhalchala is iden ical
wi h ha o D. bud zi (Ulme , 1913), D. janjulae
Oláh, 2020, D. kheled Oláh & Vinçon, sp. no .,
D. maculosus Malicky & Oláh, 1979, bu he
pe iphallic o gans o ce ci, pa ap oc , gonopod
di e ged om ha o D. bud zi, D. macula us and
D. janjulae, bu closely ela ed o D. kheled.
D usus akhalchala sp. no . di e s om D. kheled
sp. no . by he less de eloped sweeping s uc u e
on he pa ame e head. I s unc ional impo ance is
discussed in he diagnosis o D. kheled.
Desc ip ion. La ge-sized animal wi h yellow-
ish, s amineous body and wing colou . I s o e-
wing leng h is 14 mm.
Male geni alia. Segmen IX sho and high
ounded con ex an e ad wi h sudden na owing
do sad esul ed in a e y sho band-like do sum,
bu longe en um. Ce ci downwa d di ec ed
o oid se ose la ge pla e basally connec ed o he
pa ap oc . The do sal b anch o he pa ap oc
composed o an apical pai o hea ily scle o ized
den a e black ci cula lobes a ching la e ad isible
in caudal iew and a se ose do sal egion ha ing
pu a i e ce cal o igin. Gonopods almos e ical
di ec ed wi h mo e de eloped pos e ad di ec ed
do sal and less p oduced en al egions. Phallic
o gan has a obus aedeagus wi h capi a e head
and a pai o e ically la enla ged pa ame es
a med wi h a small g oup o ew spines on i s
do soapical na owing ip and co e ed on he
dis al hal wi h igh ly packed elonga ed and
decumben ine scales o sha p ip.
Female geni alia. Te gi e o used segmen s
IX and X o ming a sho ube closed en ally,
do sum wi h la ge ounded mesal excision; la e al
lobes iangula in do sal iew; he la e al se ose
lobe o s e ni e IX elonga ed wi h ounded apical
ma gin. Sup ageni al pla e (uppe aginal lip) o
segmen X well-de eloped and quad angula bo h
in la e al and en al iews. Median lobe o he
ul a scale (lowe aginal lip) p esen as a small
digi i o m slende p ocess compa ed o he much
p oduced la e al lobes.
E ymology. Coined om he name o holo-
ype’s locali y, a noun in apposi ion.
D usus alapos Oláh, 2020
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
13.VII. –27.VII. 2018, leg. GGBC membe s (1
male, IZISUT). Geo gia, Adja a, oad o Gomis-
m a, b ook, 2150–2200m, 41.8098°N, 42.1827°E,
11.VIII.2024, leg. Gilles Vinçon (3 males, 3
emales; OPC). Geo gia, Racha-Lechkhumi and
Lowe -S ane i, Akhalchala, b ook and sp ing,
2120m, 42.6928°N, 42.6447°E, 13.VIII.2024, leg.
Gilles Vinçon (4 males, OPC). Geo gia, Adja a,
abo e Gomism a, b ook, 2120–2150m, 41.8108°
N, 42.1821°E, 11.VIII.2024, leg. Gilles Vinçon (4
males, OPC).
D usus e des Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, M skhe a-M ia-
ne i, abo e K emo Mle a, nice sp ings and cas
Oláh e al.: On he T ichop e a o he Caucasus III
67
Figu es 161–165. D usus akhalchala sp. no . Holo ype male: 161 = geni alia in le la e al iew; 162 = pa ap oc in do sal iew;
163 = pa ap oc in do socaudal iew; 164 = phallic o gan in le la e al iew; 165 = pa ame es in en al iew.
Figu es 166–168. D usus akhalchala sp. no . Allo ype emale: 166 = geni alia in le la e al iew; 167 = geni alia
in do sal iew; 168 = geni alia in en al iew.
cades, 1560m, 42.4364°N, 44.4946°E, 23.IV.
2024, leg. Gilles Vinçon (6 males, OPC). Geo gia,
Gou ia, oad o Gomism a, sp ings, 1900m,
41.8317°N, 42.1556°E, 11.VIII.2024, leg. Gilles
Vinçon (2 males, OPC). Geo gia, Ime e i, abo e
Sai me, Zeka i Pass, below Didimaghala Pic, nice
b ook wi h mosses and sp ing, 2055m, 41.85°N,
42.8°E, 9.VIII.2024, leg. Gilles Vinçon (4 males,
3 emales; OPC). Geo gia, Gou ia, abo e Bakh-
ma o, b ook and sp ing, 1980m, 41.8386°N,
42.331°E, 10.VIII.2024, leg. Gilles Vinçon (5
males, 4 emales; OPC).
D usus gudau Oláh & Vinçon, sp. no .
(Figu es 169–171, Pho os 22–23)
Ma e ial examined. Holo ype: Geo gia,
M skhe a-M iane i, be ween S ephan sminda and
Gudau i, sou h Almasiani, nice b ook, 2350m,
42.5342°N, 44.4978°E, 2.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Diagnosis. Ha ing in la ed pa ame e e minal
sec ion wi h pili e ous- omen ose-pubescen pa a-
Oláh e al.: On he T ichop e a o he Caucasus III
68
Figu es 169–171. D usus gudau sp. no . Holo ype male:
169 = geni alia in le la e al iew; 170 = ce cus and do sal
a m o pa ap oc in caudal iew; 171 = phallic o gan
in la e al iew.
me e su ace and ecumben p ima y and second-
da y pa ame e spines, i belongs o he D usus
caucasicus species g oup. The pla e-shaped se ose
ce cus is used o a digi i o m se aless p ocess o
pa ap oc o igin, a cha ac e combina ion o he
D usus caucasicus species complex (Oláh e al.
2017). Mos close o D usus caucasicus Ulme ,
1907, bu di e s by ha ing pa ame e wi h mo e
in la ed head and wi h p onounced subapical
cons ic ion. The mesal a m o he bilobed used
complex ce cal s uc u e ha is he se aless used
do sal a m o he pa ap oc is e en mo e obus ,
especially in la e al iew. The neu al pe iphallic
o gan o he gonopod is sligh ly ape ing, no
ounded unca e.
Desc ip ion. Ligh b own animal wi h o e-
wing leng h o 11 mm. Segmen IX sho , e y
sho do sad and longe en ad; an e io ain-
gula lobe shi ed en ad. Basal pa o pa ap oc
wi h closed apezoid ame. Highly modi ied
do sal a m o pa ap oc used o ce ci and
de eloped pa icula ly obus in la e al iew.
Gonopods sho and ounded. Pa ame es supplied
wi h h ee ecumben spines and dila ed pili e ous
head.
E ymology. Coined om he name o he ype
locali y.
D usus kheled Oláh & Vinçon, sp. no .
(Figu es 172–174, Pho os 24–25)
Ma e ial examined. Holo ype: Geo gia, Ra-
cha-Lechkhumi and Lowe -S ane i, abo e Khe-
ledi, sp ing, 1180m, 42.8037°N, 42.6335°E, 14.
VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Diagnosis. Ha ing gigan ic pa ame es, dis ally
dila ed aedeagus and e ical plan o gonopods
his in e es ing species belongs o he small D u-
sus bud zi species g oup. This unique g oup ep-
esen s an independen lineage in he genus wi h
g ea cha ac e s a e ans o ma ions ini ia ed by
d as ic s ochas ic impac s and c ea ed du ing
in eg a i e o ganisa ion. The enla gemen o pa a-
me e sha is coupled wi h he shi o spine
o ganising cen e o he pa ame e e minal. The
basic a chi ec u al plan o he geni alia is simila
o D. maculosus Malicky & Oláh, 1979, bu he
pe iphallic o gans o ce ci, pa ap oc and gonopod
di e ged and he pa ame e is highly modi ied
wi h i s scale co e ed apical su ace and wi h i s
mesal idge. D usus kheled is a sibling, a close
ela i e o D. akhalchala bu di e s by he highly
modi ied sweeping e minal, he i illa ing powe
o he pa ame e.
Apa om he small di e gences in he pe i-
phallic o gan be ween D. akhalchala and D.
kheled he e is a signi ican modi ica ion in he
sweeping e minalia o he pa ame e. Compa ed
o D. akhalchala, D. kheled has he e minal
sweeping spine s uc u e highly modi ied and
di e si ied by p oducing mo e, o e i een hick
and long spines on pa ame e head o en o ce i s
i illa ing sweeping unc ion in he c yp ic emale
choise p ocesses. Bi d males a e ecognised and
accep ed by hei emales by ecognising hem
wi h hei song pa e n and plumage colou o
ep oduc i e isola ion ba ie s, in p ezygo ic
ma ing p ocesses.
D usus kheled male is ecognised and accep ed
by emale by i s sweeping powe du ing copu-
la ion. These adap i e di e gences seem sub le o
human capaci y, hey a e in eg a i ely o ganised
by la ge ne wo ks o DNA sequences. E en sub le
Oláh e al.: On he T ichop e a o he Caucasus III
69
Figu es 172–174. D usus kheled sp. no . Holo ype male: 172 = geni alia in le la e al iew; 173 = le pa ame e
in le la e al iew; 174 = pa ame es in en al iew.
s uc u al di e gences a e p oduced by a e y
complex combina ion o la ge numbe o quan-
i a i e ai loci; among he de ec ed 8000 genes
2261 genes we e di e en ially exp essed in small
pos e io lobe modi ica ion o geni al imaginal
disc be ween wo di e ging D osophila species
(Masly el al. 2011).
These mino shape modi ica ions es ablishing
ep oduc i e isola ion o incipien sibling species
a e c ea ed in complex in e ac i e gene ic p o-
cesses by in eg a i e o ganisa ion. In mo pholo-
gical alpha axonomy such mino shape di e gen-
ces o specia ion exp essed by specia ion magic
ai s need s abili y es by isual ai ma ix o by
simple o s e eo quan i a i e mo phome y. In
mo phological alpha axonomy o ine phenomics
he ep oduc i e isola ion o incipien sibling
species in i s ini ial s age is demons a ed by hese
mino , sub le bu s able ai di e gences. This
means ha he s abili y o he sweeping s uc u e
on he pa ame e head o D usus akhalchala needs
o be es ed bu , we need o collec mo e
specimens o his s abili y es .
Desc ip ion Medium-sized animal wi h yello-
wish, s amineous body and wing colou ; o e-
wing wi h indis inc ligh spo s. I s o ewing
leng h is 11 mm. Segmen IX sho and high
pa allel-sided wi h sudden na owing do sad
esul ed in a e y sho band-like do sum; en-
um longe . Ce ci downwa d di ec ed wi h o oid
se ose a he la ge pla e basally connec ed o he
pa ap oc . The do sal b anch o he pa ap oc
composed o an apical pai o hea ily scle o ized
black la e ad a ching lobes isible in caudal iew
and a se ose do sal egion ha ing pu a i e ce cal
o igin. Gonopods almos e ical di ec ed wi h
mo e de eloped do sal and less p oduced en al
egions. Phallic o gan has a obus aedeagus wi h
dila ed head and a pai o e ically la enla ged
pa ame es a med wi h o e i een long, hick
spines on i s do soapical hal o he head and
co e ed on he dis al egion wi h ine scaly and
sho spines; he e is a mesal idge on i s apical
mesal su ace.
E ymology. Coined om he name o he ype
locali y, a noun in apposi ion.
D usus megno Oláh & Vinçon, 2020
Ma e ial examined. Geo gia, G ea e Caucasus
Moun ains, M skhe a-M iane i egion, Pasanau i,
42.353°N, 44.709°E, 1100m, 7.X.2024, leg. Ja i
Junnilainen (1 male, OPC).
Oláh e al.: On he T ichop e a o he Caucasus III
76
Map 7. Rizeiella genus (12 species): R. ana olica Sipahile , 1986, R. bakhma a sp. no ., R. bayae Vinçon & Oláh, 2020, R.
camiliensis, Sipahile , 1999, R. e eda Oláh & Vinçon, 2020, R. keda Oláh & Vinçon, 2024, R. od a Oláh & Vinçon, 2020, R.
oldala Oláh & Ko ács, 2020, R. sashuala Oláh & Vinçon, 2024, R. a ola Oláh & Vinçon, 2020, R. be ia Oláh & Vinçon, 2024,
R. a jana Oláh & Vinçon, 2024.
(1 male, OPC). Geo gia, M skhe a-M iane i,
Zemo A ani, 1290m, 42.24664 45.01273, 10.X.
2024, leg. A. Memishishi, (1 male, OPC). Geo -
gia, M skhe a-M iane i, Be isch dili, 1160m,
42.41133°N, 44.92827°E, 10.X.2024, leg. A.
Memishishi, (1 male, OPC).
Rizeiella bakhma a Oláh & Vinçon sp. no .
(Figu es 191–197, Map 7, Pho os 20–21)
Ma e ial examined. Holo ype: Geo gia, Gou-
ia, abo e Bakhma o, b ook and sp ing, 1980m,
41.8386°N, 42.331°E, 10.VIII.2024, leg. Gilles
Vinçon (1 male, OPC). Allo ype: same as holo-
ype (1 emale, OPC). Pa a ype: same as holo ype
(1 emale, OPC).
Diagnosis. This new species has esemblance
o Rizeiella sashuala Oláh & Vinçon, 2024 and R.
a jana Oláh & Vinçon, 2024 bu di e s om
bo h by ha ing and no lacking pa ame e on he
phallic o gan, al hough he de elopmen o his
es igial s uc u e could be no s able. Also di e s
om bo h by he semici cula ce ci, he abb e-
ia ed pa ap oc and mo e elonga ed apical egion
o gonopods. I di e s also by bo h he la e al and
do sal pa e ns o he head o he s ongly scle o-
ized pai o he aedeagal apicodo sal p ocesses.
The en al lobe on he anal ube o he emale is
Oláh e al.: On he T ichop e a o he Caucasus III
77
sho e han he do sal lobe in la e al iew and he
do sal p o ile o segmen IX has wide mesal
excision.
Desc ip ion. Male and emale (in alcohol).
Yellowish pale b ownish medium-sized species
wi h pale es aceous body appendages and wi h
yellowish- es aceous wings. Fo ewing wi h less
ounded apices, e y long e ec spine-like se ae
p esen on he memb ane and on he eins; a ound
e e y second se ae on he eins s onge ; he e is
some ew pubescence p esen be ween cos al and
subcos al eins. Tibial spu numbe is 033 a male
and 133 a emale. Fo ewing leng h is 8 mm bo h
a male and emale wi h a ia ion.
Male geni alia. P o ube ance o spinulose a ea
o es i u al noncellula mic o ichia on e gi e
VIII p esen and enla ged. Segmen IX longe
middle; e y sho s ap o b idle-like do sally.
Segmen X pa ly used o basal egion o ce ci
o ming oge he a sho do sal conca i y. Ce ci
hea ily and en i ely scle o ized, semici cula in
la e al and almos ci cula in caudal iews. Pa a-
p oc sho , b oad inge -like in la e al iew and
comple ely used o he equally s ongly scle o-
ized ce ci o ming oge he a ci cula cup-like
s uc u e wi h he caudad di ec ed p ocess o
es igial en al b anch o he pa ap oc . Gonopod
sligh ly conca e apicad in la e al na owing api-
codo sad. Phallic o gan is o Rizeiella ype wi h a
single s ongly scle o ized pai o aedeagal apico-
do sal p ocesses wi h pegged apex and speci ic
la e al and en al iews; pai o pa ap oc p esen
as a single slende digi i o m p ocess wi h an
apical s ong spine.
Female geni alia. The e is a closed “anal ube”
o med by he complex o he a iously used
e gi e IX and segmen X; i s apical egion bi-
lobous bo h in la e al and do sal iews; he do sal
lobe pin-poin ed in do sal and la e al iews;
en al lobe o he ube is sligh ly sho e han he
do sal in la e al iew; he do sum deeply and
oundly excised and en um no excised apico-
mesad. Se ose s e ni e IX connec ed by less
glab ous mesal pla e; his en al su ace, he
sup ageni al pla e unc ions like he uppe aginal
lip. The lowe aginal lip, he ul a scale is
memb anous badly isible. Vaginal scle i e com-
plex isible subquad angula wi h apicola e al
lobes in do sal iew.
E ymology. Name was coined om he name
o he ype locali y, a noun in apposi ion.
Rizeiella keda Oláh & Vinçon, 2024
(Map 7)
Ma e ial examined. Geo gia, Adja a, abo e
Namonas e i, b ookle and sp ing, 1720m, 41.55
28°N, 42.0892°E, 12.VIII.2024, leg. Gilles Vin-
çon (1 male, 1 emale; OPC). Geo gia, Gou ia,
oad o Gomism a, sp ings, 1900m, 41.8317°N,
42.1556°E, 11.VIII.2024, leg. Gilles Vinçon (1
male, OPC). Geo gia, Adja a, abo e Namona-
s e i, nice b ook and sp ing, 1710m, 41.555°N,
42.0853°E, 12.VIII.2024, leg. Gilles Vinçon (1
male, OPC). Geo gia, Lesse Caucasus, Adja a,
Kin ishi, abo e he wa e all, malaise ap 11,
1637m, 41.74769°N, 42.0951°E, 21.IX –5.X.
2018, leg. GGBC membe s (3 males, OPC).
Geo gia, Lesse Caucasus, Adja a, Kin ishi, a-
bo e he wa e all, malaise ap 11, 1637m,
41.74769°N, 42.0951°E, 24.VIII.–7.IX.2018, leg.
GGBC membe s (3 males, OPC).
Rema k. This species was only known om
he ype locali y, in sou he n Adja a close o he
Tu kish bo de . I widely ex ends in he Lesse
Caucasus, up o Gou ia.
S enophylacini Schmid, 1955
Halesus caucasicus Oláh, 1985
Ma e ial examined. Geo gia, M skhe a-M ia-
ne i, Khakhma i, 1990m, 42.52761°N, 45.016
62°E, 8.X.2024, leg. A. Memishishi (3 males,
IZISUT). Geo gia, Sam skhe-Ja akhe i, Apnia,
Uppe Va dsia, Gues house Vania, 1290m,
41.374°N, 43.262°E, 5-7.X.2024 leg. J. Junni-
lainen (5 males, OPC).
Rema k. In ou o iginal desc ip ion (Oláh
1985), we ha e ecognised and d awn he wo
addi ional iny spines a he end o he pa ame e
o he holo ype. Un o una ely hese addi ional
Oláh e al.: On he T ichop e a o he Caucasus III
78
Figu es 191–194. Rizeiella bakhma a sp. no . Holo ype male: 191 = geni alia in le la e al iew; 192 = le gonopod in en al
iew; 193 = phallic o gan in le la e al iew; 194 = apex o phallic o gan in en al iew.
Figu es 195–197. Rizeiella bakhma a sp. no . Allo ype emale: 195 = geni alia in le la e al iew; 196 = geni alia
in do sal iew; 197 = geni alia in en al iew.
iny spines we e no disce nible du ing eexa-
mina ion and ed awing (Oláh e al. 2020). They
mo ed in he igh side co e o he pa ame e
e minal. Upon examining mo e specimens, we
ealised ha he posi ion o he addi ional spines
was uns able and ha he numbe o addi ional
spines could be educed o one.
Po amophylax a meniacus Mey, 1979
Ma e ial examined. Geo gia, K emo Ka li,
Kh ami Ri e a Dashbashi cascades, 1400m,
41.5945°N, 44.1238°E, 24.IV.2024 leg. Gilles
Vinçon (14 males, 2 emales; OPC). Geo gia,
Lesse Caucasus, Adja a, Kin ishi, abo e he
wa e all, malaise ap 5, 1252m, N41°44’
36.3336, E42°4’54.4296, 19.V.–1.VI.2018, leg.
GGBC membe s (2 males, OPC). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 4, 1264m, N41°44’38.8824, E42°
5’0.2904, 1.VI.–15.VI.2018, leg. GGBC membe s
(2 males, IZISUT). Geo gia, Lesse Caucasus,
Adja a, Kin ishi, subalpine meadow, malaise ap
18, 2450m, N41°45’41.8, E42°06’56.9, 29.VI.–
Oláh e al.: On he T ichop e a o he Caucasus III
79
13.VII.2018, leg. GGBC membe s (1 male,
IZISUT).
Po amophylax la ipennis (Cu is, 1934)
Ma e ial examined. Geo gia, G ea e Cau-
casus Moun ains, M skhe a-M iane i egion, Pasa-
nau i, 42.353°N, 44.709°E, 1100m, 7.X2024, leg.
Ja i Junnilainen (5 males, OPC).
S enophylax bo os Oláh & Memishishi, sp. no .
(Figu es 198–201)
Ma e ial examined. Holo ype: Geo gia,
M skhe a-M iane i, Shuapkho, 1210m, 42.41558°
N, 44.99855°E, 7.X.2024, leg. A. Memishishi, (1
males, OPC). Allo ype: same as holo ype (1
emale, OPC).
Diagnosis. Among he siblings o he S eno-
phylax nyc e obius species complex S enophylax
bo os sp. no . is mos close o S enophylax. allas
Oláh & Ko ács, 2020, bu di e s by ha ing much
la ge size, da ke body colou ; by he dis inc
od-shape s uc u e o he ee e minal o
gonopod bo h in la e al and do sal iews as well
as by he ex emely enla ged basal egion o
pa ame e.
Desc ip ion. (In alcohol) Medium-sized ligh
b own species wi h da ke body and wing colou .
I s o ewing leng h is 16 mm wi h ounded apices.
Fo ewing eins and memb ane densely co e ed
wi h s ong and long e ec hai s. Spu numbe
034.
Male geni alia. Te gi e VIII wi h much de e-
loped black se ose p o ube ance di ided by a
middle line. Segmen IX is sho and high, pa-
allel-sided wi h g adual na owing do sad
esul ed in a e y sho band-like do sum; na ow-
ing bu longe en ad. Ce ci od-like sligh ly
cu ing do sad. Pa ap oc much p oduced domi-
na ing on he geni al s uc u e; he do sal b anch
o he pa ap oc s aigh od-like, sligh ly na -
owing apicad bo h in la e al and caudal iews;
en al b anch p oduced upwa d o ming a dis-
inc shoulde p onounced bo h in la e al and
caudal iews. Gonopods almos e ical di ec ed,
Figu es 198–201. S enophylax bo os sp. no . Holo ype male: 198 = geni alia in le la e al iew; 199 = le gonopod
and pa ap oc in caudal iew; 200 = phallic o gan in le la e al iew; 201 = Allo ype emale:
used segmen s IX and X in do sal iew.
Oláh e al.: On he T ichop e a o he Caucasus III
80
na owly used o segmen IX wi h low ee od-
like o digi a e apices. Phallic o gan composed o
he upwa d cu ing aedeagus and he s ong
spine-like pa ame e wi h ex emely b oad basal
egion; upwa d cu ing apical hal o he aede-
agus is hea ily scle o ized, black and na owing
apicad; pa ame es sho e han aedeagus.
Female desc ip ion. Sligh ly smalle han
male. Ligh b own. Fo ewing leng h 15 mm.
Con a y o male o ewing wi hou long and e ec
spine-like hai s. Fused IX and X e gi e wi h
specially pa e ned do sal shape, unique among
he known species o he S enophylax nyc e obius
species complex. O he subs uc u e o he
geni alia pa ially inju ed du ing cleaning and
clea ing p ocesses.
E ymology. bo os, om “bo os” od-like in
Hunga ian, e e s o elonga ed od o s ick-like
shape o se e al geni al subs uc u es: ce ci, pa a-
p oc , gonopod and pa ame e.
S enophylax chokhiensis Oláh & Vinçon,
sp. no .
(Figu es 202–205, Pho os 30–31)
Ma e ial examined. Holo ype: Geo gia,
M skhe a-M iane i, abo e Chokhi, nice b ook
wi h mosses, 1800-1900m, 42.4858°N, 44.7384°
E, 4.VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Pa a ype: same as holo ype (1 male).
Diagnosis. Compa ison wi h i s sibling species
is based on he phallic o gan and on he pe i-
phallic o gans d awn mos ly in caudal iew. In
la e al iew he gonopod head iangula . In
caudal iew ce ci band-like wi h mesad cu ing
apical egion; pa ap oc s wi h iangula la e al
p ocess, bu wi h digi a e apical egion and
b oadening basad; gonopod wi h iangula
lobula ed la e al and longe digi a e mesal p o-
cesses. Aedeagus is igh -angled almos pa allel-
sided along i s upwa d cu ing apical hal in
la e al iew. This cha ac e combina ion deli-
nea es his new species om he siblings o he
S enophylax solo a ewi species complex. Mos
close o S enophylax kulbak Oláh & Vinçon,
2024, bu di e s by mo e pa e ned ce ci, by
bilobed caudal s uc u e o gonopod as well as by
he cu a u e o aedeagus.
Desc ip ion (In alcohol). Compa ed o o he
membe s o he species complex S enophylax
chokhiensis sp. no . is a la ge-sized ligh aded
animal wi h b ownish, s amineous body and
wing colou . I s o ewing leng h is 15 mm wi h
ligh -spo ed ma bled memb ane.
Male geni alia. Segmen IX ex emely sho
and high, pa allel-sided wi h g adual na owing
do sad esul ed in a e y sho band-like do sum;
i s na owing en um disce nible as ully used o
gonopod. Ce ci elonga ed in la e al iew; na ow
band-like bu speci ically pa e ned. Pa ap oc
b oad in la e al iew and hin in caudal iew,
apical hal o he do sal b anch almos ili o m in
caudal iew b oadening basad a med on midway
wi h la e al iangula ou g ow h; en al b anch
o he pa ap oc iangula ho izon ally and p o-
duced la e ad. Gonopods almos e ically di ec -
ed, enla ged i s do sal egion longe han segmen
IX, wi h ape ing iangula apex in la e al iew;
bi-p ocessed in caudal iew, la e al p ocess
ounded lobula ed, mesal p ocess longe digi a e.
Phallic o gan composed o he almos igh -angled
upwa d di ec ed aedeagus and spine-like pa a-
me e wi h e ically la , pla e-like, b oad-based
basal hal ; upwa d-cu ing apical hal o he
aedeagus almos pa allel-sided.
E ymology. The name was de i ed om ha o
he ype locali y.
S enophylax cla a us (Ma yno , 1916)
Ma e ial examined. Geo gia, Ming elie-High
S ane ia, d ipping ocks, abo e Ingu i lake,
42.8585°N, 42.0336°E, 590m, 3.V.2024, leg.
Gilles Vinçon (1 male, OPC). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 5, 1252m, N41°44’36.3336, E42°4’
54.4296, 7.IX.–21.IX.2018, leg. GGBC membe s
(2 males, 5 emales; IZISUT). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 5, 1252m, N41°44’36.3336, E42°4’
54.4296, 21.IX.–5.X.2018, leg. GGBC membe s
(3 males, 3 emales; IZISUT). Geo gia, Lesse
Oláh e al.: On he T ichop e a o he Caucasus III
81
Figu es 202–205. S enophylax chokhiensis sp. no . Holo ype male: 202 = geni alia in le la e al iew; 203 = le pa ap oc
in caudal iew; 204 = le gonopod in caudal iew; 205 = phallic o gan in le la e al iew.
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 5, 1252m, N41°44’36.3336, E42°
4’54.4296, 19.V.–1.VI.2018, leg. GGBC mem-
be s (1 male, 1 emale; IZISUT). Geo gia, Lesse
Caucasus, Adja a, Kin ishi, abo e he wa e all,
malaise ap 5, 1252m, N41°44’36.3336, E42°4’
54.4296, 10.VIII.–24.VIII.2018, leg. GGBC
membe s (2 males, 1 emale; IZISUT). Geo gia,
Lesse Caucasus, Adja a, Kin ishi, abo e he
wa e all, malaise ap 4, 1264m, N41°44’
38.8824, E42°5’0.2904, 24.VIII.–7.IX.2018, leg.
GGBC membe s (4 emales, IZISUT). Geo gia,
Lesse Caucasus, Adja a, Kin ishi, abo e he
wa e all, malaise ap 4, 1264m, N41°44’
38.8824, E42°5’0.2904, 27.VII.–10.VIII.2018,
leg. GGBC membe s (1 emale, IZISUT). Geo -
gia, Sam skhe-Ja akhe i, abo e Pa a a Pamaji,
sp ing, 1950m, 41.56°N, 42.927°E, 7.VIII.2024,
leg. Gilles Vinçon (1 male, 2 emales; OPC).
S enophylax me idio ien alis Malicky, 1980
Ma e ial examined. Geo gia, Sam skhe-Ja a-
khe i, Apnia, Uppe Va dsia, Gues house Vania,
41.374°N, 43.262°E, 1290m, 5–7.X.2024 leg. J.
Junnilainen (1 male, OPC).
Rema k. This species is new o Geo gia.
S enophylax pe mis us McLaclan, 1895
Ma e ial examined. Geo gia, Kakhe i egion,
Vaslo ani Na ional Pa k, Mijnisku e 100m,
41.111°N, 46.645°E, 2-3.VII.2024, leg. Ja i Jun-
nilainen (2 male, 1 emale; OPC).
S enophylax solo a ewi (Ma yno , 1913)
Ma e ial examined. Geo gia, High S ane i,
abo e Lukhi, nice sp ing, 2320m, 43.0578°N,
42.0684°E, 17.VIII.2024, leg. Gilles Vinçon (3
males, OPC).
Rema k. This species, desc ibed om Russia
(Tebe da), is new o Geo gia.
Be aeidae Walleng en, 1891
Be aea os a a (Ma yno , 1913)
Ma e ial examined. Geo gia, M skhe a-
M iane i, Jin ali, Qalilo, 950m, 42.30445°N,
44.86915°E, 25.IV.2024, leg. A. Memishishi (1
Oláh e al.: On he T ichop e a o he Caucasus III
82
Figu es 206–208. E nodes gou iensis sp. no . Holo ype male: 206 = geni alia in le la e al iew; 207 = geni alia
in en al iew; 208 = phallic o gan in le la e al iew.
male, IZISUT; 2 males, OPC). Geo gia, Same-
g elo - Zemo S ane i, Dobe adzeni, sp ing,
42.59264°N, 42.32199°E, 370m, 28.IV.2024, leg.
A. Memishishi (2 males, 5 emales, IZISUT).
Rema k. This species was i s desc ibed in
Kakhe i (Lagodekhi), bu i now appea s o ha e a
wide dis ibu ion ac oss he High Caucasus.
E nodes bakhma ensis Oláh & Vinçon, 2024
(Pho os 7–8)
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
13.VII.–27.VII.2018, leg. GGBC membe s (1
male, IZISUT). Geo gia, Lesse Caucasus, Ad-
ja a, Kin ishi, abo e he wa e all, malaise ap
11, 1637m, 41.74769°N, 42.0951°E, 29.VI. -
13.VII.2018, leg. GGBC membe s e al. (5 males,
5 emales, IZISUT; 46 male, 19 emales; OPC).
E nodes gou iensis Oláh & Vinçon, sp. no .
(Figu es 206–208, Pho os 16–17)
Ma e ial examined. Geo gia, Gou ia, Adja a,
oad o Gomism a, nice b ook, 2120–2150m,
41.8108°N, 42.1821°E, 11.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Diagnosis. Wi hou he apomo phic apicome-
sal spine-like p ocess o E nodes sal ans Ma y-
no , 1913 and i s ela i es, E nodes bakhma ensis
Oláh & Vinçon, 2024; E nodes macahelensis
Sipahile , 1997 and E nodes o dubadensis Oláh
& Ke imo a, 2020, his new species has ela ion
o E nodes izeiensis Sipahile , 1987 desc ibed
om Tu key, bu di e s by he di e en ly shaped
a i o m apex o gonopods as well as by doubled
small ili o m and igh angled mesal p ocesses o
gonopods. I is only single, no double and sha p
angled, no igh angled a E. iziensis. Mo eo e ,
he endo hecal spine pa e n di e s, al hough i is
no disce nible clea ly in nei he species.
Desc ip ion. Male (in alcohol). Scle i es on
body and legs b own. Fo ewing leng h 6 mm.
Segmen IX sub iangula , ounded con ex an-
e ad, i s e y do sum elonga ed and mo e scle o-
ized, i s en um ounded bulky. Segmen X less
scle o ized, almos memb anous, i s apex jus
disce nible. Ce ci a e e y small, elonga ed o oid.
Pa ap oc slende , a ching only shallow. Gono-
pods clea ly a i o m, wi h e y dis inc do sal
ounded semici cula head and elonga ed apical
beak a med mesad wi h wo small inge -like
p ocesses, igh angled o gonopod sha . Phallic
o gan wi h a single dis inc s ou spine and wi h
indis inc scle o ized a ching s uc u es embedded
in endo heca.
Oláh e al.: On he T ichop e a o he Caucasus III
83
E ymology. Named a e he egion o locus
ypicus.
E nodes ming eliensis Oláh & Vinçon, 2024
(Pho os 12–13)
Ma e ial examined. Geo gia, Ming elia, abo e
Koko, nice b ookle and sp ings nea he wa e
ca chmen , on limes one subs a um, 330m,
42.648°N, 42.2033°E, 3.V.2024 leg. Gilles
Vinçon (10 males, 1 emale; OPC). Geo gia,
Ming elie, abo e Koko, nice b ookle s and
sp ings nea he wa e ca chmen , on limes one
subs a um, 310m, 42.6467°N, 42.2028°E, 3.V.
2024, leg. Gilles Vinçon (1 male, 1 emale; OPC).
Geo gia, Ming elie, Sou heas J a i, nice b ook
and la e al sp ings, 340–350m, 42.6956°N,
42.0757°E, 3.V.2024, leg. Gilles Vinçon (12
males, OPC). Geo gia, Ime e i, Sou hwes Saja-
akho, ‘Be dnis Tska o’, nice sp ings and
d ipping ocks, 80m, 42.0636°N, 42.2693°E and
100m, 42.055°N, 42.275°E, 28.IV.2024, leg.
Gilles Vinçon (16 males, OPC).
Rema k. This is he i s epo om he Lesse
Caucasus.
Se icos oma oidea S ephens, 1836
Se icos oma idae S ephens, 1836
The Se icos oma idae amily consis s o ge-
ne ic axa cha ac e ized by a iable s uc u es de-
eloped on he basal segmen o he male an-
ennae and on he maxilla y palpi. These male-
speci ic s uc u es a e he ou come o unusual,
acciden al, and andom gene ic de elopmen s,
exp essed h ough sexual o ganiza ion p ocesses
in luenced bo h by p ema ing cou ship and by
c yp ic emale choice du ing copula ion. In spi e
o his “cu iosi y shop” he biza e modi ica ions
on head appendages, he Eu opean se icos oma id
males in mos gene a ha e e y simila geni al
s uc u es. These biza e modi ica ions and neo-
o ma ions a e a he species g oup speci ic and
no genus speci ic al e a ions as was demons a ed
by Wea e (2002) o Lepidos oma idae, p e-
sen ed by Oláh and Johanson (2011) o
Hyd op ilidae and discussed by Oláh and Flin
(2012) in he Leuco ichini ibe o Hyd op ilidae
amily.
Ce asma McLachlan, 1876
(Map 8)
Ce asma genus was es ablished mos ly by a
pai o biza e ho n-like o ma ion on he head o
males being p oduced pos e io ly and u nished
wi h a pencil o long hai s. Ac ually i is he
backwa d di ec ed pa o he highly modi ied
scape, he i s an ennal segmen . Bo h he gene ic
name, Ce asma (ke as ho n in G eek) as well as
he name o he nominal name-bea ing ype
species, Ce asma co nu a McLachlan, 1876
(co nu ho n in La in) e e s o his ho n-like
s uc u e on he scape. This mono ypic genus was
long ep esen ed only by he ype species de-
sc ibed om he single male holo ype collec ed
somewhe e in Gou ia, Geo gia. The second
species, Ce asma cai on was desc ibed by
Malicky (1986). Howe e , one yea la e Sipahile
& Malicky (1987: page 96, igu e 4a,b,c)
published new d awings unde he same name,
Ce asma cai on wi hou explaing why hey ha e
changed he d awings o he species and which
specimen (holo ype ed awing, pa a ypes o o he
specimen) was d awn esul ing he new d awings
wi h di e en s uc u al pa e n.
Ce asma species a e usually di icul o
collec . Mos species a e ep esen ed only by ew
specimens. Howe e , in a Malaise- ap p ojec on
he Kin ishi Ri e , Geo gia we we e lucky o
collec al oge he 412 males o Ce asma kin isha
sp. no . pe mi ing us o sea ch and s udy he
s abili y and he a iabili y anges o he mos
di e se s uc u e o he male geni alia, he
specia ion ai . Simila ly o Se icos oma genus
(Oláh e al. 2023) we ha e ound he hea ily
scle o ized i illa ing s uc u e, he pa ap oc wi h
cha ac e s a e o specia ion ai ha is being
mos di e se and mos s able gene ically p o ec ed
adap i e non neu al geni al s uc u e. Based on
he di e gences o he do sal and en al a ms o
b anches o pa ap oc he e we desc ibe six new
species and we suppose ha many mo e unknown
Oláh e al.: On he T ichop e a o he Caucasus III
84
Map 8. Ce asma genus (7 species): C. a ina sp. no ., C. bakhma a sp. no ., C. cai on Malicky, 1986, C. gode da sp. no .,
C. goma sp. no ., C. kin isha sp. no ., C. za zma sp. no .
species a e s ill wai ing o be collec ed and de-
sc ibed.
The “cu iosi y shop”, he biza e modi i-
ca ions, neo o ma ions de eloped on he second
segmen o he maxilla y palp and on he scape,
he i s segmen o an ennae also in he Ce asma
genus. The second segmen o maxilla y palpi
p essed agains he on o he ace ans e sally
and o each o he sagi ally a e con ex ex e nally
and conca e in e nally; conca i y con ains h ee
basal slende digi i o m whi e and an apical single
spoon-like black s uc u es ending in pencil o
hai s; his black s uc u e could be he hi d
segmen o he maxilla y palp. Scape, he i s
an ennal segmen highly modi ied in o backwa d
di ec ed ho nlike s uc u e o mesal conca i y
illed wi h memb anous e ec ile ex endable whi e
s uc u e ending in pencil o hai s. These mem-
b anous e ec i e s uc u es oge he wi h hai s/
scales in he conca i ies o modi ied maxilla y
palp and scape could be in ol ed in he phe o-
mone p oduc ion, dis ibu ion, o ol ac o y unc i-
ons o he scen p oducing and oconial o gan.
Ce asma a ina Oláh, sp. no .
(Figu e 209, Map 8)
Ma e ial examined. Holo ype: Tu key, il.
A in, Col Cancu alan, 800m, 1–2.VII.1996,
leg. A. Podlussány (1 male, OPC). Pa a ype: same
as holo ype (1 male, OPC).
Diagnosis. This new species has esemblance
o Ce asma cai on, bu di e s by he iangula ,
no ounded an e io ma gin o segmen X; sho ,
no long ce cus; by la e al p o ile o gonopod as
well as by s ai , no downwa d cu ing do sal
a m o pa ap oc and mo e p oduced apical
pa e n o en al a m o pa ap oc .
Desc ip ion. Male (in alcohol). Median b own
species wi h median b own appendages. Second
segmen o maxilla y palpi and scape, i s
an ennal segmen a e highly modi ied in o an
enla ged complex uni wi h deep conca i y o
phe omone p oduc ion, dis ibu ion, o ol ac o y
unc ions o he scen p ocessing and oconial
o gan. Cephalic and ho acic scle i es b own;
Oláh e al.: On he T ichop e a o he Caucasus III
85
Figu e 209. Ce asma a ina sp. no . Holo ype male:
geni alia in le la e al iew.
Figu e 210. Ce asma bakhma a sp. no . Holo ype male:
geni alia in le la e al iew.
da ke on do sum. Spu numbe 224. Fo ewing
leng h 13 mm; wing memb ane b own, densely
co e ed wi h decumben se ae.
Male geni alia. Segmen IX sub iangula in
la e al iew; en um li le longe . Segmen X less
scle o ized delimi ed la e ally by as well as pa i-
ally used o he hea ily scle o ized pai o spine-
like pa ap oc s. Pa ap oc composed o longe
do sal and sho e en al a m; do sal a m s aigh
wi h upwa d di ec ed head; i s apical ma gin ully
conca e; en al a m o pa ap oc wi h a ounded
hump and sho spine midway on apical ma gin.
Gonopods wi h much p oduced basal iangula
p ocess in la e al iew. Phallic o gan sligh ly
cu ing basad dila ed en ad on i s apical egion.
E ymology. Named a e he egion o he
locus ypicus.
Ce asma bakhma a Oláh & Vinçon, sp. no .
(Figu e 210, Map 8, Pho os 18–19)
Ma e ial examined. Holo ype: Geo gia, Gou-
ia, oad o Bakhma o, sp ing, 2040m, 41.8586°N,
42.3564°E, 10.VIII.2024, leg. Gilles Vinçon (1
male, OPC).
Diagnosis. This new species has esemblance
o Ce asma gode da sp. no . bu di e s by he
iny iangula p ocess on midway o gonopod
en um, less p oduced upwa d di ec ed head o
he do sal a m o pa ap oc and by he backwa d
di ec ed, no igh angled do soapical p ocess o
en al a m o pa ap oc .
Desc ip ion. Male (in alcohol). Median b own
species wi h median b own appendages. Second
segmen o maxilla y palpi and scape, i s an-
ennal segmen a e highly modi ied in o an
enla ged complex uni wi h deep conca i y o
phe omone p oduc ion, dis ibu ion, o ol ac o y
unc ions o he scen p ocessing and oconial
o gan. Cephalic and ho acic scle i es b own;
da ke on do sum. Spu numbe 224. Fo ewing
leng h 12 mm; wing memb ane b own, densely
co e ed wi h decumben se ae.
Male geni alia. Segmen IX ounded sub i-
angula in la e al iew; en um li le longe .
Segmen X less scle o ized delimi ed la e ally by
as well as pa ially used o he hea ily scle o ized
pai o spine-like pa ap oc s. Pa ap oc composed
o longe do sal and sho e en al a ms; do sal
a m wi h upwa d di ec ed, less enla ged head;
en al a m o pa ap oc wi h sligh ly backwa d
di ec ed do soapical spine-like p ocess. Gonopods
wi h a small iangula en al p ocess on midway
in la e al iew. Phallic o gan sligh ly cu ing
basad dila ed en ad on i s apical egion.
E ymology. Named a e he egion o he
locus ypicus.
Ce asma cai on Malicky, 1986
(Map 8)
Ce asma cai on Malicky, 1986:233–235: “Holo ypus
♂: Tü kei, P o inz Rize, 1 km südlich on O , 20
Oláh e al.: On he T ichop e a o he Caucasus III
92
Figu e 224. No idobia ekona sp. no . Holo ype male:
geni alia in le la e al iew.
Diagnosis. This new species ha ing long,
upwa d di ec ed spine-like pa ap oc is a membe
o No idobia demel i species complex (Oláh e al.
2022) and mos close o No idobia demel i
Malicky, 1974, bu di e en ia ed by he mo e
slende and downwa d u ning pa ap oc as well
as by he en al p ocess o gonopod wi h upwa d,
no downwa d di ec ed apical, subapical poi ed
p ocess.
Desc ip ion. Da k cas anean species wi h
much ligh e legs. Maxilla y palpi p essed agains
he on o he ace, con ex ex e nally and
conca e in e nally, no any silky hai s disce nible
in he in e nal ca i y. The i s an ennal segmen ,
he scape is highly modi ied as p essing agains
each o he ’s and loca ed sunken in a deep
conca i y o clypeus o ming oge he an upwa d
di ec ed join p ocess and a med wi h modi ied
se ae and whi e ou g own issue. Cephalic and
ho acic scle i es da k b own, almos black
especially on he do sum; appendages including
legs ligh e b own; haus ellum and in e scle i al
memb anous egumen s whi ish. Fo ewing leng h
10 mm; wing memb ane b own, densely co e ed
wi h decumben se ae.
Male geni alia. Segmen IX sub iangula in
la e al iew; iangula in e scle i al window in
do sal iew ha is a pu a i e s uc u e o ac o-
e gi e b ace. Segmen X less scle o ized deli-
mi ed la e ally by he hea ily scle o ized pai o
spine-like pa ap oc s. Pa ap oc s ha ing b oad
basal ho izon al egion su ounding segmen X
and a long, upwa d u ning s aigh slende spine
ob use angled on one hi d and wi h apical egion
o den al e minal. Gonopods quad angula bea a
en al p ocess wi h an apex wi h a subapical
upwa d di ec ed spine-like poin ed ou hg ow h in
la e al iew. Phallic o gan sligh ly cu ing egula
ube wi h en oapical mo e scle o ized
p ojec ion.
E ymology. ekona, om “ ékony” slim,
slende in Hunga ian, e e s o he la e al p o ile
o he do sal a m o he pa ap oc , mos slende in
he species complex
Schizopelex akhalchala Oláh & Vinçon, 2024
(Pho os 5–6)
Ma e ial examined. Geo gia, Racha-Lech-
khumi and Lowe -S ane i, below Akhalchala, 2
nice sp ings, 1870m, 42.6768°N, 42.6482°E, 13.
VIII.2024, leg. Gilles Vinçon (1 male, OPC).
Geo gia, Racha-Lechkhumi and Lowe -S ane i,
Akhalchala, o en abo e cascade, 2400m, 42.7°
N, 42.6455°E, 13.VIII.2024, leg. Gilles Vinçon (1
male, OPC). Geo gia, Racha-Lechkumi and
Lowe -S ane i, Akhalchala, b ook and sp ing,
2170m, 42.6947°N, 42.6463°E, 13.VIII.2024, leg.
Gilles Vinçon (2 males, OPC).
Rema ks. In en al iew he en al p ocess is
no unca ed and di e en in he wo specimens.
I is a he con ex apicad in he specimen om
he sp ing and wi h a minu e middle excision
apicad in he specimen om he o en .
Schizopelex ming elia Oláh & Vinçon, 2024
(Pho os 5–6, 24–25, 37)
Ma e ial examined. Geo gia, Ming elia, abo e
Koko, nice b ookle and sp ings nea he wa e
ca chmen , on limes one subs a um, 330m,
42.648°N, 42.2033°E, 3.V.2024 leg. Gilles Vin-
çon (3 males, OPC). Geo gia, Babili, sp ing,
790m, 42.79156°N, 42.74598°E, 27.IV.2024, leg.
A. Memishishi (3 males, 1 emale; IZISUT).
Geo gia, Racha-Lechkhumi and Lowe -S ane i,
abo e Kheledi, sp ing, 1180m, 42.8037°N, 42.63
Oláh e al.: On he T ichop e a o he Caucasus III
93
35°E, 14.VIII.2024, leg. Gilles Vinçon (10 males,
OPC).
Rema ks. This species ha ing used en al
p ocess wi hou den a e o g anulised su ace be-
longs o he Schizopelex ana olica species g oup
and a sibling species o Schizopelex akhalchala
Oláh & Vinçon, 2024, bu di e s by he la ge
do sal lobe o he gonopod wi hou any es igial
no ch o ma ion, shallowly s epwise do sum o
segmen X and by he bilobed, no unca e apex
o he he en al p ocess cons i u ed by he basal
pla e o gonopods and by he en al p ocess o
segmen IX. One specimen om he collec ed
h ee males o abo e Koko has less bilobed
en al p ocess, bu segmen X and he gonopod
a e ypical. Ano he specimen om he collec ed
h ee males a Babili has some es igial no ch
p esen on he gonopod, bu he bilobed en al
p ocess and segmen X a e ypical. Ten males
om abo e Kheledi ha e gonopod wi hou any
es igial no ch o ma ion, shallowly s epwise
do sum o segmen X, bu a iously (shallowly o
deeply) bilobed en al p ocess.
Schizopelex pon ica Ma yno , 1913
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 4, 1264m, N41°44’38.8824, E42°5’0.2904,
27.VII. - 10.VIII.2018, leg. GGBC membe s (1
male, IZISUT). Geo gia, Adja a, Gode dzi Pass,
a e Beshumi Bo anic Ga den, b ook and sp ing,
1970m, 41.6216°N, 42.5383°E, 7.VIII.2024, leg.
Gilles Vinçon (3 males, OPC). Geo gia, Sam-
skhe-Ja akhe i, abo e U kisubani, b ook and
sp ing, 1870m, 41.676°N, 42.5765°E, 8.VIII.
2024, leg. Gilles Vinçon (1 male, OPC).
Se icos oma g usiense Ma yno , 1913
Ma e ial examined. Geo gia, Lesse Caucasus,
Adja a, Kin ishi, abo e he wa e all, malaise
ap 5, 1252m, N41°44’36.3336, E42°4’54.4296,
29.VI.–13.VII.2018, leg. GGBC membe s (1
male, IZISUT).
Lep oce oidea Leach, 1815
Lep oce idae Leach, 1815
A h ipsodes ul ico nis Ma yno , 1913
Ma e ial examined. Geo gia, Adja a, abo e
Namonas e i, nice b ook and sp ing, 1710m,
41.555°N, 42.0853°E, 12.VIII.2024, leg. Gilles
Vinçon (1 male, OPC).
Se odes i idis new species g oup
While sea ching and inding he axonomic
posi ion and ela ion o ou new species Se odes
io icus Oláh & Memishishi sp. no . we es ablish
he e he Se odes i idis new species g oup. This
species g oup is cha ac e ized by used ce ci and
pa ap oc elonga ed and di e si ied by a ious
apical endings as well as by gonopod ha ing
di e si ied pla e-like basodo sal a m. Bo osa-
neanu & Gasi h (1971) while s udying caddis ly
auna o Is ael desc ibed Se odes i idis hulio-
hica new subspecies om Is ael and Se odes
i idis i anensis new subspecies om I an in
addi ion o nomina e species Se odes i idis
i idis Fou c oy, 1785. La e Kumansky in
Kumanski & Malicky (1976) desc ibed he ou h
subspecies Se odes i idis bulga icus om Bul-
ga ia. He e we aise hei subspecies o species
ank upplying ou uni ied phylogene ic species
concep and aking subspecies and ace ou o
science (Oláh e al. 2018). Species o Se odes
holoce cus Na as, 1923 desc ibed om Spain and
Se odes kugle i Bo osaneanu & Gasi h, 1983
desc ibed om Is ael seem o belong o his
species g oup as well.
Se odes io icus Oláh & Memishishi, sp. no .
(Figu es 225–226)
Ma e ial examined. Holo ype: Geo gia, Ka-
ke hia, Chachuna Managed Rese e, Ioi i Ri e
alley, Chachuna ho el, 260m, 41.220°N, 45.971°
E, 28–30.VI.2024 ligh leg. Ja i Junnilainen, (1
male, OPC).
Oláh e al.: On he T ichop e a o he Caucasus III
94
Figu es 225–226. Se odes io icus sp. no . Holo ype male: 225 = geni alia in le la e al iew; 226 = geni alia in do sal iew.
Diagnosis. Ou new species Se odes io icus sp.
no . di e s om all known species by ha ing
e y di e ged and complex head o he used ce ci
and pa ap oc . This pa icula ly p oduced used
s uc u e is he mos di e se cha ac e s a e in his
species complex.
Desc ip ion. (In alcohol) This new Se odes is a
yellowish ligh b own e y slende and agile
animal wi h o ewing leng h o 8 mm. Male
geni alia. La e al p o ile o segmen IX egula
iangula . Segmen X less scle o ized and na -
owing elonga ed; i s e minal egion na ow lying
in be ween he la e al wings p oduced on he
middle egion o phallic o gan. Fused ce ci and
pa ap oc , he mos di e se, he e o e he p obable
specia ion ai o he Se odes i idis species
complex has p oduced a di e gen e y pa i-
cula ly o ganised apical egion o widened semi-
ci cula head a med wi h an apical poin ed spine;
he head is hea ily and densely co e d wi h sho
ee h. Basodo sal a m o gonopod densely co-
e ed wi h longe spine-like se ae o ele a ed
al eoli, sligh ly bu dis inc ly bilobed apicad;
lowe a m o gonopod s aigh . Phallic o gan
cu ing hal ci cula wi h a pai o do sal ele a ed
ounded wings midway housing he apical egion
o segmen X; i s basal egion connec ed o used
ce ci and pa ap oc do sad as well as o gonopods
en ad.
E ymology. Named a e he egion o he lo-
cus ypicus.
T iaenodes kaw aiskii Ma yno , 1909
Ma e ial examined. Geo gia, Kake hia, Cha-
chuna Managed Rese e, Ioi i Ri e alley,
Chachuna ho el, 260m, 41.220°N, 45.971°E, 28-
30.VI.2024, ligh leg. Ja i Junnilainen, (6 males, 5
emale; OPC).
Rema k. The genus and species a e new o he
Geo gian auna.
Acknowledgemen – We would like o exp ess ou
g a i ude o he CaBOL p ojec eam membe s o hei
assis ance du ing he ieldwo k and hei help wi h so ing
he malaise ap ma e ial a Ilia S a e Uni e si y. We a e
g a e ul o Finnish lepidop e ologis , Ja i Junnilainen o he
caddis ly specimens he has collec ed on ligh in Geo gia.
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