E-mail: lubomi .ko [email protected]
Fi s eco d o a ep esen a i e o he o de Palpig adi
(Chelice a a: A achnida) om Se bia
D agan ANTIĆ1,2, Ľubomí KOVÁČ3
1 Uni e si y o Belg ade—Facul y o Biology, Ins i u e o Zoology, S uden ski T g 16, 11000 Belg ade, Se bia
2 Se bian Biospeleological Socie y, T g Dosi eja Ob ado ića 2, 21000 No i Sad, Se bia
3 Depa men o Zoology, Ins i u e o Biology and Ecology, Facul y o Science, Pa ol Joze Ša á ik Uni e si y,
Š obá o a 2, 041 54 Košice, Slo akia
Biologia Se bica, 2025, 47: 49-53
O iginal pape
DOI: 10.5281/zenodo.17679386
Summa y. This con ibu ion e e s o he i s eco d o he genus Eukoenenia Bö ne , 1901 (A achnida: Palpig adi)
in Se bia. One adul male was disco e ed in P o alija Ca e, a pa o he Ce janska Ca e Sys em, nea Niš in eas e n
Se bia. Based on diagnos ic mo phological cha ac e s, habi a p e e ences and he dis ibu ion o simila axa, we con-
clude ha i belongs o he Eukoenenia spelaea species complex.
Keywo ds: Balkan Peninsula, Ce janska Ca e Sys em, Eukoeneniidae, mic owhip sco pions, oglobion s.
Accep ed: 7 No embe 2025 / Published online: 11 No embe 2025
INTRODUCTION
Palpig adi, o mic owhip sco pions, a e iny, unpig-
men ed and eyeless a achnids ha a e o en o e looked in
hei na u al habi a . Species o his o de inhabi he soils
o opical and sub opical egions o he wo ld, bu a e also
ound in sub e anean habi a s (Condé 1996). In Eu ope,
he e a e nume ous eco ds o palpig ades associa ed wi h
ca es (Mammola e al. 2021), wi h hei no he nmos e-
po ed dis ibu ion limi a 49° no h in he Wes e n Ca -
pa hians in Slo akia (Ko áč e al. 2002). To da e, Palpig adi
ha e been ound in mos o Cen al and Sou he n Eu ope,
bu no indings ha e been epo ed om Se bia. Du ing e-
cen biospeleological explo a ion o ca es in Se bia, an adul
male was disco e ed in P o alija Ca e, which is pa o he
Ce janska Ca e Sys em nea Niš. This is he i s eco d o
Palpig adi om Se bia.
MATERIAL AND METHODS
Si e cha ac e is ics – ca e desc ip ion
The Ce janska Ca e Sys em is one o he la ges unde -
g ound sys ems in Se bia; and is loca ed in he no he n pa
o he Kala a Massi in he eas e n pa o he coun y, no h
o Niš (Fig. 1). Due o i s excep ional geomo phological, hy-
d ological and aes he ic alue, he ca e sys em has been p o-
ec ed as a na u al monumen since 1998, unde he name
‘Ce janska Ca e’ by dec ee o he Go e nmen o he Republic
o Se bia. The en i e sys em de eloped in he limes one o
Lju i V h (Nešić 2023a). The i s documen ed explo a ions
o he ca e da e back o he ea ly 1970s. The ca e is ichly
deco a ed wi h speleo hems, including s alac i es, s alagmi es,
d ape ies, ca e co als, and c ys al lowe s. In addi ion, he-
lic i es a e especially abundan , and a e one o he mos e-
Fi s eco d o a ep esen a i e o he o de Palpig adi (Chelice a a: A achnida) om Se bia
D. An ić e Ľ. Ko áč
ma kable ea u es o Ce janska Ca e. Acco ding o he mos
ecen da a, Ce janska Ca e is he hi d longes ca e sys em
in Se bia, wi h 7,170 me e s o explo ed passages. Fou mo -
phological and gene ic uni s can be dis inguished: P o alija
Ca e, K a ljanska Pi , he in low passages, and Peću ina
(Nešić 2023b; Nešić and Jo ić 2023).
P o alija Ca e is he dominan componen o he sys-
em. I ep esen s a la ge adose pono ca e, wi h a o al
leng h o 5,703 m, including 3,675 m o ac i e i e passage.
I has a simple mo phology and consis s mainly o a i e
channel, which ex ends o e wo le els and has a leng h o
4,903 m, as well as se e al side passages wi h a combined
leng h o 800 m (Nešić 2023b). The i s biospeleological in-
es iga ions o he Ce janska Ca e Sys em we e conduc ed
a he end o 1999. Be ween 1999 and 2015, esea che s om
he Ins i u e o Na u e Conse a ion o Se bia, in collabo a-
ion wi h speleologis s, su eyed se e al sub e anean si es
wi hin he Kala a Massi , including P o alija Ca e. In 2017,
biospeleological s udies we e conduc ed by membe s o he
Se bian Biospeleological Socie y, and mos ecen ly, in 2024,
by biospeleologis s om he Uni e si y o Belg ade – Facul y
o Biology. Acco ding o Pa iće ić e al. (2023), 20 a h opod
species ha e been eco ded om P o alija Ca e, he majo -
i y o which a e oglophiles and ogloxenes. Howe e , ou
oglobi ic species a e known om he ca e: he ha es man
Pa anemas oma bu eschi (Roewe , 1926); he millipedes Daz-
bogosoma naissi Maka o & Ću čić, 2012 and Balkanodes-
minus se bicus An ić, Vagalinski, S oe & Akka i, 2022; and
he ca abid bee le Du alius anjensis p o alijae Pa iće ić,
Za ezalo & Popo ić, 2016 (Maka o e al. 2012; An ić e al.
2022; Pa iće ić e al. 2023).
Reco d speci ica ion
Se bia, Niš, P o alija Ca e, abou 1,000 m om he ca e
en ance, 1 adul male, on a e y we s one, 19.11.2024, leg.
D. An ić.
One pe manen slide has been deposi ed a he Ins i-
u e o Zoology, Uni e si y o Belg ade – Facul y o Biology,
Belg ade, Se bia.
Pe manen slide p epa a ion and mo phology s udy
Mo phology o he specimen was s udied using a Ca l
Zeis Axiolab 5 ligh mic oscope equipped wi h phase con-
as . Fo his pu pose, he specimen was boiled gen ly in an
e apo a ion glass dish in 95% e hyl alcohol o app oxima ely
1 minu e o emo e a om he body. Fo clea ing, he speci-
men was ans e ed o a 10% KOH aqueous solu ion in a
conca e glass dish o ca. 1 minu e and hen ans e ed o a
dish con aining chlo ophenol un il i was ully anspa en .
Finally, he specimen was moun ed in Swann medium on
a pe manen slide and le o d y. A e d ying, he co e
glass was inged wi h Canada balsam o seal he moun ing
medium and p e en desicca ion.
Fig. 1. Loca ion o P o alija Ca e.
50 Biologia Se bica 47
Fi s eco d o a ep esen a i e o he o de Palpig adi (Chelice a a: A achnida) om Se bia
Map and pho og aphy
The dis ibu ion map was c ea ed in Google Ea h P o
( e sion 7.3.6.10201) and Adobe Pho oshop CS6. Pho os o
he li ing male in si u we e aken wi h an Olympus S ylus
Tough TG-6 digi al came a. Images o mo phological s uc-
u es we e eco ded using a Leica DM2500 ligh mic oscope
equipped wi h DIC (di e en ial in e e ence con as ). The
inal images we e edi ed and combined using Adobe Pho-
oshop CS6.
RESULTS
Taxonomy
Class A achnida
O de Palpig adi
Family Eukoeneniidae
Genus Eukoenenia Bö ne , 1901
Diagnosis. Rep esen a i es o he genus exhibi he ol-
lowing diagnos ic cha ac e s: opis hosomal s e ni es IV–VI
wi hou en al sacs; segmen IX is na owe han VIII, bu
wide han XI; and he opis hosomal s e ni es IV and VI
each o m a p o ube ance (Ba anco and Ha ey 2008).
Eukoenenia spelaea species complex
Ch is ian e al. (2014) de ined he Eukoenenia spelaea
complex based on he ollowing cha ac e s: la e al o gan
wi h 3 o mo e blades; coxae o legs I–IV wi h 0–4–4–1 hick
se ae; leg I a3 wi h 4 o ked se ae; leg IV b a wi h 4 se ae;
me apel idium wi h 3 se ae; e gi e II wi h 2+2, e gi es III-
VI usually wi h 3+3 se ae , median se a absen ; s e ni es
IV–VI wi h 3+3 o mo e se ae a; emale wi h 3+3 apical se ae
on he i s geni al lobe and 3 se ae on he second geni al
lobes; male wi h 3 se ae on he second and 4 on he hi d
geni al lobes; opis hosoma wi h a dis inc na owing be ween
segmen s VIII and IX.
Mo phological analysis
The male om P o alija Ca e shows: chelice a wi h 8
ee h (Fig. 2A); la e al o gan on each side o he p osoma
wi h 3 blades (Fig. 2B); deu o i os e num wi h 5 se ae a -
anged in he shape o he le e V; se al pa e n o opis -
hosomal s e ni es IV–VI as a1, a2, a3, s1, s2; s e nal po es
p esen ; segmen XI o opis hosoma wi h 11 se ae; pa e n
o o ked se ae on a sal segmen 3 on leg I (leg I a3): 1+1+2
o ked se ae, se a s in la e al posi ion (Fig. 2C); se al pa e n
o male geni al pla e: each hal o 1s lobe wi h 2 s e nal se ae,
Fig. 2. Mo phological cha ac e s obse ed in he male o Eukoenenia c . spelaea (Peye imho , 1902) om P o alija Ca e. A, e minal seg-
men o chelice a ( inge ); B, la e al o gan; C, pa e n o o ked se ae on a sal segmen 3 o leg I; D, male geni al pla e. (Scale ba s: 0.01 mm).
Biologia Se bica 47 51
Fi s eco d o a ep esen a i e o he o de Palpig adi (Chelice a a: A achnida) om Se bia
D. An ić e Ľ. Ko áč
9 se ae on lobe ex ension (3 la e al, 6 submedian) and 2 api-
cal usules, se al pa e n o 2nd and 3 d olds unclea (Fig. 2D);
lagellum wi h 7 segmen s.
The lagellum is a mo phological s uc u e wi h some
axonomic signi icance. Due o i s agili y, i some imes
b eaks o pa ially in li ing animals and is o en comple ely
los in p ese ed specimens. Bo h apply o he animal iden-
i ied om P o alija Ca e. Be o e i was cap u ed, i had a
b oken lagellum wi h 7 segmen s (Fig. 3), ins ead o he
12–15 segmen s ound in in ac specimens o he E. spelaea
complex.
DISCUSSION
Because his was he i s eco d o a palpig ade in Se -
bia, e o s we e made o iden i y he specimen a he spe-
cies le el. Ini ial inspec ion o he eshly moun ed animal
e ealed ha he geni al pla e belonged o an adul male. A
se al pa e n o a1, a2, a3, s1, s2 on he opis hosomal s e -
ni es IV–VI s ongly indica ed a ilia ion wi h he E. spelaea
complex. The p esence o 3 o mo e se ae ‘a’ pe hemis e ni e
is one o he (albei no exclusi e) ea u es o his complex
(Ch is ian e al. 2014). Only E. aus iaca (Hansen, 1926),
which appea s o be ela ed o E. spelaea, has 2 se ae ‘a’, like
many o he Eukoenenia species.
Du ing he examina ion, he sample was damaged
unde he co e glass, ende ing some mo phological de ails
un ecognizable. Howe e , wha we we e able o disce n was
compa ible wi h an assignmen o he E. spelaea complex:
he pa e n o he o ked se ae on a sal segmen 3 o leg I,
he numbe o blades in he la e al o gan, and he numbe
and a angemen o se ae on he deu o i os e num. Only he
numbe o se ae on opis hosoma XI was sligh ly highe han
in mos popula ions o he E. spelaea complex (11 s. 9–10).
Un il u he diagnos ic cha ac e is ics can be examined in
o he palpig ades om P o alija Ca e, we e e o his popu-
la ion as Eukoenenia c . spelaea (Peye imho , 1902).
The known dis ibu ion ange o Eukoenenia spelaea
wi h i s ou subspecies ex ends om F ance, wes o he
Rhône, ac oss he Alpine a c o he wes e n Ca pa hians
and he no he n Dina ides. The o he species o he E. spe-
laea complex (E. aus iaca, E. bonadonai Condé, 1979, E.
lanai Ch is ian in Ch is ian e al. 2014, E. oscia Ch is ian in
Ch is ian e al. 2014, E. s ina ii Condé, 1977) inhabi much
smalle a eas and a e dis ibu ed wi hin he la ge ange o
E. spelaea, wi h he excep ion o he Sou h Ca pa hian E.
ma ga e ae O ghidan, Geo gesco & Sâ bu, 1982 and E. con-
dei O ghidan, Geo gesco & Sâ bu, 1982. The eco d o E. c .
spelaea in P o alija Ca e ex ends he dis ibu ion ange o
he en i e species complex o he sou h.
The species in he E. spelaea complex a e only mod-
e a ely oglomo phic: mo phological adap a ions o ca e
Fig. 3. Habi us o he collec ed male o Eukoenenia c . spelaea (Peye imho , 1902) om P o alija Ca e. Pho os: D. An ić.
52 Biologia Se bica 47
Fi s eco d o a ep esen a i e o he o de Palpig adi (Chelice a a: A achnida) om Se bia
li e, such as elonga ed ex emi ies, an elonga ed lagellum
and a highe numbe o blades in he la e al o gan a e no
e y p onounced. Like he as majo i y o Eu opean palpi-
g ades, hey a e oglobion , and es ic ed o sub e anean
habi a s. The ac ha wo o hem (E. spelaea and E. aus-
iaca) ha e been ound in Aus ia, no only in ca es bu also
nea he su ace in loose sedimen s (Ch is ian 2004) does
no con adic his: such habi a s a e conside ed ‘shallow
sub e anean habi a s’ sensu Cul e and Pipan (2014). The
species iden i ied in P o alija Ca e can also be conside ed
o be oglobion .
ACKNOWLEDGMENTS
We a e deeply g a e ul o he anonymous e iewe and
especially o E ha d Ch is ian (Aus ia) o hei co ec ions
and commen s, which g ea ly imp o ed he i s e sion o
his manusc ip .
The pe mi o explo a ion o P o alija Ca e was issued
by he manage o he p o ec ed Na u al Monumen “Ce jan-
ska Pećina”, Public En e p ise “Di ec o a e o Cons uc ion
o he Ci y o Niš” e e ence numbe 03-4937-2/2021, da ed
Oc obe 29, 2021.
The p esen s udy was pa ly suppo ed by he Minis y
o Science, Technological De elopmen and Inno a ion o
he Republic o Se bia (451-03-137/2025-03/ 200178).
D agan An ić g a e ully acknowledges he company o
S ećko Ću čić and Aleksanda Pegan du ing explo a ion o
P o alija Ca e in No embe 2024.
REFERENCES
An ić DŽ, Vagalinski B, S oe P, Akka i N. 2022. A e iew o he ca e nico
-
lous T ichopolydesmidae (Diplopoda, Polydesmida) om he Ca -
pa hian-Balkan a ch and he Rhodope Moun ains, wi h he desc ip-
ion o wo new gene a and h ee new species. ZooKeys. 1097:1–46.
h ps://doi.o g/10.3897/zookeys.1097.83916.
Ba anco P, Ha ey MS. 2008. The i s indigenous palpig ade om Aus-
alia: a new species o Eukoenenia (Palpig adi: Eukoeneniidae). In-
e eb a e Sys ema ics. 22:227–233. h ps://doi.o g/10.1071/IS07031.
Ch is ian E. 2004. Palpig aden (Tas e läu e ) – Spinnen ie e in eine Wel
ohne Lich . Denisia. 12:473–483. h ps://www.zoboda .a /pd /deni-
sia_0012_0473-0483.pd .
Ch is ian E, Isaia M, Pasche a M, B uckne A. 2014. Di e en ia ion among
ca e popula ions o he Eukoenenia spelaea species-complex (A ach-
nida: Palpig adi) in he sou hwes e n Alps. Zoo axa. 3794(1):52–86.
h ps://doi.o g/10.11646/zoo axa.3794.1.2.
Condé B. 1996. Les Palpig ades, 1885–1995: acquisi ions e lacunes. Re ue
Suisse de Zoologie. ol. ho s sé ie: 87–106.
Cul e DC, Pipan T. 2014. Shallow sub e anean habi a s. Ecology, E olu-
ion, and Conse a ion. Ox o d and New Yo k: Ox o d Uni e si y
P ess.
Ko áč Ľ, Mock A, Ľup áčik P, Palacios-Va gas JG. 2002. Dis ibu ion o
Eukoenenia spelaea (Peye imho , 1902) (A achnida, Palpig adida) in
he Wes e n Ca pa hians wi h ema ks on i s biology and beha iou .
In: Tajo ský K, Balík V, Pižl V, edi o s. S udies on soil auna in Cen-
al Eu ope. P oc. 6 h Cen al Eu opean Wo kshop on Soil Zoology,
Ins i u e o Soil Biology AS CR, České Budějo ice. p. 93–99.
Maka o SE, Ću čić BPM, Milinčić M, Pecelj M, An ić D, Mi ić BM. 2012.
A new genus and species o he amily An h oleucosoma idae om
Se bia (My iapoda, Diplopoda, Cho deuma ida). A chi es o Bio-
logical Sciences, Belg ade. 64(2):801–806. h ps://doi.o g/10.2298/
ABS1202793M.
Mammola S, Souza MF, Isaia M, Fe ei a RL. 2021. Global dis ibu ion o
mic owhip sco pions (A achnida: Palpig adi). Jou nal o Biogeog a-
phy. 48:1518–1529. h ps://doi.o g/10.1111/jbi.14094.
Nešić D. 2023a. Posi ion, geological composi ion, elie and hyd olog aphy
o he no he n pa o Kala a . In: Nešić D, Jo ić D, edi o s. Na u al
Monumen Ce janska Ca e. Su oundings, Na u al His o y, P o ec-
ion. Ins i ue o Na u e Con e sa ion in Niš, Niš, Se bia. p. 13–21.
Nešić D. 2023b. The main speleological cha ac e is ics o he Ce janska Ca e
Sys em. In: Nešić D, Jo ić D, edi o s. Na u al Monumen Ce janska
Ca e. Su oundings, Na u al His o y, P o ec ion. Ins i ue o Na u e
Con e sa ion in Niš, Niš, Se bia. p. 53–84.
Nešić D, Jo ić D. 2023. In oduc ion. I In: Nešić D, Jo ić D, edi o s. Na u al
Monumen Ce janska Ca e. Su oundings, Na u al His o y, P o ec-
ion. Ins i ue o Na u e Con e sa ion in Niš, Niš, Se bia. p. 9–11.
Pa iće ić D, Za ezalo A, Popo ić M. 2023. Biospeleological esea ch o
Ce janska Ca e and he no he n pa o Kala a a ea. In: Nešić D,
Jo ić D, edi o s. Na u al Monumen Ce janska Ca e. Su oundings,
Na u al His o y, P o ec ion. Ins i ue o Na u e Con e sa ion in Niš,
Niš, Se bia. p. 109–119.
Biologia Se bica 47 53
