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The karst areas in Croatia, part of the Dinarides, have been defined biospeleologically as an area of high biodiversity in world terms, covering 26 thousand km2 in all. In this monograph we give data about the biodiversity of subterranean habitats and their biota primarily from a taxonomic perspective with special attention being paid to regional diversity. The diversified geomorphology, hydrology and climate have resulted in a remarkable range of different underground habitats in Croatia, viz. inland and coastal caves, superficial and deep phreatic networks, interstitial - hyporheic substrates and other infiltration zones, etc. In Croatia more than 500 obligate subterranean species and subspecies are known from caves and interstitial habitats. The majority of subterranean species in Croatia are terrestrial biota. There are 299 troglobiont and 170 stygobiont taxa (species and subspecies) recorded. Most subterranean species have very restricted ranges, almost 70% of them endemic to Croatia. Cave fungi are poorly known. Among them there are several important parasitic troglobionts and troglophilic species on cave coleopterans and troglophilic moths in Croatian caves. Some saprotrophic species could be troglobiotic too but their taxonomic and/or ecological status must be clarified in future research. With a respect to the number of troglobionts, the five largest animal groups are Coleoptera, Pseudoscorpiones, Araneae, Gastropoda and Diplopoda. The beetles (Coleoptera) are predominant. More than 100 cavernicolous beetles (species and subspecies) are known in Croatia. Among stygobionts, the Crustacea predominate. At least half of species described are Crustacea. Among Crustacea, the orders Syncarida and Thermosbaenacea are exclusively stygobiotic. Numerically, among crustacean stygobiotic Amphipoda dominate in caves and Copepoda in interstitial habitats. The only known stygobiotic sponge Eunapius subterraneus Sket et Velikonja inhabits several caves in Croatia. Also, the only known stygobiotic clam Congeria kusceri Bole inhabits a series of caves in Croatia. Other important groups include hydrozoa, planarians (Temnocephalida and Tricladida), snails (Mollusca: Gastropoda) and the only European stygobiotic Chordata Proteus anguinus Laurenti. The high taxonomic diversity of the subterranean biota of Croatia can be attributed partly to the geographical heterogeneity and a rather unique combination of various geological and ecological phenomena.
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AN OVERVIEW
OF THE CAVE AND INTERSTITIAL
BIOTA OF CROATIA
Supplementum
Hrvatski
prirodoslovni
muzej
Croatian
Natural History
Museum
NAT. CROAT. VOL. 11 Suppl. 1 1¿112 ZAGREB December, 2002
PUBLISHED BY / NAKLADNIK
CROATIAN NATURAL HISTORY MUSEUM / HRVATSKI PRIRODOSLOVNI MU-
ZEJ, HR-10000 Zagreb, Demetrova 1, Croatia / Hrvatska
EDITOR IN CHIEF / GLAVNI I ODGOVORNI UREDNIK
Josip BALABANI]
EDITORIAL BOARD / UREDNI[TVO
Marta CRNJAKOVI],ZlataJURI[I]-POL[AK, Sre}ko LEINER,NikolaTVRTKOVI],
Mirjana VRBEK
EDITORIAL ADVISORY BOARD / UREDNI^KI SAVJET
W. BÖHME (Bonn,D),I.GU[I] (Zagreb, HR), Lj. ILIJANI] (Zagreb, HR), F. KR[I-
NI] (Dubrovnik, HR), M. ME[TROV (Zagreb, HR), G. RABEDER (Wien, A), K. SA-
KA^ (Split, HR), W. SCHEDL (Innsbruck, A), H. SCHÜTT (Düsseldorf-Benrath, D), S.
[]AVNI^AR (Zagreb, HR), T. WRABER (Ljubljana, SLO), D. ZAVODNIK (Rovinj, HR)
ADMINISTRATIVE SECRETARY / TAJNICA UREDNI[TVA
Marijana VUKOVI]
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Hrvatski prirodoslovni muzej
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According to the DIALOG Information Service this publication is included in
the following secondary bases: Biological Abstracts ®, BIOSIS Previews ®,
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The Journal appears in four numbers per annum (March, June, September,
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NATURA CROATICA
Vol. 11, Suppl. 1, 2002
HRVATSKI CROATIAN
PRIRODOSLOVNI NATURAL HISTORY
MUZEJ MUSEUM
AN OVERVIEW
OF THE CAVE AND INTERSTITIAL
BIOTA OF CROATIA
Editor
SANJA GOTTSTEIN MATO^EC
This publication is financed by
Ministry of Science and Technology
of the Republic of Croatia
and
Croatian Biospeleological Society
CONTENTS
PREFACE ....................................3
INTRODUCTION ................................3
1. GENERALITIES (by SANJA GOTTSTEIN MATO^EC)...............3
2. KARST AND CAVES .............................4
2.1. The Karst (by MLADEN KUHTA)......................4
2.2. Caves (by BRANKO JAL@I] and ROMAN OZIMEC).............8
3. BIOSPELEOLOGY ..............................11
3.1. The history of biospeleology
(by ROMAN OZIMEC and SANJA GOTTSTEIN MATO^EC).........14
3.2. Biospeleological organisation in Croatia
(by SANJA GOTTSTEIN MATO^EC).....................18
4. THE FLORA OF CAVE ENTRANCES (by SUZANA BUZJAK).........20
5. CAVE FUNGI (by NEVEN MATO^EC)
Ascomycota, Basidiomycota and Anamorphic (Mitosporic) fungi.......21
5.1. Introduction ...............................21
5.2. Major taxa with notes on cave biodiversity and ecology.........22
5.3. Preliminary Checklist of troglobiotic fungi and fungi occuring
both in and out of cave underground in Croatia ............25
6. HYPOGEAN FAUNA.............................27
6.1. Hypogean aquatic fauna .........................27
INVERTEBRATES...............................27
Porifera (by TATJANA BAKRAN-PETRICIOLI)..................27
Hydrozoa (by SANJA GOTTSTEIN MATO^EC).................28
Turbellaria .................................28
Temnocephalidea (by SANJA GOTTSTEIN MATO^ EC)............28
Tricladida (by ROMAN OZIMEC).....................29
Mollusca ..................................30
Gastropoda (by TON]I RA\A)......................30
Bivalvia (by BRANKO JAL@I]).......................30
Polychaeta (by BRANKO JAL@I]).......................30
Oligochaeta (by MLADEN KEROVEC).....................33
Hirudinea (by MLADEN KEROVEC)......................34
Crustacea..................................35
Copepoda (by IVAN^ICA TERNJEJ)....................35
Ostracoda (by SANJA GOTTSTEIN MATO^E C)...............35
Syncarida (by SANJA GOTTSTEIN MATO^E C)................36
Isopoda (by SANJA GOTTSTEIN MATO^E C).................36
Amphipoda (by SANJA GOTTSTEIN MATO^E C)...............37
Thermosbaenacea (by SANJA GOTTSTEIN MATO^E C)............40
Decapoda (by SANJA GOTTSTEIN MATO^E C)................40
Hydracarina (by ROMAN OZIMEC)......................41
Diptera (by SANJA GOTTSTEIN MATO^E C and ROMAN OZIMEC)........41
VERTEBRATES ................................42
Pisces and lampreys (by NIKOLA TVRTKOVI] and MARIJANA FRANI^EVI])..42
Amphibia (by EDUARD KLETE^KI,BRANKO JAL@I] and TON]I RA\A)....42
6.2. Hypogean terrestrial fauna........................44
INVERTEBRATES...............................44
Mollusca ..................................44
Gastropoda (by VESNA ŠTAMOL ).....................44
Crustacea..................................47
Isopoda (by JANA BEDEK and SANJA GOTTSTEIN MATO^E C)........47
Arachnida (by ROMAN OZIMEC).......................49
Myriapoda (by ROMAN OZIMEC)......................59
Insecta ...................................63
Apterygota (by ROMAN OZIMEC).....................63
Pterygota ................................66
Orthoptera (by ROMAN OZIMEC)...................66
Lepidoptera and Trichoptera (by MLADEN KU^INI])..........68
Coleoptera (by BRANKO JAL@I])....................69
Diptera (by ROMAN OZIMEC).....................73
Hymenoptera (by ROMAN OZIMEC)..................75
VERTEBRATES ................................75
Aves (by JELENA KRALJ)...........................75
Mammalia (by NIKOLA TVRTKOVI]).....................76
7. FAUNA OF MARINE AND ANCHIHALINE CAVES ............77
7.1. Marine caves (by TATJANA BAKRAN-PETRICIOLI and PETAR KRU@I])...77
7.2. Anchihaline caves (by SANJA GOTTSTEIN MATO^EC)...........79
8. PROTECTION OF KARST AND CAVES BIOTA
(by SANJA GOTTSTEIN MATO^EC).......................80
9. BIBLIOGRAPHY ...............................81
ISSN 1330-3430
UDK 575.856:551.442(497.5)
AN OVERVIEW OF THE CAVE AND
INTERSTITIAL BIOTA OF CROATIA
SANJA GOTTSTEIN MATO^EC1(ed.), TATJA N A BAKRAN-PETRICIOLI1,
JANA BEDEK2,DRAGAN BUKOVEC2,SUZANA BUZJAK2,MARIJANA
FRANI^EVI]3,BRANKO JAL@I]2,MLADEN KEROVEC1,EDUARD
KLETE^KI2,JELENA KRALJ4,PETAR KRU@I]1,MLADEN KU^INI]1,
MLADEN KUHTA5,NEVEN MATO^EC6,ROMAN OZIMEC6,TON]I RA\A7,
VESNA [TAMOL2,IVA N^ I C A TERNJEJ1&NIKOLA TVRTKOVI]2
1Department of Zoology, Faculty of Science, University of Zagreb,
P.O. Box 933, Rooseveltov trg 6, 10000 Zagreb, Croatia
(e-mail: sgottst@zg.biol.pmf.hr)
2Croatian Natural History Museum, Demetrova 1, 10000 Zagreb, Croatia
3Institute of oceanography and fisheries, [etali{te I. Me{trovi}a 63,
P.O. Box. 500, 21000 Split, Croatia
4Institute of Ornithology, Ilirski trg 9, 10000 Zagreb, Croatia
5Institute of Geology, Sachsova 2, 10000 Zagreb, Croatia
6Croatian Biospeleological Society, Demetrova 1, 10000 Zagreb, Croatia
7Speleolo{ko dru{tvo »[piljar«, Ulica Slobode 28, 21000 Split, Croatia
Gottstein Mato~ec, S. (ed.), Bakran-Petricioli, T., Bedek, J., Bukovec, D., Buzjak, S., Frani~e-
vi}, M., Jal`i}, B., Kerovec, M., Klete~ki, E., Kralj, J., Kru`i}, P., Ku~ini}, M., Kuhta, M., Mato~ec,
N., Ozimec, R., Ra|a, T., [tamol, V., Ternjej, I. & N. Tvrtkovi}: An overview of the cave and in-
terstitial biota of Croatia. Nat. Croat., Vol. 11, Suppl. 1, 1–112, 2002, Zagreb.
The karst areas in Croatia, part of the Dinarides, have been defined biospeleologically as an
area of high biodiversity in world terms, covering 26 thousand km2in all. In this monograph we
give data about the biodiversity of subterranean habitats and their biota primarily from a taxo-
nomic perspective with special attention being paid to regional diversity. The diversified geomor-
phology, hydrology and climate have resulted in a remarkable range of different underground hab-
itats in Croatia, viz. inland and coastal caves, superficial and deep phreatic networks, interstitial –
hyporheic substrates and other infiltration zones, etc. In Croatia more than 500 obligate subterra-
nean species and subspecies are known from caves and interstitial habitats. The majority of subter-
ranean species in Croatia are terrestrial biota. There are 299 troglobiont and 170 stygobiont taxa
(species and subspecies) recorded. Most subterranean species have very restricted ranges, almost
NAT. CROAT. VOL. 11 Suppl. 1 1¿112 ZAGREB December, 2002
Croatian Natural History Museum, Demetrova 1, Zagreb, Croatia
70% of them endemic to Croatia. Cave fungi are poorly known. Among them there are several im-
portant parasitic troglobionts and troglophilic species on cave coleopterans and troglophilic moths
in Croatian caves. Some saprotrophic species could be troglobiotic too but their taxonomic and/or
ecological status must be clarified in future research. With a respect to the number of troglobionts,
the five largest animal groups are Coleoptera, Pseudoscorpiones, Araneae, Gastropoda and Di-
plopoda. The beetles (Coleoptera) are predominant. More than 100 cavernicolous beetles (species
and subspecies) are known in Croatia. Among stygobionts, the Crustacea predominate. At least
half of species described are Crustacea. Among Crustacea, the orders Syncarida and Thermosbae-
nacea are exclusively stygobiotic. Numerically, among crustacean stygobiotic Amphipoda dominate
in caves and Copepoda in interstitial habitats. The only known stygobiotic sponge Eunapius sub-
terraneus Sket et Velikonja inhabits several caves in Croatia. Also, the only known stygobiotic clam
Congeria kusceri Bole inhabits a series of caves in Croatia. Other important groups include hy-
drozoa, planarians (Temnocephalida and Tricladida), snails (Mollusca: Gastropoda) and the only
European stygobiotic Chordata Proteus anguinus Laurenti. The high taxonomic diversity of the sub-
terranean biota of Croatia can be attributed partly to the geographical heterogeneity and a rather
unique combination of various geological and ecological phenomena.
Key words: subterranean biota, biospeleology, stygobionts, troglobionts, biodiversity, karst, cave,
interstitial habitat, terrestrial habitat, groundwater, distribution, Croatia
Gottstein Mato~ec, S. (ed.), Bakran-Petricioli, T., Bedek, J., Bukovec, D., Buzjak, S., Frani~e-
vi}, M., Jal`i}, B., Kerovec, M., Klete~ki, E., Kralj, J., Kru`i}, P., Ku~ini}, M., Kuhta, M., Mato~ec,
N., Ozimec, R., Ra|a, T., [tamol, V., Ternjej, I. & N. Tvrtkovi}: Pregled {piljskih i intersticijskih
organizama Hrvatske. Nat. Croat., Vol. 11, Suppl. 1, 1–112, 2002, Zagreb.
Kr{ko podru~je Hrvatske ukupne povr{ine 26 tisu}a km2, s dijelom dinarskog kr{a, biospele-
olo{ki je odre|eno kao podru~je velike biolo{ke raznolikosti u svijetu. U ovoj monografiji podastiru
se podaci o biolo{koj raznolikosti podzemnih stani{ta i organizama primarno s taksonomskog
aspekta i s posebnim osvrtom na regionalnu raznolikost. Geomorfolo{ka, hidrolo{ka i klimatolo{ka
raznolikost Hrvatske rezultirala je osobitim rasponom razli~itih podzemnih stani{ta, kao {to su
kontinentalne i priobalne {pilje, mre`a plitke i duboke freati~ke zone, intersticij – hiporei~ka zona i
drugi tipovi infiltracijskih zona, itd. Iz {pilja i intersticijskih stani{ta Hrvatske poznato je preko 500
obveznih podzemnih vrsta i podvrsta. Ve}ina podzemnih vrsta u Hrvatskoj su kopneni organizmi.
Utvr|eno je 299 troglobionata i 170 stigobionata. Ve}ina podzemnih vrsta ima vrlo ograni~eno
podru~je rasprostranjenosti, gotovo 70 % je endemno za Hrvatsku. [piljske gljive su vrlo slabo
poznate. Me|u njima je nekoliko troglobionata i troglofila koji su paraziti na {piljskim kornja{ima i
troglofilnim leptirima. Neke subtroglofilne vrste mo`da su tako|er troglobiontske, no njihov takso-
nomski i/ili ekolo{ki status morala bi razjasniti budu}a istra`ivanja. U pogledu najve}e brojnosti
troglobionata, pet je velikih skupina `ivotinja – Coleoptera, Pseudoscorpiones, Araneae, Gastropoda i
Diplopoda. Kornja{i (Coleoptera) su najdominantniji. U Hrvatskoj je poznato vi{e od 100 kaver-
nikolnih kornja{a (vrsta i podvrsta). Me|u stigobiontima dominiraju raci. Gotovo polovica opisanih
vrsta su iz skupine Crustacea. Me|u racima, skupina Syncarida i Thermosbaenacea su isklju~ivi
stigobionti. Broj~ano gledano, me|u stigobiontskim racima skupina Amphipoda dominira u {pilja-
ma, a skupina Copepoda u intersticiju. Jedina do danas poznata stigobiontska spu`va Eunapius
subterraneus Sket et Velikonja nastanjuje nekoliko {pilja u Hrvatskoj. Tako|er jedini poznati stigo-
biontski {koljka{ Congeria kusceri Bole nastanjuje niz {pilja u Hrvatskoj. Ostale va`ne skupine su
obrubnjaci, virnjaci (Temnocephalida i Tricladida), pu`evi (Gastropoda) i jedini poznati stigobiont-
ski kralje{njak Proteus anguinus Laurenti. Velika taksonomska raznolikost podzemnih organizama
Hrvatske pripisuje se djelomi~no geolo{koj heterogenosti i posebno jedinstvenom spoju razli~itih
geolo{kih i ekolo{kih pojava.
Klju~ne rije~i: podzemni organizmi, biospeleologija, stigobionti, troglobionti, biolo{ka razno-
likost, kr{, {pilja, intersticijsko stani{te, kopneno stani{te, podzemna voda, rasprostranjenost,
Hrvatska
2Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
PREFACE
An overview of the cave and interstitial biota of Croatia was assembled from
available literature data, museum collections and from numerous notes made in
various field researches of the present authors.
This Overview provides data on Croatian underground biodiversity and a taxo-
nomic basis for further monographic publications of Underground Biota and ex-
tended activities, such as developing a Croatian Biospeleological Database (CBSD)
and the preparation of The Atlas of Cave Fauna.
As previous review - Croatia - of similar subject matter published in the III Vol.
of Encyclopaedia Biospeologica was printed in rather limited number of copies an
effort was done to make an extended monograph more available to regional and in-
ternational scientists, to applied scientists and society in general. The present Over-
view emerged as the result of this effort as a supplement enlarged and improved
with many new data included (such as: additional records for some species, records
on new species, Checklists, updated systematics and classification of the numerous
groups of organisms, colour photographs etc.).
INTRODUCTION
According to CAMACHO (1992) and HOLSINGER (1994) we used the term tro-
globiont for terrestrial, cave-limited species and stygobiont for aquatic species, which
by definition is a species generally restricted to hypogean waters: in caves and also
in interstitial habitats. The term cavernicolous species (= cave-dwelling species) is
used for some terrestrial species where more precise terms - troglobiont and tro-
glophile were inapplicable because neither troglobiotic nor troglophilous status can-
not be assigned for those species specifically. We expect for some species that are
not limited only to the caves. Also, criteria that distinguish cavernicolous species
are inexact in some cases and the real nature of many cavernicolous species is so
far virtually unknown.
1. GENERALITIES
by SANJA GOTTSTEIN MATO^E C
Croatia, although a small country (56.610 km2of land surface), has a wide diver-
sity of ecosystems and habitats that is reflected in richness of flora, mycoflora and
fauna. Such an abundance is a result of the geographic characteristics of Croatia, as
it is at the border of several biogeographic regions, various types of relief, geologic,
pedologic, hydrologic and climatic types.
However, human beings have also had an influence on the diversity. According
to the natural characteristics, one can divide Croatia into four regions: the Croatian
lowlands, the Croatian highlands, the Mediterranean coast, and finally – the islands
and the Adriatic Sea. The Croatian lowland is a flat area surrounded by rivers Sava,
Nat. Croat. Vol. 11, Suppl. 1, 2002 3
Mura, Drava and Danube, and by the outer Panonian hills (54.4% of the land area).
It consists of numerous regions; of them, Slavonija, the Zagreb region, @umberak,
Pokuplje, Kordun and Banovina are the most important from the biospeleological
point of view. The Croatian Highlands is a part of the Dinarides, with its highest
peak of Dinara (1,831 m). This region is a high karstic belt from mountain to foot-
hill area, with »islands« of impenetrable rocks, karst valleys and river valleys (14%
of the land area). A peculiarity of the region is the great geomorphologic diversity
of the karst phenomena: caves, potholes, rocks, crevasses, canyons and dolines. It
consists of the regions Gorski kotar and Lika. The Mediterranean coast and islands
of Croatia make a narrow coastal belt with islands, separated from the inland with
high mountains (31.6% of the land area). The Croatian coast is the most diverse
coast of the Mediterranean with 1,200 islands and rocks across the 5,835 km of
coastline. It has the following regions: Istra, Hrvatsko primorje with Kvarner and
Dalmacija. The Adriatic Sea is a distinct unit well known for its biodiversity, in-
cluding marine caves with deep water and relict fauna, and for the great depth of
the Jabuka basin of the South Adriatic (RADOVI], 1999).
2. KARST AND CAVES
2. 1. The Karst
by MLADEN KUHTA
In the development of a branch of science nothing is fortuitous; this includes the
use of Croatian karst phenomena terms such as ponor, dolina, polje, jama, etc., in
the international geological vocabulary. The karst relief covers a major part of the
Croatian national territory. The systematic research of karst has a history of over
two hundred years. This region, with its numerous, well-developed and diverse
karst forms, both on the surface and underground, is unique in the world, a »locus
typicus«. Furthermore, geological scientists from all over the world, when referring
to the Dinaric karst refer to it as the »classical karst«.
The Croatian territory south of Karlovac, as well as the Dinaric karst belt that ex-
tends from Slovenia through Croatia to Albania is the region of the classical karst.
The karst covers about 26.000 km2in Croatia, which is 46% of the national territory.
The geology of the karst terrain is dominated by Mesozoic and Tertiary carbonate
rocks (limestone and dolomite) and clastic Paleozoic-Triassic rocks and Cretaceous-
Paleogene carbonate flysch rocks are subordinate.
According to recent analysis (HERAK, 1986; 1991), the Croatian Dinaric region
consists of four paleo-dynamic and paleo-structural belts (Fig. 1). Carbonate rocks
and karst phenomena are situated within the Adriaticum and the Dinaricum struc-
tural units. These are areas of former carbonate platforms, where carbonate deposi-
tion lasted during the whole of the Mesozoic and continued even later. The Epi-
adriaticum is of small spatial extent (tectonic reduction) and consists of basinal and
marginal, predominately clastic sedimentary rocks. The Supradinaricum contains
only sporadic occurrences of karst.
4Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
The geotectonic position is interpreted in the light of the mobilistic theory, i.e., of
plate tectonics. The shaping of the Dinaric fabric was gradual, and caused by the
movement of the Adriatic micro-plate and the deformation of the earth’s crust in
the marginal zone of the European plate (BIJU-DUVAL &MONTADERT, 1977; DEWEY,
1987; MOORES &TWISS, 1995). In the context of global geotectonic movements a
subduction zone was formed and elements of the Epiadriaticum and the Adriati-
cum were subject to underthrust beneath the Dinaricum carbonate mass. This pro-
cess began during the transition into the Paleogene and the convexial currents
Nat. Croat. Vol. 11, Suppl. 1, 2002 5
Fig. 1. Simplified general tectonic map of the Croatian part of the Dinarides with the
distribution of karst. Key to symbols: A – Adriatic carbonate platform (Adriaticum),
D – carbonate platform (Dinaricum), S – Eudynamic area (Supradinaricum), P – Geolog-
ical structures of the Pannonian basin (basic data after HERAK 1986, RADOVI] 1999,
modified by D. BUKOVEC, design by OIKON Ltd.).
caused intensive folding and the formation of nappe structures. As a consequence,
the major part of the Croatian classical karst is manifested as a complex orogenic
karst accumulation. The present structural-tectonic fabric was formed by neotec-
tonic movements, characterised by radial tectonics, accompanied by further disrup-
tion of rock masses and the differential downthrow or uplift of individual blocks.
The hydrogeological relationships in the Dinarides are complex. The vast thick-
ness of the carbonate deposits allows the karst aquifers to be open at depths. Rocks
of lower permeability, the position of which depends on the structural-tectonic fab-
ric of the terrain, sometimes act as lateral or partial barriers, of varying length and
strike, and as such have a major influence on the groundwater dynamics. The prin-
cipal retention zones are formed within the mountain massifs, and the groundwater
drains towards their margins. The type of groundwater flow ranges from fast verti-
cal percolation in the unsaturated zone, or turbulent flow through networks of
subsurface channels or slow siphonal water movement in the deeper parts of the
karst aquifers. The velocity of groundwater flow determined by numerous tracing
tests ranges from several millimeters to tens of centimeters per second, with mean
velocities of 4 cm/s. In the discharge/outflow zones typical karst springs with
large yield occur. One of the largest of such outflows is the Ombla spring near
Dubrovnik with a maximal yield of 154 m3/s.
A large number of submarine springs (vrulja) exist in the coastal region. Their
present position is the consequence of the change in the level of the Adriatic Sea;
the sea level has risen approximately 100 m since the time of the Würm glacial pe-
riod ([EGOTA, 1968). In addition to resulting in the submergence of springs and
older speleological objects, the rise in the sea level has had an effect on specific re-
lationships in coastal and island aquifers. One such phenomenon is the Vrana Lake
on Cres Island. With a surface of 5 km2, and a volume of 220 million m3of potable
water of high quality and a depth of 60 m below the sea level, it is an unique karst
phenomenon in the Dinarides (BIONDI] et al., 1998).
The present karst relief is relatively young in a geological sense. Its development
was favored as a consequence of the tectonic disruption of carbonate rocks, which
is one of the major conditions contributing to the development of karstification. The
karst regions in Croatia are a belt parallel to the coastline. The most prominent
geomorpholohical and orographic feature of the region is the Dinarides with alti-
tudes up to 1800 m (Mt Dinara, 1831 m). The vicinity of the sea and the mountains
results in the formation of condensation and large quantities of rain, with an an-
nual rainfall ranging from 1300 to 2000 mm (from 1600 mm on the coast to 3600
mm inland in Gorski kotar). The corrosive and erosional water action on the frac-
tured carbonate bedrock has resulted in the formation of numerous surface and
subsurface morphological phenomena.
On the surface, in addition to relatively unimportant forms such as karren and
solution pans (kamenica) the most frequent are sinkholes and uvalas. In intensively
karstified regions sinkhole density can be over 160 sinkholes per km2([ARIN &
HRELI], 1984), some of which are several hundred meters in diameter and more
than 100 m deep. The largest morphological phenomena are the poljes (valleys).
Fourteen major karst poljes with an area in excess of 10 km2are located in Croatia.
6Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
Nat. Croat. Vol. 11, Suppl. 1, 2002 7
Fig. 2. The jama-system Lukina jama – Trojama (–1392 m) on Mt Velebit is among the
deepest caves in the world (drawing by D. BAK[I] &V.KU[AN).
The largest polje is the Li~ko polje with an area of 465 km2. Due to the fast infiltra-
tion of meteoric water, the surface hydrography is poorly developed and the flow
is confined to a few rivers. In contrast to this, large quantities of water are retained
in the karst underground, which has formed a rich hydrographic network and nu-
merous speleological phenomena. So far, about 7000 caves have been investigated.
From a morphological point of view, jamas (potholes or pits) predominate, account-
ing for over 70%, while 30% of the morphological phenomena are horizontal caves.
Although most of these explored speleological phenomena are dry features, water
is permanently or periodically present in 1842 of them (GARA[I], 1991). The ponors
play an important role within the karst, on such cave-ponor system, Ðulin ponor-
Medvedica, near Ogulin, with an explored length of 16,396 m, is also the longest
cave in Croatia. Very deep jamas (potholes) have been found in the karst mountain
region. The jama-system Lukina jama – Trojama (–1392 m) and Slova~ka jama
(–1301 m) on Mt Velebit are among the deepest speleological phenomena in the
World (Figs. 2 & 29).
The karst regions, due to the lack of surface water and the small arable soil sur-
faces, and their failure to be incorporated into the transportation network, have al-
ways been sparsely populated. Today, in view of the increasing vulnerability of the
natural environment, this seems to be an advantage. The karst region of Croatia in
most part has been preserved. The developed relief of this region, its extension
through various climatic zones ranging from the Mediterranean type to the moun-
tain type, its high degree of natural preservation and finally the large number of
speleological phenomena, make it an immensely interesting terrain for biospeleo-
logical research.
2. 2. Caves
by BRANKO JAL@I] and ROMAN OZIMEC
There are more than 7500 registered caves in Croatia, and it is estimated that at
least twice as many exist. Annually, 50 to 100 new caves are discovered and ex-
plored. Such a large number of caves and potholes is a consequence of the charac-
teristic karst fabric of the region. In the northern, continental part of Croatia only
small areas of isolated karst are found, and the continuous karst zone lays to the
south of Zagreb. The Croatian karst consists of high mountain ranges and moun-
tain plateaus, but also of low, fluvial and shallow karst. Most of the caves are lo-
cated in numerous mountains and many are still not adequately surveyed. The
caves are developed in limestone and dolomite bedrock, or in combinations of
both. Caves that have developed in marls, flysch and sandstone are very rare, but
some are of importance in world terms. The cave of Piskovica in Istra is located
within flysch deposits and is 1036 m long (Fig. 30) (JEKI] &KOVA^EVI], 1988), and
the [pilja cave near [u{njari in Banovina province has developed in marl deposits
(GARA[I] &KOVA^EVI], 1991).
Most of the caves developed as a result of the combined action of erosion and
corrosion of waters percolating through tectonically predestinated bedrock. Along
the Adriatic coast, abrasion widened these caves and today they are submerged be-
8Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
low the sea surface. Some of the more famous submarine caves are the cave Me-
dova bu`a on the island of Rab and the submarine vertical cave Zmajevo uho near
Rogoznica. Since some of the phenomena are located within the tidal zone they of-
ten produce sonorous effects and the local people call them rikavica (she that roars)
(STRA@I^I], 1954). »Blue« caves are also frequent, such as the Modra {pilja cave on
the island of Bi{evo. Caves of volcanogenic origin have not been found in Croatia.
Artificial caves derive from human activity during the construction of roads, mili-
tary facilities and hydroelectric power plants (BO@I^EVI], 1984). During tunnel and
quarry excavations numerous natural caves were discovered, previously inaccessi-
ble, and sometimes of immense dimensions (such as the Cavern in the U~ka Tun-
nel, 1490 m long; the Cave in the Tounj quarry, 8487 m long (Fig. 27).
Historical cavities include mines, wells, subterranian quarries, catacombs, etc. A
unique facility is the Roman water supply system on the island of Pag, from the
first century AD (BO@I], 1987), as well as the smithsonite mine with several hori-
zons from the 19th century, located in northwestern Croatia (Fig. 28) (OZIMEC, 1994).
The morphological and hydrological features of Croatian caves were described
by GARA[I] (1991).
The most frequent speleo-morphological phenomena are vertical caves – jamas –
(70%), horizontal caves account for approximately 28% and there are about 2% of
combined caves. In a morphological sense there are simple features (20%), knee
type (40%), branching (30%), in several horizons (9%) and other system forms ac-
counting for a share of less than 1%. Among them the rarest are traverses (the tra-
verse Mate{i}eva-Popova~ka cave in the Kordun province). Of all the described
speleological features, 65.8% are dry, while 34.2% are hydrologicaly active. In accor-
dance with their hydrologic function there are permanent (0,7%) springs, intermit-
tent springs (3%), permanent ponors (5%) and intermittent ponors (5%), also there
are estavellas (7.5%) and the most frequent (17%) percolating objects. All of these
caves are situated either within the Black Sea drainage basin or the Adriatic Sea
drainage basin.
A special group of subterranean habitats consists of submarine springs (vrulje),
anchihaline caves, ice caves as well as chambers with elevated CO2concentrations.
The submarine springs (vrulje) are located along the whole Adriatic coast, with the
highest occurrence frequency in the Velebit and the Biokovo channels, and in the
latter the largest submarine spring Vruja is located (ALFIREVI], 1969). Most of them
have small entrances, and some twenty have been explored so far. The study of
submarine spring fauna dates to more recent times (PETRICIOLI et al., 1994). Anchihaline
objects also occur along the Adriatic coast at the area of contact between fresh and
saline water. These were investigated from Istra to Dubrovnik, and they contain
both marine and fresh water faunas, but also some endemic species (SkET, 1986a, b).
Features with elevated concentrations of CO2and SO2have been identified in
Istra and on some Dalmatian islands. Recently in one such jama on the island Lasto-
vo a new taxon from the family Pselaphidae (ordo Coleoptera) was discovered.
Ice caves in Croatia are found in large numbers. In the specific microclimatic
conditions numerous caves contain permanent snow and ice. In these conditions
Nat. Croat. Vol. 11, Suppl. 1, 2002 9
typical troglobiotic fauna is present, often with glacial relicts. Numerous toponyms
exist for these phenomena: Snije`nica, Ledenica, Ledena {pilja, etc.
Large caves and cave systems are found in the region of shallow karst (Kordun
and the Ogulin plateau), as well as in the outer karst belt as terminal parts of fossil
subsurface river channels (N and S Dalmacija).
The deepest potholes, up to as much as 1400 m, are found in the regions of the
highest mountains especially in the Velebit and Biokovo massifs (Tab. 1). A unique
phenomenon is Crveno jezero (the Red Lake) near Imotski, which probably devel-
oped as a consequence of the ceiling collapse of an underground cavern.
Thirty-one caves are protected by nature conservation legislation; 29 are classi-
fied as geomorphologic monuments of nature and two as palaeontological sites.
Fourteen caves are open to the public (Bo@I], 1999). The caves have been the sub-
jects of archaeological and paleontological studies. In this context over 100 caves
have been researched, including the site with the world’s most abundant remnants
of Neanderthal man(M
ALEZ, 1984), Hvar culture and many other finds (^E^UK &
DRECHSLER-BI@I], 1984).
Studies of inorganic environmental features of underground features in Croatia
are rare and mostly deal with temperature and humidity measurements. In terms
10 Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
Tab. 1. The list of the largest caves in Croatia.
Name Region Length & depth (m)
The longest caves
1. The \ulin ponor – Medvedica system
2. The Panjkov ponor – Kr{lje system
3. The cave in the Tounj quarry
4. Veternica
5. The Jopi}eva {pilja – Bent system
6. Muni`aba
7. The Vilinska {pilja – Ombla system
8. Gospodska {pilja
9. Donja Cerova~ka {pilja
10. Slova~ka jama
The deepest caves
1. The Lukina jama – Trojama system
2. Slova~ka jama
3. Amfora
4. Meduza
5. Stara [kola
6. Vilimova jama (A-2)
7. Patkov gu{t
8. Ledena jama
9. Ponor na Bunovcu
10. Jama Olimp
Ogulin
Rakovica, Kordun
Tounj, Kordun
Zagreb, Mt Medvednica
Brebornica, Kordun
Crnopac, Mt Velebit
Dubrovnik, Dalmacija
Vrlika, Cetinska krajina
Gra~ac, Lika
Mali kuk, Mt Velebit
Hajdu~ki kukovi, Mt Velebit
Mali kuk, Mt Velebit
Mt Biokovo, Dalmacija
Ro`anski kukovi, Mt Velebit
Mt Biokovo, Dalmacija
Mt Biokovo, Dalmacija
Gornji kuk, Mt Velebit
Lomska duliba, Mt Velebit
Bunovac, Mt Velebit
Mt Velebit
16,396
12,385
8,487
7,100
6,590
3,700
3,060
3,060
2,682
2,414
–1,392
–1,301
–790
–707
–576
–572
–553
–536
–534
–531
of temperature, speleological features can be classified as ice caves (ledenice) with
temperatures below 1°C, high mountain objects at altitudes over 1000 m (1–5°C),
continental (5–10°C), sub-Mediterranean (10–14°C), and Mediterranean (14 to 20°C).
JAL@I] (1984) performed the first systematic studies of fauna and temperature de-
pendence. The humidity in most caves ranges from 85 to 100%, and only in the
Mediterranean region does it drop below 60%. Some features are characterized by
air circulation, especially in the continental parts of Croatia and on the Velebit mas-
sif, which has led to the creation of toponyms for caves with air circulation (Veter-
nica cave, Puhaljka pothole).
3. BIOSPELEOLOGY
3.1. The history of biospeleology
by ROMAN OZIMEC and SANJA GOTTSTEIN MATO^E C
The first information regarding cave organisms in Croatia dates from the first
half of 19th century. For that period, the typical methods were sampling and giving
a short description of sample material. Numerous findings are marked by the name
of region only, for example, »Dalmatia« – which makes it difficult today to judge
whether they are members of Croatian fauna, because of changes in administrative
regionalization and of frontiers. The troglomorphic snail Lanzaia elephantota
(Muhlfeldt) (syn. Turbo elephantota Muhlfeldt) was described in 1824 as a marine or-
ganism of the Dalmatian coast. More than 100 years later it was discovered to be a
member of the subterranean fauna (KU[^ER, 1933). The troglophilic cave cricket
Dolichopoda araneiformis (Burmeister) (syn. Phalangopsis araneiformis Burmeister) was
described in 1838, most probably, from a cave in the environs of Dubrovnik
(KaRAMAN, 1958). The olm Proteus anguinus Laurenti was found in Dalmacija by F.
CARRARA from Split in 1840 (the source of Goru~ica River by Sinj). On the basis of
this finding, plus a sample from an unnamed cave in the Neretva area, FITZINGER
(1850) described the species Hypochton carrarae Fitzinger,which was later not recog-
nised as being a separate species of Proteus.
During the second half of 19th century faunistic surveys led by numerous re-
searchers began in Croatia. From material collected in Dalmacija by J. ERBER from
Vienna, new species of cave beetles and spiders were described (MILLER, 1861a, b;
KEYSERLING, 1862). In caves of Lika in 1862, the Slovene N.HOFFMAN defined the
first accurate finding of troglomorphic beetles for Croatia (PRETNER, 1973). J.
SAPETZA and J. STIEGLER from Karlovac collected in caves in the Pokuplje area and
discovered new species of cave Coleoptera (MILLER, 1867;HAMPE, 1870). S. BRUSINA
collected material in caves of Kordun and Lika, and noted the first sample of
troglobiont snail in caves (HIRC, 1902), and the first cavernicolous millipedes. In the
Lika region, on July 7th, 1879 the olm was found, for which BRUSINA (1880) defined
a new taxonomic category, Proteus croaticus nom. nud. Faunistic research into
phreatic waters in the environs of Zagreb (VEJDOVSKY, 1882), and various interstitial
biotopes in other parts of Croatia ([O[TARI], 1888) were started. M.PADEWIETH of
Nat. Croat. Vol. 11, Suppl. 1, 2002 11
Senj collected beetles in Lika and the Mt Velebit region and described new taxa
(PADEWIETH, 1891). Unfortunately, due to the commerce with cave organisms, he
hid the locations or named and marked them wrongly, so his data are mostly not
useful. Cave data, and descriptions of the flora and fauna of entrance zones can be
found in numerous papers of D.HIRC (e. g. 1898, 1900). As well as by these re-
searchers, cave fauna was collected by J. SCHLOSSER-KLEKOVSKI and F. ERJAVEC in
continental parts of Croatia, by M. PAVE L in the Gorski kotar region, J. STUSSINER, A.
STOSSICH,andA.VALLE in Istra, A. GOBANZ, E.REITTER, R.STURANY, L.BIRO and A.
LANGHOFFER in the Mt Velebit and Lika region, and by I. NOVAK, N.DAMIN and A.
GOBANZ in Dalmacija. The first systematic biospeleological study was performed by
A. E.JURINAC who collected, in the period from 15th till 31st August, 1883 items of
fauna in five caves in the Kordun region. The results are published in »Prilog
hrvatskoj fauni ogulinsko-slunjske okolice i pe}ina«(J
URINAC, 1887b). Among other find-
ings, a new species of stygobiotic amphipod Niphargus croaticus (Jurinac) (syn.
Eriopis croatica Jurinac) (Fig. 3) is described. This survey earned him a Ph.D. in Jena
(JURINAC, 1888). He also surveyed the cave fauna in the surroundings of Krapina,
including fauna in the wells of the Slavonija region (JURINAC, 1886, 1887a).
In the first half of the 20th century biospeleologic research became much more
vigorous and thorough. In the period from 1902 to 1913, A. LANGHOFFER led sur-
veys on the grandest scale to date. Rich animal material was collected from caves in
Kordun, Gorski kotar, Lika and Zagreb and the surrounding regions. A review of
cavernicolous taxa known up to that time and defined by numerous Croatian and
foreign researchers was published in two parts under title »Fauna cavernarum
Croatiae«(L
ANGHOFFER, 1912, 1915b). From 1912 to 1914 similar research was per-
12 Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
S. Brusina D. Hirc
formed in Dalmacija by U. GIROMETTA. A list of fauna collected in more than 50
caves in Central Dalmacija was published in two works, the latter, entitled »Fauna
cavernarum Dalmatiae«(G
IROMETTA, 1914) being more detailed. V. STILLER (1911–1918)
in the paper »Meine Höhlenexkursionen in Kroatischen Montangebiet«, described fauna
collected in eight caves in Lika, Gorski kotar and the region around Zagreb. D.
POLJUGAN (1915) redescribed the species and described the male of the troglobiotic
spider Parastalita stygia (Joseph) (syn. Stalita gracilipes Kulcz.) (Fig. 4). The fauna of
water tanks and wells was surveyed by L.CAR (1901), and subterranean decapods
were reviewed by K. BABI] (1922). In 1932 the Slovene malacologist L.KU[^ER
(1933) conducted the first fairly large-scale research in subterranean molluscs in
Croatia. Intensive biological research into interstitial underground waters was
started in the 30s by S. KARAMAN who studied the topic till his death in 1959.
Alongside F. VEJDOVSKY and P. A. CHAPPUIS he is considered to be the pioneer of re-
search into interstitial underground waters. Mention should be made of Croatian
cave fauna collectors: in Continental Croatia – I. HOCHETLINGER, M. [NAP, R.
WEINGÄRTNER, V. SLABNIK, F. OPERMAN, B. KOSI], E. RÖSSLER, PEUPELMANN, S.
PLAN^I], J. POLJAK, @. KOVA^EVI], E. PRETNER and V. REDEN[EK, in the Hrvatsko
primorje region (Kvarner) – G. DEPOLI and A. GOIDANICH, and in Dalmacija – P.
NOVAK, S.KARAMAN, Z.KARAMAN, P.MARDESI], S.SVIR]EV, K.MUSSAP and G.
NONVEILLER. Of foreign researchers, a major contribution to knowledge about Cro-
atian subterranean fauna was made by Czech biospeleologist K. ABSOLON who sur-
veyed the Dinarides with breaks from 1901 to 1933, and Croatia from 1908 to 1922.
In his collection named »Biospeologica Balcanica«, the most significant part was ob-
Nat. Croat. Vol. 11, Suppl. 1, 2002 13
A. Langhofer L. Car
tained in Croatia (PRETNER, 1976). As well as his own, material obtained by B. H.
VRSALOVI] (island of Bra~), L.WEIRATHER and U. GIROMETTA (Dalmacija), G. DEPOLI
(Kvarner), and J. OBENBERGER, V.STILLER, R.MEUSELand A. LANGHOFFER (Continen-
tal Croatia) was included in the collection (ABSOLON, 1916). As well as in numerous
papers of his own, the majority of the material is described in a large number of
publications by distinguished European specialists: M. BEZZI, S. J. SCHMITZ, J.
KRATOCHVIL, J. LANG, M. BEIER, B. FOLKMANOVA, C. ATTEMS, J. KOMAREK, A. J. WAG-
NER, C. WILLMANN,etc.G.MÜLLER collected cave beetles in the Hrvatsko primorje
region from Istra to Dalmacija up to World War II. He described numerous new
taxa of beetles and pseudoscorpions, as well as of crustaceans (MÜLLER, 1931a,
1931b). At the beginning of the 20th century, in Southern Dalmacija V. APFELBECK re-
searched into coleopterans. He had a job in Ludbreg (Croatia) before he got a posi-
tion in the Sarajevo Museum. During 30s, the Czechs J. KRATOCHVIL and F. MILLER
started to survey spiders in Dalmacija, while in 1935 and 1936 F. PAX described the
fauna of two caves in Lim Channel, in Istra (PAX, 1937, 1938). R. FRANKENBERGER
and V. BALTHASAR surveyed terrestrial isopods in Dalmacija, and also described
isopods collected by J. KRATOCHVIL (FRANKENBERGER, 1938a, 1939). L. WEIRATHER in
the period 1908 – 1938 visited about 500 caves, many of them in Croatia. The flora
of cave entrances on the Kvarner islands was surveyed by A. HARA^I] and F. MOR-
TON. As well as these researchers, in Istra, Hrvatsko primorje and the Kvarner is-
lands there were also F. J.SCHMIDT, A.WINKLER, G.HORVATH, L.GYLEK, L.MADER, F.
NETOLITZKY, H.KREKICH-STRASSOLDO, E.STOLFA, K.STRASSER, O.CHENDA, CIRKOVICH,
G.RAVASINI , A.SCHATZMAYR, C.LONA, P.PARENZAN andH.SPRINGER.InLika,the
major researchers are: R. MEUSEL, F.TAX and A. WINKLER. The most vigorous biospe-
leologic research was carried out in Dalmacija, where subterranean fauna was col-
lected by numerous coleopterologists, while specialists for other groups are less nu-
merous (NONVEILLER, 1989). The best known scientists from all the groups were: G.
PAGANETTI-HUMMLER, F.TAX, H.KRAUSS, K.A.PENECKE, O.WETTSTEIN, J.STAUDACHER,
14 Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
Fig. 3. Amphipoda, Niphargidae. Niphargus croaticus (Jurinac) (after JURINAC 1887a,
modified).
C.MAYER, H.KREKICH-STRASSOLDO, A.SCHATZMAYR, M.GRABOWSKI, E.DOMBROWSKI,
J.KLIMESCH, J.ROUBAL, F.BLÜHWEISS, A.HOFFMAN, A.WINNEGUTH, G.MESSA, F.
HEIKERTINGER, P.CZERNY, A.WINKLER, J.MATCHA, F.NETOLITZKY, F.NEUMANN, E.
KOHLMEYER, E.STOLFA and P. A.REMY.
In the second half of the 20th century specialist biospeleological research was
conducted. After World War II, cave Coleoptera were surveyed by E. PRETNER from
Postojna, but he also collected other cave material such as millipedes, pseudoscor-
pions, etc. Coleoptera were collected by V. REDEN[EK from Zagreb, while spiders
were surveyed by F. NIKOLI] from Dubrovnik. In 1954, M. ME[TROV started system-
atic research into the interstitial fauna in the Sava River valley and on Mt Med-
vednica (ME[TROV, 1957, 1958, 1960a, b, 1961, 1962, 1964). He discovered and
described a new type of underground habitat, the hypotelminorheic zone –»lebio
-
tope hypotelminorhéique« (Fig. 5), in Mt Risnjak and Mt Medvednica (ME[TROV,
1962). Later, with his colleagues, he did intensive faunistic-ecologic research into in-
terstitial waters next to the rivers of the Adriatic basin, and the rivers Sava and
Drava. Studies of vertical and horizontal distribution and density of population de-
pending on ecological factors, the mutual influence between a polluted river and its
hyporheic zone, etc., were also carried out (ME[TROV &LATTINGER-PENKO, 1981;
ME[TROV et al., 1976, 1983). In the period from 1962 to 1967, and later, supported by
SAZU (the Slovene Academy of Sciences and Arts) the Dinarides was surveyed
systematically by Slovene bispeleologists,J.BOLE, B.DROVENIK, J.MATJA[I^, A.PO-
LENEC, E.PRETNER, B.SKET, K.TARMAN, F.VELKOVRH, etc. Rich cavernicolous mate-
rial was collected, only partially described (MATJA[I^ &SKET, 1969). From 1961 to
Nat. Croat. Vol. 11, Suppl. 1, 2002 15
K. Babi} P. Novak
1988 C. DEELEMAN–REINHOLD from the Netherlands did research into the spiders of
the Dinarides. She surveyed about 400 caves in nearly 30 years of research. Beside
description of numerous new taxa of cave spiders she described taxa from several
16 Gottstein Mato~ec, S. (ed.): An overview of the cave and interstitial biota of Croatia
Fig. 4. Araneae, Dysderidae. Parastalita stygia (Joseph). Key: A – dorsal view, female;
B – ventral view of cephalothorax, male (after POLJUGAN 1915, modified).
Fig. 5. Schematic presentation of the interstitial habitat – the hypotelminorheic zone in
Mt Medvednica, in non-limestone rocks. Arrows indicate the direction of water flow
(basic data after LATTINGER, 1988; ME[TROV, 1962; modified and drawingbyS.GOTTSTEIN
MATO^E C).
other groups (e. g. isopods) (DEELEMAN–REINHOLD, 1965, 1971a, b, 1978, 1983, 1985,
1993). Arachnology research was carried out by J. KRATOCHVIL and F. MILLER until
1977. They produced a monograph on cave spiders of the Dysderidae family and a
review of the cavernicolous spiders of the Dalmatian islands (KRATOCHVIL, 1970,
1978). Since the late 1960s, cave fauna in the whole of Croatia has been collected by
the coleopterologist B. JAL@I]. Among his numerous works on the taxonomy and
distribution of coleopterans he has contributed to the research into stygobiotic fauna
(JAL@I], 1984, 1993, 1998, 2001; JAL@I] &PRETNER, 1977). Since the end of the 70s,
mostly in Dalmacija, cavernicolous fauna has been collected by the malacologist T.
RA\A. From work on material obtained from these two researchers, numerous new
taxa of pseudoscorpions, millipedes, crustaceans and other groups have been de-
scribed (]UR^I], 1988;G.KARAMAN, 1989a;MR[I], 1992; etc.). Faunistic and ecologi-
cal research into interstitial subterranean waters and other underground water
biotopes, focused on the taxonomy and ecology of aquatic isopods, has been done
Nat. Croat. Vol. 11, Suppl. 1, 2002 17
F. Nikoli}
by R. LATTINGER (R.LATTINGER-PENKO, 1970, 1972a, b, 1979). Cavernicolous snail
fauna has been surveyed by H.SCHÜTT and the Slovene malacologists J. BOLE, F.
VELKOVRH, recently by R. SLAPNIK and the Croatian malacologist V. [TA MOL . The
spiders of Dalmacija have been investigated by M. BRIGNOLI, and those in the
Gorski kotar region by F. GASPARO. Crustacean fauna has been investigated byG.
KARAMAN, B.SKET and F. STOCH, and recently by S. GOTTSTEIN MATO^EC.B.SKET has
started research into anchihaline caves (SKET, 1981; 1986a, b, 1996). Millipede fauna
was surveyed by N.MR[I], and the olm by B. SKET, T.RA\A, T.KOVA^EVI], B.
JAL@I] and E. KLETE^KI. Bat fauna has been surveyed by B.\ULI], N. TVRTKOVI]
and D.KOVA^I]. Recently, numerous young researchers have been dealing with
subterranean fauna in Croatia, most of them having contributed to the preparation
of the data for this paper. In 1996 the Croatian Biospeleological Society (Hrvatsko
biospeleolo{ko dru{tvo) was founded; this unites biospeleologists of Croatia. And
without co-opera