I
t
CAVELIFE
LubomirKov66
It is widelyknownthat cavesdo not represent
a
deadenvironment
withoutanytracesof lifeandthat
thereis eventhe chanceto meetsomeanimals,
for example
bats,frogsor salamanders.
onlya few
peopleareawareof the presence
oftinyorganisms
withtheirimmensediversity
andspeciesrichness
in thecaves.
An independent
scientificdiscipline
focusedon
research
intosuchlifeformsandtheirrelations
with
- t ospeteolo8S/
the surrounding
environment
was
established
on the basisof this phenomenon.
We
can meettypicalcaveanimals,oftenwith bizarre
shapes,in tropicaland subtrooical
cavesof the
world.In Europerich underground
faunaoccurs
predominantly
in cavesof the Baikankarstmountains. Intensivebiospeleological
researchin the
last tlvo decadeshasconflrmed
the Dresence
of
Flyi.g bat, Donica Cale. Photo: P, Bella
ru
E
uniquesubterranean
animalsinthekarstregions
of
Slovakia.
lvlanyof theminhabitcavesof the World
NaturalHeritage.
TheSlovakKarstrepresents
an
important
evolutionary
centrcof the subterranean
faunawithintheWestern
Carpathians.
THECAVEECOSYSTEM
Characterization
of thesubterrarea[(cave)ecosystemis necessary
if we wantto understand
the
phenomena
crucial
of the distributionof living
organisms
in caves.Amongthe most important
aDrotrcfactors(agentsof the "lifeless'environment)arelight,temperature
andair humidity.
Light
is a limitingfactorfor the occurenceof vegetation
in the entrancepartsof thecaves.Themajority
of
underground
faunaescapes
fromthe lightto the
dark,a phenomenon
calledphotoptroDra.
However,
it wasobserved
thatlightdoesnotnecessarilyconstitutea limitingfactor.Forexample,
aquaticcrustaceansof the genusNrphargus
regularly
occur
in karstspringswheretheyfind richerand more
accessible
food,but alsomorenaturalenemies.
Caveair temperature
reflectsthe yearlyaverage
valueof the externalenvironment
as a resultof
the latitudeandaltitudeofthegivensite.Thetemperature
range,
whichtheanimals
areabletotolerate,differswiththe species.
Subtenanean
fauna
is ableto withstandlowertemoeratures
{closeto
point)moreeasilythanvaluesabove
the free2ing
20 "C. Air humidityis an importantfactorruling
the distribution
of true cavefaunathat prefers
highvalues(95 100%).Drierspacesarepoorer
for specialized
subterranean
forms.In caveswith
highair humiditydue to percolating
water,such
95
F...r f1.! rr ..re e, ftor ,f..t
t.h.t
l\1 Cti)..|.t at 2DA2
Cave,Niphargirs
as the Don ca or Kresnohorsk6
ln waterpoolsol even
specmefs ntaybe observed
nEreat vely
movingon the wetcaveflooTovercorn
ong0rstances.
may lre
All ivLngcomponents
of the ecosyste..
c assfied as biotic factors.Autotrophicorgansms
ol
are the category
assocatedwth the presence
ght (algae. osses, chensand hgher pants).
organc substances
as a resLr
t ol prro
TheyprodLrce
perfornred
in the r bodies.
Heterotrophlc
tosynthesls
of thesesubstances
and
organisTns
areconsumers
prlmariy
Thrs
on the autotrophs.
so are
dependent
g r o u pi n c u d e sb a c t e r a m
. c r o s c o p if cu n g ia n d
is
of autotrophic
organisms
an mals.Thepresence
parts0Ttrrecavesexposed
linitedt0 the entrance
however,
to the ght on whichtheyaredependent.
theyafeabsentn the lnternacavespacesandthe
foodwebis basedony on the nutrentsI theforin
in the surfacekarst
remnants
originatlng
of organic
horizons.
A s mpllfied
foodwebdominated
bydetri.
tophagous
animas {thosefeedngon deadorganlc
lorms s chat
materalslandcarnvorous(predator)
acteristicoi caves.
96
CAVEVEGETATION
and gorgeshavespecial
Caves,
deepsinkholes.
Therapidchangesn the envTonnrental
vegetauon.
conditonsgve rise to the wellknownphen0nl
enon of inversestfttification of the fora. n the
S ovakKarstthe greatentrancepart of the SLcka
L'€fuon5
'rrib,rr.rs or rr.
Mad$copb
gnlpetdous funei in the xr'snohotsk'
Cave.
Mucormocedo-. colonyof the nicroscopt. funEicovetingbat cuano,
Iadnica
Cave,
evolved
byfailure
ofthecaveceiling,
is
good
phenomenon.
a
example
of this
Counting
from
the surfaceof the karst plateauto the end of
the floor ice lt is 79 m deep.The air temperatureat the bottomof the monumental
entrance
of
ablssalcharacter
fluctuates
onlynegligibly
around
0 'C. Wemayobserve
herewellmarked
vegetation
associations.
Atthemargin
oftheptateau
withdirect
sUnlight,
a eupftotc zorewithpredominanty
higher,
plantsoccurs.ln the twilight- dlsptotc
flowering
At&e - pe.t of the tano twa, Donlca Cave.
zorp - wemayfindferns,mosses,
liverworts,
lichens
andalgaegrowing
ontheprimarysoilor
rocks.
Inthe
lowerpartof thisspaceonlyiceandtalusdeposits
arepresent.
Inthe zoneof full darkness
of cavesthe aptDuczore - vegetation
is completety
absent.
Thatis whyit is notcorrectto classiry
anyplantas
a truecaveorlFnismalthough
prefer
somespecies
conditions
closetothecaveentrances.
Deeper
caves
primitive
areinhabited
by
microscopicfung
{moulds),
light'independent
bacteria
andinsomespecialcases
alsobygreenalgae.
Theyareveryimponant
asfood
sources
for manyanimals.
Onrottenwoodwe may
observefruiting
bodies
of ligniperdous
fungi.Incaves
the colonies
of microscopic
fungisuchas Mucot
mucedomaybe easilyrecognized
on the surfaceof
thebatguano.
plantsaroundlightbulbs,soGroMhs
of primitive
calledtampIto'a,represent
artificialcommunities
incaves.
Lampflora
is composed
mainly
of diatoms,
blueandgreenalgae,mosses
andsometimes
also
of ferns.lt is an undesirable
andunnatural
compc
nent,development
of whichis apparenfly
limitedby
usingof coldlightandis removed
eithermecnan!
callyor chemically.
CLASSIFICATION
OFTHECAVEFAUNA
(rockcreviceinhabiting)
Cavernlcolous
animats
dwellingin subterranean
habitatsare adaDted
to
this envkonment
to differentextents.Thus,severalecologcal
categories
wereouflinedthat parfly
reflectadaptatonsof theparticular
faunalspecies.
proposed
0f the
classifications,
divisionof thesubterranean
faunaintofourcategories
is widelyused
97
at present.Terrestrialanimal forms enteringthe
cavesonlyat random,perhapstransportedbywater
streamfrom the surfaceor falleninto abysses,are
calledtrcgloxenes.Animalsthat may be found in
cavesmorereguladyare classifiedas troglophiles.
They representa broadercategoryinvolvingtwo
ratherdifferentgroups.Subtaoglophlles
are associated with cavesonlyduringcertainstagesof their
life cycleand theydo fot possesanyspecificadap
tationsto thjs kind of environment.
Somedipteran
flies or bats may serve as examplesof animals
dwelling
in cavesonlyduringacertainseasonofthe
yearor duringsomepart of the day.Eutroglophlles
are animals permanentlypreferringthe subterra
neanenvironment
and are welladaptedby external
bodyconstitutionand usuallyalso by internatlife
functions.However,
they may marginallylive and
reproduce
alsobeyondthe caves,as do somemites,
collembolans
and otherarthropods.
0bviousadaptationsto the subterranean
worldare characteristic
of troglobites,which spend their whote tives in
cavesor similarhabitats.
The similarclassification
may be usedfor the
aquaticfauna, wherewe may distinguishstygoxenes, stygophllesand eustygophites.organisms
strictlylimitedto occufiencein underground
water
are classifiedas stygobites.The prefix of these
wordsis derivedfrom the StyxRiver,that in creek
m}{hologyrepresented
the riverthroughwhichdead
peopleenteredthe underworld.We may register
Suchanimalsin the Styxof tl'e DomicaCaVeOrin
the underground
streamsof the Kr6snohorskd
and
Hrusovsk6caves.However,
these expressions
are
notstrictlyconnected
withthe cavefauna.Forexamp e. lhe phreatobltesthat lve In lhe tiny crevtces
amongthe
sandparticles
andgravels
in submerged
sedimentsof the caveand abovegroundstreams
also belongto the stygobites.
lt is not atwayseasy
to allocate
animas occurring
in the underground
to
one of the above-mentioned
categories,
sincethere
are manyintermediate
cases.Thisdivislonof the
subterranean
fauna suggeststhe necessityof protectionof the specificmicro-environments
inhabited
bythis fauna.
ADAPTATIONS
OF THE CAVEFAUNA
Adaptatlonsto life in the slbterraneanspaces are obviouspredominantly
found in animals
classifiedas troglobites.Interestingly
very similar
adaptations
are observedin verydifferentfaunal
groups.we may recognizeadaptationsin external
98
bodyshape(morphological),
internallife processes
(physiological),
behaviour(ethologacal)
and heri
tage (genetic).The morphological
adaptationsare
very marked and often very bizafie, especially
in the warmerregionsof the Earth.Reductionor
completelost of the visualapparatusis characteristic of the majorityof true caveanimals.Theyare
paleor whiteas the consequence
of the reouclon
or absenceof pigmentationin coverbodylayers.
In many arthropodswe may observeapparently
elongated
antennaeand legs.0n theseextremitres an increasednumber of elongatedsensory
setaeis distributed.Thefunctionof theseexternal
bodychangesis to strengthensensoryorgansthat
areessentialfor
lifein complete
dafkness,
namely
sensitivity
to air humidity
andodour(chemical)
signals.Longerlegsenablethe animalto movefaster
whensearchingfor food,sourcesof whichare usuallypoorin the caveenvironment.
Amongthemost
pronouncedphysiologicalchangesthe following
are concerned:
elimination
of the daily rhlthms
in animalactivities,
inhibitionof the metabolism
and abilityto starvefor longerperiods,and lower
numberof eggs laid. Ethologicaladaptationsare
connected
with the mode in whichthe animals
colonize
underground
habitats.
THEUNDERGROUND
ENVIRONMENT
AND CAVEFAUNA
Terrestrial
subterranean
environment
Theterm"caveecosystem"
wasusedfora long
timeforthepatsofthenatural
underground
spaces
reachableby man.However,
cavesare not the only
environment
inhabitedby the cavernicolous
fauna.
The terrestrialunderground
environments
may be
dividedintothreemaingroups:
1. naturalcaves
2. epikarst
3. artificialcavesmadeby man
Naturalcavesusuallyrepresentspacesthat may
hide even larger animals bats or amphibians.
The presenceand distributionof the animalsin
them is determined
by variousecological
factors.In
manycaseswe maydistinguish
severalwelldefined
associations
of the animalsoccurringin caves.Cave
entrances
aremarkedly
influenced
bytheexternalclimate.Theseusuallywetterand protected
sitesserve
as refugesfor some bird species(pigeons,owls,
passerine
birds).Someanimalsnestor staycloseto
the entrances,
for example,
dormice,frogs,salamanders,snails,insectsand otherarthropods.
Fromthe
entranceto the beginningof the aphoticzone,cave
wallsand ceilingsare inhabitedby a heterogeneous
communityof animalsdefinedas pa etal fauna.
Theyrepresentoccasionalcaveanimalsusingcave
entrancesonly as temporarysheltermainlyduring
summer and winter (troglophiles):
dipterans,butterflies,trichopterans,
spidersand opilionids.the
compositionof such communitiesis dependenton
the size,orientation
and altitudeofthe entrance,
fur,
theronthestructureof the entrancepartsof the cave
and on the seasonalchangesof the above-ground
climate.Internalpartsare usedby bat coloniesthat
produceexcrementsrich in nutrientsduringtheir
activityphase- guano,an importantfoodsourcefor
a widespectrumof invertebrates.
Depending
on the
degreeof association
with such micrcenvironment
theymaybe classifiedeitherguanophilesor guanc
bltes.0n the pilesor smallerguanoaccumulations
we mayrecognize
earthworms,
mites,collembolans,
woodlice,millipedes,beetlesand dipterans.Thus,
the populations
of suchanimalsare primarilyassc
ciatedwith the presenceof bats dependingon the
amountandthe ageofthe guano.
obligatecaveformsofthefauna livein the deeper
partsof cavesin completedarkness,stabletemperaturesand highair humidity.Wemayobservethem
onthesurfaceof sedimentsandspeleothems,
under
rocks,on the organicremnantsor on the surfaceof
the waterpools.In mostcasesthey representtiny
animalsthat maypenetrateintoeprkarstconsisting
of the systemof interconnected
microcavernstn
fragmentedbedrock.lt is situatedin the upperzone
of the karstmassifunderthe superficialsoil layers.
This environment
type links caveswith the rest of
poorly
the karsticsystem.lt has been biologically
exploredsinceit is hardlyaccessible
to man.
In the tefiitoryof Slovakia,the third underground
- the
environmenttype
hasbeenpoorlyunderstood
artificialshafts and adits of mines.The biological
observations
in that environment
mayadd consider,
ablyto understanding
of the distributionand evolu,
tion of cavefaunain the country.
Theaquaticsubterranean
environment
This environment
type may be classifiedinto
two maintypes:waterscirculating
in porousrock
(alluvialterrains)and watersflowingin disconnected,predominantly
karsticrock.Bothtypesare
inhabitedby differentfaunawithspecificadaptationsto lifein theseenvironments.
Animalsofthe
first type are phreatobites,fascinatingformsthat
havebeeninsufficiently
knownup to now.Their
bodiesare considerably
modifiedenablingthem
to live in the interstitlal envlrcnment,namely
the micro-spaces
betweensand and gravelparticles.The crustaceanBarhynellanatans ts a
goodexampleof suchfauna.Watercirculating
in
present
fragmented,
predominantly
erodedrockis
anthe karst regions.We recognizetwo hydro,geologicalzoneswithinthis environment
type.The
vadosezorreincludesthe upperpart of the karst
massifwithmainlyverticalflowof the water.Here
lhe faunais welladapledto Ife in smal'crevices,
it may occasionally
appearin water pools,too.
Theyare chieflytiny animalswith elongated
and
planarians,
flattenedbodies,suchas
nematodes,
oligochaete
worms,andfromcrustaceans
cyclops
good
and amphipods
serveas
examples.
Wemay
also detectlargerforms in such habitats,like
amphipodcrustaceans
of the genusNiphargus
the lengthofwhichcanexceed2 centimetres.
The
phreaticzone coversthe part of the massifwith
mainlyhorazontal
waterflow.Thezonels usually
populated
by a higler numberof fauna,specres.
This is the kingdomof largercrustaceans,
for
example,
lhe dbovementroned
Nipha/gus.
DIVERSITYOF CAVEORGANISIVIS
fhe blodivebiaylspeciesdiversltyof the organisms)0f cavesprimarilydependson the abioticfactors of the environment.
Heterogereityof the given
habitatis alsoveryimportantsincea widespectrum
of micro-habitats
createsrnorepossibilities
for colonizationbyvariousformsof organisms.
Thesizeof the
karstareais crucialin this respectsincelargerareas
arenormallyinhabitedbya highernumberofspecies
of organisms.
Thisis alsothe caseof the vast plateausof the SlovakKarst.Thebiodiversity
of caves
also dependson stabilityof the environment.
Less
stablecaveswithratherfluctuating
conditions
during
the year creatediversemicr+habltatsthus populatedby faunalcommunities
richerin species.Food
principal
accessibility
is oneofthe
factorsregulating
biodiversity
of the subterranean
habitats.Eltrophrc
spaceswith sufficientsupplyof organicmatterare
rich in faunaand micrcorganisms,
for example,
the
plant
bottomsofthe entranceshaftswithfallen
remnants.or cavepartswtthgualo deposits.
Howeve(In
thesepartstrogloxenes,
troglophlles
or guanophiles
prevail.Ongotroptrrc
cavesare poorin organicmate
populared
rialsand are
by faunapoorerin species.
0n the otherhand,suchcaveshosthighernumberof
trogobiticanimalscompared
to the preceding
case.
99
100
ORIGIN,EVOLUTION
AND DISTRIBUTION
OFTROGLOBITES
THEPECULIARITIES
OF LIFEIN CAVES
OFTHEWORLDHERITAGE
Whyand how did animalspenetrateinto the
subterranean
world?Howis it possiblethat they
wereableto adaptto sucha harshenvironment?
Thereare two main theoriestrying to answer
these questions.Both considerevolutionfrom
ancestorsoriginatingin abovegroundhabitats
(soil,surfacewaterstreams).
0neofthemregards
caveformsof faunaas remains- relrctsof the
preceding
geological
periods,so-called'living
fossils".Forsuchformsthe cavesarethe refugeg
againstunfavourable
externalconditions.
Based
on this theory,surfacefaunawasforcedto look
for sheltersdeeperunderground
withthe startof
periods
the long
of the coldand dry Pleistocene
climateroughly
2 millionyearsago.Thesecond
theory,
ontheotherhand,considers
activecolonizationof cavesbyfauna,a process
thatcontinues
up till the present,However,
not all animalsare
permaable to populatesuch an environment
nently.In somecasesthis wasthe wayto the evo,
lutionof a highernumberof relatedformsfrom
one ancestor
withina relatively
smallarea,the
processcalledadapttreradtatlon.
evolutionary
Thisis the caseof carabidbeetlesof the genus
Duvalius
andits subspecies
in the territoryof the
Western
Carpathians.
Colonization
of the subteranean
habitatsby
passively
thesurface
fauna,either
astheconsequence
of unfavourable
climateor byactivepenetration,is accompanied
by speclatron,that is
bythe evolutionof newspecies.,soratlonof the
premise
spaceor areais an unavoidable
for the
evolution
of newspecies.
Populations
oftheoriginalspecies
isolated
in subterranean
spaces
for
a relatively
long time (tens-or hundred-thousandsof years)becomedifferentfrom
the ancestorduringsuccessive
adaptations,
species
being
product
process.
the final
of this
In manykarst
regionsthe populations
of the sane species
are isolatedfromthe surfacepopulations.
l\4any
examples
of the originof independent
species
withinonecavesystemareknown.
Suchunique
forms,limitedin theiroccurrence
to a relatively
smallarea,are callederderrrcs.Theyrepresent
the rarestand most vulnerable
formsof life.
Wewillconcentrate
on themmorethoroughly
in
part involving
the following
the peculiarities
of
the cavelife withinthe WorldHeritage
sitesin
Slovakia.
Inthischapterthe biotaof the bestknowncaves
is stressed:Domica,Gombaseck6.
Krdsnohorska
andJasovsk6
caves,
Silickil'adnica
Cave.ochtinska
Aragonite
Caveandthe Dobsinsk6
lceCave.
Wewill
alsomention
somepeculiar
animalsof othercaves
or abysses
of the SlovakKarstthatarethe partof
theWorldHeritage.
In recentyearsa widespectrumof mlcro-orgaF
lsns wererecorded
in the cavesof the Slovakf€rst.
Green
algaearetypic€
| components
ofthelampflora,
withStichococcus
bac,r,as,Chlorella
minutissima,
ol
species
of the genusDesmococcus
sp.l\4icroscopic
fung areextraord
inaryintheirdivelsity.
Theyserveas
the important
foodsource
for manysmallerinverte
brates,
suchasmitesandcollembolans.
Thespecies
of thegenusMucordominates
amongthefirstcolq
nizersof bat guano.Entomopathogenic
Beauveth
brcngnianii
c@at€svatia
blemacroscopic
colon
ieson
limestonerocksandsinietlillings,Pidoplitchkovie
a
tenlcolaisolatedfrom isopodexcrements
is a very
raresaprotrophic
microfungal
species.
Theunicellular
animals- protbb (Protista)
have
not beenexplored
sufficiently
in the cavesof the
Slovakiayet, so we havealmostno information
(Turbellaria)
on this animalgroup.Planarlans
of
genus
the
Dendrocoelum
are ratherrareamong
the aquaticfauna,theyare predators
feedingon
Planaia. wm of the genLs Dendro@lum frcm the Farbeni ponor
Cave,Photo:J. Stankovld
tiny invertebrates,
including
deadbodies.Among
the smallformsof annell& oligochaete
worms
(Annelida)
that live in the claysediments
of the
underground
streams,
or in decaying
organicmat+
rialswithintheterrestrial
habitatsno uniqueforms
yet.Largerannelids- earthhavebeenobserved
wormsrepresentpermanentcomponents
of the
faunaof manycavesbeingpreferably
associated
withthe presence
of wood.Theyare ableto consumegreatamounts
of claysedimenttogether
with
organicremains.
Inthe Domica
Caveseveral
earthwormspecieshavebeendetected,
but notspeciaf
izedcaveforms.Remarkable
tracesof theiractivity
arethecasts(earthworm
excrements)
in manysites
in the cave,especially
in the riverbedof the Styx,
oftenevenon the sintersurfaces.Leacheshave
occasionally
beenregistered
in the underground
Ior
waters, example,
thesurfacespecies
Haemopis
sarguisuga
inthe Domica
cave.
(lvlollusca)
Molluscs
mayberatherabundant
inthe
glaberis an
cavesof Slovakia.
Thesnailoxychilus
abundant
inhabitant
of the entrance
micro-habitats.
In karstspringswe maymeetthe snailSad/eriana
- anendemic
pannonica
oftheSlovak
andAggtelek
Karst,
andBiikkl\4ts.
inHungary
aswellasEythinel/a
austriaca.
A sofar undescribed
species
of cavesnail
of the genusHauffenra
has beenreportedfrom
severalkarstspringsof the Silickiiand Plesivskd
plateaux.0n
thesurface
of banksediments
ofunder
groundstreams
tinybivalves
of the genusPisid,um
maybe observed
by the nakedeye,theyare very
abundant
in the river-bed
of theeiernyBrookof the
Iadnica
Silick6
Cave.
Parp,iqrade
Eukoenenia
spelaea(bodytenEth7.5 nm,
Phato:G.C.iah4hut ahdL. t<avAa
several
caves,including
the Domica,
Gombaseck6,
Kr6snohorsk6,
Hru5ovsk6
and Jasovsk6
caves.lt
primitive
represents
anevolutionarily
arachnid
form
demonstrating
typicalcharacteristics
of the obligatecavedwellers.
A longandsegmented
flagellum
on theendof theabdomen
is oneof thepeculiar
group.The
morphological
featuresof thisarachnid
territoryof Slovakiais the northern-most
region
withthe presence
of palpigrades
in the world.The
modeof their reproduction
andfood preferences
- predatory
remainunknown.
Theotherarachnids
pseudoscorplons
(Pseudoscorpionida)
usuallydo
Snat Oxychilus
Elaberlbody/enEth74 nm), KresnohoBkeCaee.
Thehighestspeciesdiversityof caveinvertebrates
is concentrated
in the animalphyllumArthropoda.
Severaluniqueanimalsofthe WorldHeritagecaves
belongto the arachnids(Arachnida).
PalpiErades
(Palpigardi)have been registeredin Slovakiain
one sDecies- EukoeneniasDe/aeadistributedin
Beudosco,pionNeobsilm (Slothrls) slovacun (body teh4th4 nn).
701
Har€srneh lschyrcpelis manica\a often pdet?tinq
into the cavet Photo: P, lupt4iik
deepu
Sptder Meta menadi, inhabifant al the caw ennanc*. lhoto: p tuptaaik
Ircglobit c spider Pod.omma p@lundom,Oom,€ Car€,photoj I Srankolrd
Oribatd nite Pantelozetes€vari.us (body tength O.5 ntu).
Photo: G. CsiAtu'rova and P, LuptAiik
702
not livein caves.Troglobite
(Blothfus)
Neobisium
s/ovacum,characterized
by stronglyelongated
pincerpedipalps,
havebeenobservedin many
abysses
of the Ple5ivsk6
Plateau,
and in a limited
numberof caveson the HornivrchandZadielska
plateaus.
Thespecieswasdescribed
byJ. culiaka
in the 1970s.Thiswasan imoortantlandmark
in
Slovakbiospeleolo$/
sinceit represented
one of
the mostvaluable
confirmations
of originaltroglobiticfaunain the Slovakcaves.Fromthe haruestm6n(Opiliones),
,schyropsalis
manicata
is a sma er predatory
arachnidwith remarkable
chelicerae
inhabiting,
for example,
the DomicaCave.Spiders
suchasMetamenardiarefoundin large
{Araneae)
numbers
in caveentrances,
wheretheycatchflying
insectsin theirwebs.Somesmallerformsof spidersareadapted
to lifein the deepercavespaces
wheretheylookfor theirprey,mostoftencollembolans,mitesor insectlarvae.Ihe mostpeculiar
of thembelongto the genusPonhomma
of which
P. profunclum
is considered
troglobiticspecies.
Thespecieswasdescribed
fromthe Domica
Cave
whereit established
stable pooulations.
Mltos
represent
an arachnidgroupwithstrongly
diversified communities,
Oribatidmitesconsumealgae,
fungiand rottenorganicmalerials(wood,guano).
Pantelozetes
cavaticusis closelyassociated
with
caves.lt wasfirstdescribed
fromthe Certova
diera
CavenearDomica.
lt is a eutroglophite
miteoften
occurring
in closevicinityto bat guano.oribatid
miteDamaeuslengensdotfi
is a largerformwith
elongated
legs,an inhabitant
of the DomicaCave.
ln the samecave,the uropodid
mitesWoobovella
advenaand Cyrtolaelapschiropterceare among
the principalconstituents
of the bat guanoassociations.Predatory
mitesare represented
by the
eutroglophile
form Parasitus/onbatus,a regular
inhabitant
oftheSlovak
caves,
or rhagidiid
mitesof
the genusRobustoche/es
registered
in the oomica
Cavein recentyears.
(Crustacea)
Ctustacean3
are predominantly
aquaticanimals.Theterrestrial
eutroglophile
isq
podeMesoniscus
gran€er(lsopoda)
is anexception
populations
establishing
abundant
in the Domica
and Jasovsk6
cavesas the importantconsumer
of the bat guano.Theisopodewasdiscovered
by
Slovaknativel. Frivaldszky
in 1865in the Earadla
CavenearAggtelek,
a partof the Domica-Baradla
caveslstem.Anongaquatic
crustaceans
Niphargus
tatrensis
is worthmentioning.
lt is distributed
inthe
majorityof Slovakcaveswith standingor flowing
waters.
Asa symbolofstygobitic
animats
it js abun-
/sopodMeso.lscusgEniger Photo:). Stankovia
sq€obnt c/usraceanN pha/gustanenss. PhoroiJ. slankovid
F{r@d,'La l)]edr
in the St)r(River(Domica)where
dantespecially
the
individualsare notablefor their unusualsizeof up
to 4 centimeters.
Richcommunities
of cladocerans
penetrateinto the DomicaCaveby the
(Cladocera)
flow of the DomickyBrookespeciallyduringhea!ry
rainfalls.However,in the undergroundwatersin
Slovakiano obligatecavespecieshas beendiscoveredyet.Ofthe relatedcopepods(Copepoda),
three
speciespopulating
the StyxRivermaybe classified
as phreatobitic:
Acantocycrops
venustus,
Diacyclops
lanEuidoidesand Microcyclopsrubellus.Bathynella
groupof crusnatansbelongingto the Bathynellae
livingin the microspaces
taceansis a phreatobite
of submergedsedimentsof the underground
river'
beds.lt maymarginally
occurin lakessuchas in the
BielaHallof the Dobsinsk6
lce Cave.
The pale and blind mllllpede{Diplopoda)
of the
genusIyphloiuluswas detectedin the Domicaand
Gombaseckd
cavesthus representing
one of the
mostvaluablediscoveries
of cavefaunain Slovakia
in recentyears.lt is olr largesttroglobitewith a
bodylengthof 2.6 cm and 147 pairsof leg.ThemiL
lipedeBrcchychaeteuma
bradaeis a typicalinhabitant of the DomicaCave.TheDobsinsk6lce Caveis
inhabitedbyAiiorhiscosoma
sphinx,a cavernicolous
milliDede
endemicto the WesternCaroathian
caves.
Millipedesare importantconsumersof decaying
organicmaterialsdepositedin caves.
Collsmbolans(Collembola)
are amongthe most
abundantarthroDods
of the subterranean
habitats.
Several
speciesin our cavesexhibita highdegreeof
Theypreferably
feed
adaptation
to that environment.
on fung coveringthe surfaceof decayingorganic
remains,or eventhe surfaceof the speleothems.
HypoEastturacassaeEranulatadobsinensisis an
inhabitantof the inversed
entrancemossvegetation
of the DobSinsk6
lce Cave.lt is a glacialrelictsurvivgaciation.Several
ing he.efromthe lastPleistocene
troglobiticforms of Collembola
may be recognized
within the generaDeuteraphorura,
Pseudosine//a
Barh'+lla nM6(body[.nErn L5 mm) ry'ofo r Hr-dec
hoElabitic mi ipede of the gen!s Typhloiulus,Domica Cave.
ltaElobitic @llenbolan Pseudosinellaaggteekiensis (body,ength 2 mnl,
103
(Coleoptera)
and Anhopalites.Deutercphorurc
krctochviliis
BeetleE
areamongthe mostremarDob5insk6
lce
a typicalcollembolan
of the
Cave.In
kablecavernicolous
arthropods.
E, csiki and E.
the cavesof the SlovakKarstthe speciesis replaced Bokorprovided
the first information
on the cave
bVa closely
related
undescribed
one.Deuteraphorufa beetlesof the SlovakKarstat the begnning
of the
quedius,
schoenviszlryi
ocurringin the Gombaseck6
Caveis
lastcentury.
In cavesthegeneraTrechus,
a troglobite
as wellandan endemic
species
of the
AthetaandCholevaate the mostoftenrepresentaSlovakandAggtelek
Karsl Pseudosinela
agtelekientivesandthe majority
ofthemmaybeclassified
as
is
sis anendemic
species
distributed
inthesame
area
troglophiles.
Inthe Domica
cavelrechusaustriacus
whereit isoftenabundant.
Inkarstregions
ofthecenand ouediusmesomelinus
createratherdense
tralWestern
Carpathians
it is replaced
bytheclosely populations.
ThegenusDuvaiiuscoversthe only
relatedP pacnithat livesin the Doblinsk6lceCave, obligate
cavebeetles
in ourcountry.
Thespecies
of
too.ThegenusArrhopaiites
includes
several
obligate thegenusareendemics
Western
Carpathian
of the
c€veformsinourterritoryEutroglophilous
A.pygmae
mountain
rangespreferably
inhabiting
the superfi(deeper
partsofthe
usis a frequentspecies
ofthecavecommunities.
0n
cialunderground
environment
the otherhand,troglobiticA. aggte,ekiensis
is distristonydebris),in cavestheyare rarer.Interestingly,
butedina limited
number
of undistulbed
caves
where two differentspecieslivetogetherin the Jasovska
theymayberathernumerous,
asintheoertova
die€
CavenearDomica
andin Kr6snohorsk6
lt
Cave. is an
endemic
species
of the SlovakParadise,
Murenska
Plateauand SlovakKarst.Duringbiospeleological
research
in recentyealssomeunknown
collembolan
species
havebeendscovered
suchasa newspecies
of the genusMesaphorura
in the KrSsnohorski
Cave
Neelides
Megalothorax
in
orthat of
and
theJasovske
Cave.
Suchdiscoveries
confirm
howlimited
ourknowlpopulating
edgeof suchsmalllivingcreatures
subterranean
habitats
stillis.
Staphylhid beete Quedi6 msmeli.us,
Co enbota on su.faceof a watetpoot.Kt6shohotsk'c.ave(bodyle gth
of snar/erMegalothorax
sa o 5 hn). F,hoto:.1.stahkoia
TtoCtobiticcoltenbolah Anhoparitesaggterekiensis{body lencrh 2 mm),
104
Oomie CrE PlDto: P tuptidiik
ltoCtobitb catabid tuetle ol the ge.us Duvalius, (.dsnohoEld
CaE,
Caue- D. hunEaicusandD. bokori.Dlptsranfllss
(Diptera),
buttertlles(Lepidoptera)
andcaddbflle3
(Trichoptera)
represent
an importantpart of the
parietalfaunawithseasonal
emergence
in caves
(subtroglophiles).
Of dipteransTriphleba
anticola
is a stablecavedwellerofthe Jasovskd
Caveassociatedwiththe batguanoaccumulations.
ThebutterffiesScoliopterr4libathix andTriphosadubitata
are partof the typicalfaunaof the entrance
communities,beingnumerousespecially
duringthe
regarded
partof caves.In
themas an inseparable
conditions
with little knowledge
therewerea lot
of legends
and reportsaboutthem,Nowadays
we
knowthatbatslookforcavesasoneofthepossjble
shelters
period,
duringtheirresting
eitherduringthe
dayinthewarmmonths,
or inthewinterseason
duringrrlDemat
or (winterresting
stage).
Theyarcone
of the fewgroupsof warm-blood
animalsthatstay
duringthe winterin a stageof truewintersleepin
whichbloodcirculation
andrespiration
areseveral
Aude.ft Scoliopteryx
libaMr inhabiart of the caveenttancT.
timesslowerandbodytemperature
dropsto 10 1 "C. Batsusually
fall intowinterrestin November
(Amphibia)
Amphlblans
guestsof the
areregutar
asdobutterflies
andbeetles,
theirmainfoodsource.
DomicaCave- salamanders
and frogspassively Incavestheyareratherprotected
against
theirnatufloodedinsidethe caveduringheavierrains.The
ral enemies(manens,
weasets,
owls).Batsusually
alarmingly
coloured
Salamandrc
salamandra
hibet- catchinsectsduringflight.orientation
in the dark
natesin numerous
aggregations
in the entrance spaces
is enabled
byechorocat
or, namelythetype
passageof the Gombaseckd
cave.Of mammals oforientation
basedonthereflection
ofsupersound
(lvammalia)
smallformsof rodents,
dormice,
weawavesthat are emittedin regularintervalsand
selsand martensare regularguestsof the caves aresubsequently
received.
Thusthe baGareable
withan unusualsensefor orientation
in complete to distinguish
barriersor preyduringflightin the
darkness.
Themartensare important
predators
of
darkness.
Species
of thefamilyRhinolophidae
emit
batsin caves.Batsarethe bestknownand most
signals
bynose,thoseofthefamilyVespenilionidae,
popularcavemammals.
FromancienttimesDeoole on the otherhand,bymouth.Thereflected
signals
arereceived
bythe ears.Thesoundsaresituated,
depending
on the particularspecies,withinthe
extentof 15 - 115 kHz,so theycanonlyparflybe
heardbythe humanear.Echolocatjon
signals
emitted bybatsareusually
adapted
to flightconditions
and spacestructuresthat considerably
impede
- oevtces
theiridentification
using"BAT"detectors
enabling
analysis
ofthe batsounds.
Danubecated .ewt - IlituG dobrogtcls,Odne Cave.fto0oj p Llpnatt
Common frog - Ra.a temporatiat Doni@ Caw
ronSJeSged
bar ri.l - rxods v6per| ionisrna,el, exte/ra,pa€site of
105
Guano pile in the Domica Cave.Photo: A. No6ko@
Gteater hoseshoe bat in detait, Photo: P.LuotAeik
Auruhn coloty af the EtearethoAeshoebat - Rhinoloph
us
feriumequnum.PholoiP Llpbaik
Considering
the caveanimalsit is impossible
to
omit the surfaceparasitesof batsthat are sucking
their blood- ticks,bugs,fleasand tiny wingless
fliesof the familyNycteribidae.
Sulfacemammal
speciesare the other hostsof theseparasites.Bat
tick /xodesvesperti,ionisis the exceptionsince it
is unableto liveoutsidethe cavesso that it is an
obligateparasiteof bats. Similarlyto other cave
elongated
legs.
animalsit hasstrikingly
The DomicaCaveis well-knownfor its bat colo
nies. In the cave we may observelarger guano
pilesand guanopots - smallerspheroidalcavities
evolvingon the sintersas a resultof the actionof
acidicbat excrements.
In the Domicacavesystem
16 bat specieswereregistered.
Themostimportant
horseshoebat Rhinolophus
is the lvlediterranean
euryalepopulatingthe caveduringthe wholeyear
and establishinglarge coloniesof up to 2,000
106
individuals.The caveis one of the northern-most
distribution
sitesofthis batsoeciesin Eurooe.Inthe
neighbouring
CertovadieraCave,partofthe Domica
system,the southern bent'wingbal Minioptetus
schreibersii
wasveryabundantin the past.lt istypikarstareas
cal ofthis speciesthat in moreextensive
it inhabitsonly a few caveswith favourableambient conditions.lt is an extraordinary
socialspecies
livingin coloniesduringthe wholeyearround.In
the above'mentioned
caveit createdcolonieswith
300 - 9,000 individuals.
Thickguanodepositsare
the traces of their activityin the cave. However,
since 1989 hibernationof this specieshave not
beendetected,the possibleconsequence
of instal
lationof an unsuitablegate on the caveentrance
in the precedingyears.The JasovskdCavehosts
a highnumberof bat species.lt servesas the most
importanthibernationplacefor the greaterhorseshoe bat Rh,noioohus
fenumeouinumin Slovakia
with250 individuals.
TheDobsinsk6
lce Caverepre'
sentsoneofthe mostimportanthibernation
sitesof
the prevailingly
cold adaptedforestspeciesMyots
myotisa^d M. brardtii in CentralEuropewith 422
individualscountedin 1999. In glaciatedpafts of
the cavea populationofthe northernbat Eptesicus
nilisonidominates.
BASICMETHODS
OFCAVEFAUNA
RESEARCH
isdeposited.The
meshis putoverafunnelfixed
into
a stand,a vesselwith
conservation
liquidissituated
underthe funnel.With progressive
dryingof the
Caveinvertebrates
are most often collected substratethe
faunatriesto escape
bymovingdown
directlyby visualsearching
usingpincers,brush towards
themeshandfallingthrough
it intotheves.
or gxtausto.,the deviceby whichwe are ableto
selwhereit is subsequently
fixed.
sucksmallerindividuals
intoa vesselwithfixation
Aquaticinvertebrates
are collectedwithtweezers.
iiquid.Baits(pieces
of meat,cheese,
fruits),usually Smaller
formsmaybe co ectedbyfiltering
of water
place
deposited
on the
for longertime,areusedto
throughthe planktonic
net.Thedeviceconsists
of
attractfauna.Themajorityof smalleranimalsare
softsilkclothputon a circular
framewitha handle.
Thenet progressively
narrowsdownintothe neckof
a smallplasticvial.Inthe vadose
zonethewateris
filteredwithsucha netfromwaterpoolsandperce
latingwater.
Inlargerlakesand
undergroundstreams
theplanktonic
netis usedjn thesamewayasin the
catching
of waterfaunain aboveground
habitats.
In suchmicrohabitats
the densityof the faunais
ratherlow'so collecting
withthe nethasto bedone
repeatedly
in different
hydrological
conditions
during
thet€ar lt is alsonecessary
to watchthesurface
of
cavebottoms
andwallsfor theaOuatic
tuuna.Baits
areusedfor attraction
andsubsequent
collection
of
thewaterfauna.During
collecting
weshouldkeepin
mindthe ethicalsideof the investigations
andthe
Cottection
of terest at [email protected]]s,Dohica Cave,photo:p. Lupteeik
aspectof the faunaconservation.
Thus,it is highly
desirable
to modirymethods
to prevent
remarl€ble
losses
inthefaunapopulationsand
to prevent
distur,
banceof theexisting
particular
equilibrium
between
components
of thesubterranean
slstem.
Occasional
collectingof invertebrates
in a cave
cannotin factseriously
threaten
the subterranean
communities
sincethec€veis onlya windo{intoan
ammense
system
ofcrevices
inthelimestone
bedrock.
just
Thus,
theorganisms
foundintheca\r'e
represent
a minutefractionof whatreallylivesin the wnote
karstmassif.Massive
deathof thec€vefaunamaybe
caused
bynaturaldisturbances.
Forexample,
during
hea\,y
rainsthousands
of animals
areflooded
outof
thecaveswheretheydieinthesurfacewaters.
calectionof aquattcfauha,Dobainske
be cave.Photo:p t:u aajk
Explorations
of bats are carriedout by three
basicmethods.
Thefirstoneis the directcounting.
However
this methodis difficultto usein batsDefixedandtransported
in vialswithethyl€lcohol
or
preferring
ciesor particular
individuals
inaccessible
formaldehyde,
beetlesin vesselswithwoodshav- or hardlyaccessible
spacesor crevices
for resting.
ingssaturated
byethylene
acetate.Pitfalltrapping Thefurthermethodis identification
ofspecies
using
'BAT"detectors
is anothercollecting
method- a plasticvesselwith
basedon the characteristic
suoerfixationliquidis dugintothe softercavesediment soundsignalsof the particular
species.
N4arginally,
(loam,smallerstonydebris)sothatthesediment
is
as the additional
method,collecting
intothe nets
levelled
withthevesseluppermargin.
Forco ecting is sometimes
used.Themethodis, however,
tess
of tinyanimalformsdwellingin bat guanoor rotsuitable
sinceit canstressthebatsandmayapparten wooda photo€clectorapparatusis used.The
entlyinfluence
thebehaviour
ofthelnvestigated
bat
device
consists
ofthemeshonwhichstudymaterial colonyinthecave.
707
CONSERVATION
OF THE CAVEBIOTA
waters. Similarly,negativeeffects are obserued
when soent carbideor batteriesare deoositedin
Thedistributionof the subterranean
organismsis
caves.Excessive
organicsubstancesoriginatingin
in fact not limitedto caves,it also coversepikarst
thecattlefarmingor
inthe rinseofchemicals
during
(the systemof crevicesin disconnected
rock under
wintersaltingof roadsmaythreatenwaterbiotaof
surfacesoil layers),submergedspacesin the
the subterranean
habitats.Construction
of damsin
ohreaticzone, as well as micro-soaces
between
the watershedcauseschangesin the profileof the
particlesof softer sedimentsof the subterranean riverflows and in the accumulationof sediments.
running waters (interstitialmicro-habitats).
Since
Interstitialmicro-environments
with uniquefauna
the resources
of the subterranean
systemoriginate
may be completely
destroyedby fine clayparticles
on the surfacefromthe oroduction
ofthe olantsand
originating
in damssincetheyfill up minutespaces
animals.the conseruation
shouldalsocoversurface
betweensand and gravelparticles.Populations
of
habitatsdirectlyconnectedwiththe cavesystems.
caveinvertebrates
may be threatenedby intensive
Conservation
ofthe cavebiotaresultsmainlyfrom
collectionusingpitfalltraps,especiallyif they are
the fact that manyspeciesof organismsare "living
left in the cavefor longerperiods.
fossils".Theydisappeared
from the surfacewaters
Dropin populationsof particularbat speciesis
and terrestrialhabitatsmillionsyearsago surviving
causedespeciallyby disturbanceduring hibernain caves,undergroundwatersand other types of
tion and nursingof the young,localvandalism,
subterraneanhabitatsthus protectedagainstthe
lossof habitatby closingof cavesand minesor by
harsh weatheroutside.The other reasonfor the
changeof the cavemicroclimate
by the openingof
protectionis the uniqueness,rarityand increased
new entrances.changeof the landscapesurface
vulnerability
fauna,especially with decreaseof plant diversityand subsequent
of the cavernicolous
endemicformsandtroglobites.
Thefollowingspecies decreaseof insectsthat serveas the foodbasishas
deservespecialprotectionin our caves:palpigrade negativeeffects upon bat populations.Moreover,
Eukoeneniaspe/aea,pseudoscorpion
Neobisium
batsmaybethreatened
directlybyexcessive
collects/ovacum,millipedeof the genusTyphloiulus
ingfor scientificpurposes.
and
Pseudosinella
collembolans
aggtelekiensis,
P paclSeveralinternational
Conventions
can be aoolied
ti, Arrhopalites
aggtelekiensis
and A. irtermedius
to subterranean
speciesand habitats,for example,
(13 June,1992)for biodiversity
since they representuniqueforms with very limthe RioConvention
protection.
(23lune,
ited distribution.Thevulnerability
of theseanimals
In Europethe EonnConvention
resultsfromtheir lowecological
flexibilityand small
1979)for theconservation
wild
migratory
of
species
distributionranges.Thus,theyare threatenedeven
is relevantto troglophilous
bats.Lawno.543/2002
by localchangeof the ecologicalconditionsor by
of the NationalCouncilof the SlovakRepublicon
pollution
of the environment.
the conservation
of natureand the landscapealso
Directdestruction
ofthe caveenvironment
in limeaooliesto all caveanimals.Basedon this law it is
stone quarriesis the most seriousdangerfor the
forbiddento interruotDrotectedanimals in their
subterranean
In
way,
locality
organisms. this
the
of
naturaldevelopment,
especially
to kill, injure,catch
the first discovery
of the subterranean
carabidbeeor movethem. Moreover,
it is forbiddento destroy
tle Duvaliusbokori(LeontinaCavenearGombasek) and to harm their biotopesand dwellings.Many
almosttotallydisappeared.
Chemicalpollutionisthe
formsof the cavefaunaare directlyincludedin the
mostdrasticfor the waterfaunasincewaterreprelistof protected
animals.However,
we shouldrealize
sentsa mediumof fasttransportforpollutionagents.
that orderingof the givenspecieson the list of the
Theseare mainlypesticidesappliedin agriculture, orotectedfaunadoesnot havemuchimDortance
if
pollutionby petrolsubstancesduringaccidentson
we are not ableto guaranteeprotectionof the par'
water-courses
and pollutionof watersby heavymetticulararea,the wholesystemwith its components
als or aggressive
colorationsubstancesusedduring
whichis populatedby this faunaand on whichit is
investigations
of the hydrology
of the underground essentiallv
deoendent.
108
CAVES
OFTHEWORLDHERITAGE
IN SLOVAKIA
STATENATURECONSERVANCY
OFTHESLOVAI(REPUBLIC
SLOVAKCAVESADMII{ISTRATION,
LIFIOVSI(VMIKUIAS
Edltors:DoGRl{Dr lo[bf Jakiil, Dlsq
Rl{Dr.PavelB€lla,PhD.
UptovskiMikulSS
Author$Rl{Dr.PavelE€lla,PiD. - SlovakCalesAdmlnlstr€tlon,
Llptot6ldMlkuliiS
RNor.fudovlt Ga6l- Slov€kCav€6Admlnlstr8tion,
Mdr.DadrnarHavlarov6- slo\€k Cdv€sMmlnlsuatlon,LlpbvskyMlkul6a
llpto6k, Mlkul6S
Ing.Joaf Hlae6d- slovakca\,€sAdmlnlsuation,
Insttute of SqS,Btatislava
Doc.Rt{Dr,.rozefjak6l, DEc, - Geographlcal
Ko6lce
Doc,RNDr.tubomlr Kov6d,csc. - Facultyof llatutal scl€nc€s,P.J. Saf6rlkUntu€rslty,
Ing lrarcel Lall(ovL,CSc,- Facultyof NaturdlScl6nce5,M. B€l Unlw6rty.Bansk6Stlavnlca
PhDr.Mad6nSoldlqPhD.- ArcheologcalInsd$le of SAS,SplSskSNor6Ves
Llpto/sldMlkulSS
Rl{Or.J5nZellnka- SlovakCav€ Admlnistratlon,
Revisver$RNDr.v6clavCllek,CSc.,Prof.PhDrVliclavFumSnek,Drsc.,
csc.,
Doc.RNDi.zdenkoHochmdh,csc., RNDr.vladlmlrK0d€1,
Doc.RNDr.MllanL'apin,csc., RNDr.LadlslavNo\otny,Ing MlkulSSRozloznlk
Ceer pagBphotogapk: GombasecMCa!€.Photo:M. Rengevld
Hruaovsk6
cave.Photo:l. stanloue
lssuedbryO S'Hs NatureCorcervancyofthe SlovakRepubllc,
Slo/akCav6 Admlnlstatton,UptovBldMlkul6sin 20Oa
In KnEn6cemrum,Prcdm6tsk651, 010 0l Zlllna
Mgr.BohuslavKortman
Redacuon:
T€nslatlon:Ing FbterGatlk,D,oc.RNDr.Lubomiiyor/64,(S6.
corecuons:Martlnc. styan
Graphlc$
Ing.J6nf€s6k
Prlnt PolWIafpdnr,Preao,
|sB Cra-AO-aOA4-308-4
I
CONTENS
(lozefHlav66)
Foreword
5
(JozefJak6l,
lntroduction
Pavel
Bella)........
9
SlovakKarstNational
Park- Remarkable
Landscape
(Pavel
ontheSurface
andLjnderground
Bella,
JozefJak6l)......... lL
S o l u b l e R o c k s - T h e E n v i r o n m e n t f o r K a r s t a n d C a v e D e v e l o p m e n t ( L u d o v i!7
tGa6l),.....
(JozefJakSl,
Water Creative
Agentof KarstandCaves
PavelBella)
25
(Jozef
TheSurfacel\4orpholog/
of KarstLandscape
Jak6l).. . . . . . . . . . . . . .
(Pavel
Passages,
Labyrinths
andVerticals
undertheEarth's
Surface
Bella).. . . . . . . . . . . . . . . . . . . 47
(Ludovit
TheStoneBeauty
of Caves
Ga6l)
67
(J6n
CaveMicroclimate Zelinka).
............
a7
C a vLei f e ( L u b o m i r K.o. .v. 6. .6. .) . . . . . . . . .
95
(l\4ari6n
CaveSettlement
Soj6k)... . . . . . . . . . .
109
(l\4arcel
History
of Knowing
andDiscoveringthe
Caves
Lalkovii).. ....
123
(Pavel
Thelvostlmportant
Caves
andAbysses
of theWorldHeritage
Bella). . . . . . . . . . . . . . . . . . . 139
(JozefJak6l,
Protection
of KarstandCaves
[udovit
Ga6l,
Dagmar
Haviarov6)
..................
Selected
Literature
L67