Caves of the World Thereis Eventhe Chanceto Meetsome Animals, Naturalheritage

I t

CAVELIFE LubomirKov66 ru E

It is widelyknown that cavesdo not representa uniquesubterranean animals in the karst regions of deadenvironment without any traces of lifeand that Slovakia.lvlany of theminhabit caves of the World thereis eventhe chanceto meetsome animals, NaturalHeritage. The Slovak Karst represents an for examplebats, frogs or salamanders.only a few importantevolutionary centrc of the subterranean peopleare aware of the presenceoftiny organisms faunawithin the Western Carpathians. withtheir immense diversity and species richness in thecaves. THE CAVEECOSYSTEM An independentscientific discipline focused on researchinto such life forms and their relations with Characterizationof the subterrarea[ (cave) eco- the surroundingenvironment - t ospeteolo8S/was systemis necessaryif we wantto understandthe establishedon the basisof this phenomenon.We crucialphenomena of the distributionof living can meettypical cave animals, often with bizarre organismsin caves.Among the mostimportant shapes,in tropicaland subtrooicalcaves of the aDrotrcfactors (agents of the "lifeless'environ- world.In Europerich undergroundfauna occurs ment)arelight, temperature and air humidity.Light predominantlyin caves of the Baikankarst moun- is a limitingfactor for the occurenceof vegetation tains. Intensivebiospeleological research in the in the entranceparts of thecaves. The majority of last tlvo decadeshas conflrmed the Dresenceof undergroundfauna escapes from the lightto the dark,a phenomenoncalled photoptroDra. However, it wasobserved that light does not necessarilycon- stitutea limitingfactor. For example, aquatic crus- taceansof the genusNrphargus regularly occur in karstsprings where they find richerand more accessiblefood, but also more natural enemies. Caveair temperaturereflects the yearlyaverage valueof the externalenvironment as a resultof the latitudeand altitude ofthe given site. The tem- peraturerange, which the animals are abletotoler- ate,differs with the species.Subtenanean fauna is ableto withstandlower temoeratures {close to the free2ingpoint) more easily than values above 20 "C. Air humidityis an importantfactor ruling the distributionof true cavefauna that prefers highvalues (95 - 100%). Drier spaces are poorer for specializedsubterranean forms. In caveswith Flyi.g bat, Donica Cale. Photo: P, Bella highair humiditydue to percolatingwater, such

95 F...r f1.! rr ..re e, ftor ,f..t t.h.t l\1 Cti)..|.t at 2DA2

as the Donca or Kresnohorsk6Cave, Niphargirs CAVEVEGETATION specmefs ntaybe observed ln waterpools ol even movingon the wetcave flooT overcorn nE re at vely Caves,deep sinkholes. and gorgeshave special ong0rstances. vegetauon.The rapid changes n the envTonnrental All ivLngcomponents of the ecosyste..may lre conditonsgve rise to the wellknown phen0nl c assfied as biotic factors.Autotrophic organ sms enon of inversestfttification of the fora. n the arethe categoryassoc ated wth the presenceol S ovakKarst the greatentrance part of the SLcka ght (algae. osses, chensand hgher pants). TheyprodLrce organ c substancesas a resLrt ol prro tosyntheslsperfornred inthe r bodies.Heterotrophlc organisTnsare consumers of thesesubstances and so areprlmariy dependent on the autotrophs.Thrs groupincudes bactera. mcroscopic fungi and an mals.The presence of autotrophicorganisms is linitedt0 the entranceparts 0T trre caves exposed to the ghton whichthey are dependent. however, theyafe absent n thelnterna cave spaces and the foodweb is basedon y onthe nutrentsI theforin of organicremnants originatlng in thesurface karst horizons.A s mpllfiedfood web dominated by detri. tophagousanima s {thosefeed ng on deadorganlc materalsl and carn vorous (predator) lorms s chat L'€fuon5 'rrib,rr.rs or rr. acteristicoi caves.

96 Mad$copb gnlpetdous funei in the xr'snohotsk' Cave. At&e - pe.t of the tano twa, Donlca Cave.

zorp - wemay find ferns, mosses, liverworts, lichens andalgae growing on the primarysoilor rocks. In the lowerpart of thisspace only ice and talus deposits arepresent. In the zoneof full darknessof caves- the aptDuczore - vegetationis completetyabsent. Thatis whyit is notcorrect to classiryany plant as a truecave orlFnism although some species prefer conditionsclose to the caveentrances. Deeper caves areinhabited by primitive microscopicfung {moulds), light'independentbacteria and in some specialcases alsoby green algae. They are very imponant as food sourcesfor manyanimals. On rotten wood we may observefruitingbodies of ligniperdous fungi. In caves the coloniesof microscopicfungi such as Mucot mucedomay be easilyrecognized on the surfaceof thebat guano. GroMhsof primitiveplants around light bulbs, so- calledtamp Ito'a, represent artificial communities incaves. Lampflora is composed mainly of diatoms, blueand green algae, mosses and sometimes also of ferns.lt is an undesirableand unnatural compc Mucormocedo -. colonyof the nicroscopt. funEi covetingbat cuano, nent,development of which is apparenfly limited by usingof coldlight and is removedeither mecnan! callyor chemically. IadnicaCave, evolved byfailure ofthe cave ceiling, is a goodexample of thisphenomenon. Counting from CLASSIFICATIONOFTHE CAVE FAUNA the surfaceof the karstplateau to the end of the floor ice lt is 79 m deep.The air tempera- Cavernlcolous(rock crevice inhabiting) animats tureat the bottomof the monumentalentrance of dwellingin subterraneanhabitats are adaDtedto ablssalcharacter fluctuates only negligibly around this envkonmentto differentextents. 'C. Thus,sev- 0 Wemay observe here wellmarked vegetation eralecologcal categories were ouflined that parfly associations.Atthe margin ofthe ptateau with direct reflectadaptat ons of theparticular faunal species. sUnlight,a eupftot c zorewith predominanty higher, 0f the proposedclassifications, division of thesub- floweringplants occurs. ln the twilight- dlsptotc terraneanfauna into four categories is widely used

97 at present.Terrestrial animal forms enteringthe bodyshape (morphological), internal life processes cavesonly at random,perhaps transported by water (physiological),behaviour (ethologacal) and heri streamfrom the surfaceor falleninto abysses,are tage (genetic).The morphologicaladaptations are calledtrcgloxenes. Animals that may be found in very marked and often very bizafie, especially cavesmore regulady are classified as troglophiles. in the warmerregions of the Earth.Reduction or They representa broadercategory involving two completelost of the visualapparatus is character- ratherdifferent groups. Subtaoglophlles are associ- istic of the majorityof true caveanimals. They are ated with cavesonly during certain stages of their paleor whiteas the consequenceof the reouclon life cycleand they do fot possesany specific adap or absenceof pigmentationin coverbody layers. tationsto thjs kindof environment.Some dipteran In many arthropodswe may observeapparently flies or bats may serve as examplesof animals elongatedantennae and legs.0n theseextremi- dwellingin caves only duringa certain season ofthe tres an increasednumber of elongatedsensory yearor duringsome part of the day.Eutroglophlles setaeis distributed.The function of theseexternal are animals permanentlypreferring the subterra bodychanges is to strengthensensory organs that neanenvironment and arewell adapted by external areessentialfor life in completedafkness, namely bodyconstitution and usuallyalso by internatlife sensitivityto airhumidity and odour (chemical) sig- functions.However, they may marginallylive and nals.Longer legs enable the animal to movefaster reproducealso beyond the caves,as do somemites, whensearching for food,sources of whichare usu- collembolansand other arthropods. 0bvious adap- allypoor in thecave environment. Amongthe most tationsto the subterraneanworld are characteristic pronouncedphysiological changes the following of troglobites,which spend their whote tives in are concerned:elimination of the dailyrhlthms cavesor similarhabitats. in animalactivities, inhibition of the metabolism The similarclassification may be usedfor the and abilityto starvefor longerperiods, and lower aquaticfauna, wherewe may distinguishstygox- numberof eggs laid. Ethologicaladaptations are enes, stygophllesand eustygophites.organisms connectedwith the modein whichthe animals strictlylimited to occufiencein undergroundwater colonizeunderground habitats. are classifiedas stygobites.The prefixof these wordsis derivedfrom the StyxRiver, that in creek THEUNDERGROUND ENVIRONMENT m}{hologyrepresented the riverthroughwhich dead AND CAVEFAUNA peopleentered the underworld.We may register Suchanimals in theStyx of tl'e DomicaCaVe Or in Terrestrialsubterranean environment the undergroundstreams of the Kr6snohorskdand Theterm "cave ecosystem" was used for a long Hrusovsk6caves. However, these expressionsare timeforthe pats ofthe natural underground spaces notstrictlyconnected with the cavefauna.For exam- reachableby man.However, caves are not the only p e. lhe phreatobltesthat lve In lhe tiny crevtces environmentinhabited by the cavernicolousfauna. amongthesand particles and gravels in submerged The terrestrialunderground environments may be sedimentsof the caveand abovegroundstreams dividedinto three main groups: also belongto the stygobites.lt is not atwayseasy 1. naturalcaves to allocateanima s occurringin the undergroundto 2. epikarst oneof the above-mentionedcategories, since there 3. artificialcaves made by man are manyintermediate cases. This divislon of the Naturalcaves usually represent spaces that may subterraneanfauna suggeststhe necessityof pro- hide even larger animals bats or amphibians. tectionof the specificmicro-environments inhabited The presenceand distributionof the animalsin bythis fauna. them is determinedby variousecological factors. In manycases we maydistinguish several welldefined ADAPTATIONSOF THE CAVEFAUNA associationsof the animalsoccurring in caves.Cave entrancesare markedly influenced bythe externalcli- Adaptatlonsto life in the slbterraneanspac- mate.These usually wetter and protected sites serve es are obviouspredominantly found in animals as refugesfor some bird species(pigeons, owls, classifiedas troglobites.Interestingly very similar passerinebirds). Some animals nest or stayclose to adaptationsare observed in verydifferent faunal the entrances,for example,dormice, frogs, salaman- groups.we mayrecognize adaptations in external ders,snails, insects and other arthropods. From the

98 entranceto the beginningof the aphoticzone, cave bodiesare considerablymodified enabling them wallsand ceilingsare inhabitedby a heterogeneous to live in the interstitlal envlrcnment,namely communityof animalsdefined as pa etal fauna. the micro-spacesbetween sand and gravelpar- Theyrepresent occasional cave animals using cave ticles.The crustaceanBarhynella natans ts a entrancesonly as temporaryshelter mainly during goodexample of suchfauna. Water circulating in summer and winter (troglophiles):dipterans, but- fragmented,eroded rock is presentpredominantly terflies,trichopterans, spiders and opilionids.the anthe karst regions.We recognizetwo hydro,geo- compositionof such communitiesis dependenton logicalzones within this environmenttype. The the size,orientation and altitude ofthe entrance,fur, vadosezorre includes the upperpart of the karst theron the structure of the entranceparts of the cave massifwith mainly vertical flow of the water.Here and on the seasonalchanges of the above-ground lhe faunais welladapled to Ife in smal'crevices, climate.Internal parts are usedby bat coloniesthat it may occasionallyappear in waterpools, too. produceexcrements rich in nutrientsduring their Theyare chiefly tiny animals with elongated and activityphase - guano,an importantfood source for flattenedbodies, such as planarians,nematodes, a widespectrum of invertebrates.Depending on the oligochaeteworms, and from crustaceans cyclops degreeof associationwith such micrcenvironment andamphipods serve as goodexamples. We may theymay be classifiedeither guanophiles or guanc also detectlarger forms in such habitats,like bltes.0n the pilesor smallerguano accumulations amphipodcrustaceans of the genusNiphargus we mayrecognize earthworms, mites, collembolans, the lengthofwhich can exceed 2 centimetres.The woodlice,millipedes, beetles and dipterans.Thus, phreaticzone coversthe part of the massifwith the populationsof suchanimals are primarilyassc mainlyhorazontal water flow. The zone ls usually ciatedwith the presenceof bats dependingon the populatedby a higlernumber of fauna,specres. amountand the ageofthe guano. This is the kingdomof largercrustaceans, for obligatecave forms ofthefauna live in the deeper example,lhe dbovementroned Nipha/gus. partsof cavesin completedarkness, stable temper- aturesand highair humidity.We may observe them DIVERSITYOF CAVEORGANISIVIS onthesurface of sediments and speleothems, under rocks,on the organicremnants or on the surfaceof fhe blodivebiaylspecies diverslty of the organ- the waterpools. In mostcases they representtiny isms)0f cavesprimarily depends on the abioticfac- animalsthat maypenetrate into eprkarst consisting torsof the environment.Heterogereity of the given of the systemof interconnectedmicro caverns tn habitatis alsovery important since a widespectrum fragmentedbedrock. lt is situatedin the upperzone of micro-habitatscreates rnore possibilities for coloni- of the karstmassif under the superficialsoil layers. zationby various forms of organisms.The size of the Thisenvironment type links caveswith the restof karstarea is crucialin this respectsince larger areas the karsticsystem. lt has been biologicallypoorly arenormally inhabited by a highernumber ofspecies exploredsince it is hardlyaccessible to man. of organisms.This is alsothe caseof the vast pla- In the tefiitoryof Slovakia,the third underground teausof the SlovakKarst. The biodiversity of caves environmenttypehas been poorly understood - the alsodepends on stabilityof the environment.Less artificialshafts and adits of mines.The biological stablecaves with rather fluctuating conditions during observationsin that environmentmay add consider, the year creatediverse micr+habltats thus popu- ablyto understandingof the distributionand evolu, latedby faunal communities richer in species.Food tion of cavefauna in the country. accessibilityis oneofthe principalfactors regulating biodiversityof the subterraneanhabitats. Eltrophrc Theaquatic subterranean environment spaceswith sufficientsupply of organicmatter are This environmenttype may be classifiedinto rich in faunaand micrcorganisms,for example,the two maintypes: waters circulating in porousrock bottomsofthe entranceshafts with fallen plant rem- (alluvialterrains) and watersflowing in discon- nants.or caveparts wtth gualo deposits.Howeve( In nected,predominantly karstic rock. Both types are theseparts trogloxenes, troglophlles or guanophiles inhabitedby different fauna with specific adapta- prevail.Ongotroptrrc caves are poor in organicmate tionsto lifein theseenvironments. Animals ofthe rialsand are popularedby faunapoorer in species. first type are phreatobites,fascinating forms that 0n the otherhand, such caves host higher number of havebeen insufficiently known up to now.Their trogobiticanimals compared to the precedingcase.

99 ORIGIN,EVOLUTION AND DISTRIBUTION THEPECULIARITIES OF LIFEIN CAVES OFTROGLOBITES OFTHE WORLD HERITAGE

Whyand how did animalspenetrate into the Inthis chapter the biotaof the bestknown caves subterraneanworld? How is it possiblethat they is stressed:Domica, Gombaseck6. Krdsnohorska wereable to adaptto sucha harshenvironment? andJasovsk6 caves, Silicki l'adnica Cave.ochtinska Thereare two main theoriestrying to answer AragoniteCave and the Dobsinsk6lce Cave. We will these questions.Both considerevolution from alsomention some peculiar animals of othercaves ancestorsoriginating in aboveground habitats or abyssesof the SlovakKarst that are the partof (soil,surface waterstreams). 0ne ofthem regards theWorld Heritage. caveforms of faunaas remains- relrctsof the In recentyears a widespectrum of mlcro-orgaF precedinggeological periods, so-called'living lsns wererecorded in the cavesof the Slovakf€rst. fossils".For such forms the cavesare the refugeg Greenalgae a re typic€ | com ponents of the lamp flora, againstunfavourable external conditions. Based withStichococcus bac,r,a s, Chlorellaminutissima, ol on this theory,surface fauna was forced to look speciesof the genusDesmococcus sp. l\4icroscopic for sheltersdeeper underground with the startof fung areextraord ina ry intheir divelsity. They serve as the longperiods of the coldand dry Pleistocene the importantfood source for manysmaller inverte climateroughly 2 millionyears ago. The second brates,such as mites and collembolans. The species theory,on the other hand, considers active coloni- of thegenus Mucor dominates among the first colq zationof cavesbyfauna, a processthat continues nizersof bat guano.Entomopathogenic Beauveth up till the present,However, not all animalsare brcngnianiic@at€svatia blemacroscopic colon ies on able to populatesuch an environmentperma- limestonerocks and siniet lillings,Pidoplitchkovie a nently.In somecases this wasthe wayto the evo, tenlcolaisolated from isopodexcrements is a very lutionof a highernumber of relatedforms from raresaprotrophic microfungal species. oneancestor within a relativelysmall area, the Theunicellular animals - protbb (Protista)have evolutionaryprocess called adapttre radtatlon. not beenexplored sufficiently in the cavesof the Thisis the caseof carabidbeetles of the genus Slovakiayet, so we havealmost no information Duvaliusand its subspeciesin the territoryof the on this animalgroup. Planarlans (Turbellaria) of WesternCarpathians. the genusDendrocoelum are ratherrare among Colonizationof the subteraneanhabitats by the aquaticfauna, they are predatorsfeeding on thesurface fauna, either passively as the consequenceof unfavourableclimate or byactive pen- etration,is accompaniedby speclatron,that is bythe evolutionof newspecies. ,soratlon of the spaceor areais an unavoidablepremise for the evolutionof newspecies. Populations ofthe orig- inalspecies isolated in subterraneanspaces for a relativelylong time (tens-or hundred-thou- sandsof years)become differentfrom the ances- torduring successive adaptations, species being the final productof this process.In manykarst regionsthe populationsof the sane species populations. areisolated from the surface l\4any Planaia. wm of the genLs Dendro@lum frcm the Farbeni ponor examplesof the originof independentspecies Cave,Photo: J. Stankovld withinone cave system are known. Such unique forms,limited in theiroccurrence to a relatively smallarea, are called erderrrcs. They represent tiny invertebrates,including dead bodies. Among the rarestand mostvulnerable forms of life. the smallforms of annell& oligochaeteworms Wewill concentrate on themmore thoroughly in (Annelida)that live in the claysediments of the the followingpart involving the peculiaritiesof undergroundstreams, or in decayingorganic mat+ the cavelife within the WorldHeritage sites in rialswithin the terrestrial habitats no uniqueforms Slovakia. havebeen observed yet. Larger annelids - earth-

100 wormsrepresent permanent components of the faunaof manycaves being preferably associated withthe presenceof wood.They are ableto con- sumegreat amounts of claysed iment together with organicremains. In the DomicaCave several earth- wormspecies have been detected, but notspeciaf izedcave forms. Remarkable traces of theiractivity arethe casts (earthworm excrements) in many sites in the cave,especially in the riverbed of the Styx, ofteneven on the sintersurfaces. Leaches have occasionallybeen registered in the underground waters,Ior example, the surface species Haemopis sarguisugain the Domicacave. Molluscs(lvlollusca) may be ratherabundant inthe cavesof Slovakia.The snail oxychilus glaber is an abundantinhabitant of the entrancemicro-habitats. In karstsprings we maymeet the snailSad/eriana pannonica- an endemic ofthe Slovak and Aggtelek Karst,and Biikk l\4ts. in Hungary aswellas Eythinel/a austriaca.A so far undescribedspecies of cavesnail Parp,iqradeEukoenenia spelaea (body tenEth 7.5 nm, of the genusHauffenra has beenreported from Phato:G. C.iah4hut ahdL. t

Snat OxychilusElaber lbody /enEth 74 nm), KresnohoBkeCaee.

Theh ighest species d ive rsity of caveinvertebrates is concentratedin the animalphyllum Arthropoda. Severalunique animals ofthe WorldHeritage caves belongto the arachnids(Arachnida). PalpiErades (Palpigardi)have been registeredin Slovakiain Beudosco,pionNeobsilm (Slothrls) slovacun (body teh4th4 nn). one sDecies- EukoeneniasDe/aea distributed in

701 not livein caves.Troglobite Neobisium (Blothfus) s/ovacum,characterized by stronglyelongated pincerpedipalps, have been observed in many abyssesof the Ple5ivsk6Plateau, and in a limited numberof caveson the Hornivrch and Zadielska plateaus.The species was described by J. culiaka in the 1970s.This was an imoortantlandmark in Slovakbiospeleolo$/ since it representedone of the mostvaluable confirmations of originaltroglo- biticfauna in the Slovakcaves. From the haruest- Har€srneh lschyrcpelis manica\a often pdet?tinq deepu m6n(Opiliones), ,schyropsalis manicata is a sma- into the cavet Photo: P, lupt4iik er predatoryarachnid with remarkablechelicerae inhabiting,for example,the DomicaCave. Spiders {Araneae)such as Meta menardi are found in large numbersin caveentrances, where they catch flying insectsin theirwebs. Some smaller forms of spi- dersare adapted to lifein the deepercave spaces wherethey look for theirprey, most often collembolans,mites or insectlarvae. Ihe mostpeculiar of thembelong to the genusPonhomma of which P. profunclumis consideredtroglobitic species. Thespecies was described from the DomicaCave whereit establishedstable pooulations.Mltos representan arachnidgroup Sptder Meta menadi, inhabifant al the caw ennanc*. lhoto: p tuptaaik withstrongly diversi- fied communities,Oribatid mites consume algae, fungiand rottenorganic malerials (wood, guano). Pantelozetescavaticus is closelyassociated with caves.lt wasfirst described from the Certovadiera Cavenear Domica. lt is a eutroglophitemite often occurringin closevicinity to bat guano.oribatid miteDamaeus lengensdotfi is a largerform with elongatedlegs, an inhabitantof the DomicaCave. ln the samecave, the uropodidmites Woobovella advenaand Cyrtolaelapschiropterce are among the principalconstituents of the bat guanoasso- ciations.Predatory mites are representedby the eutroglophileform Parasitus/onbatus, a regular inhabitantofthe Slovak caves, or rhagidiidmites of Ircglobit c spider Pod.omma p@lundom,Oom,€ Car€, photoj I Srankolrd the genusRobustoche/es registered in the oomica Cavein recentyears. Ctustacean3(Crustacea) are predominantly aquaticanimals. The terrestrial eutroglophile isq podeMesoniscus gran€er (lsopoda) isan exception establishingabundant populations in the Domica and Jasovsk6caves as the importantconsumer of the bat guano.The isopode was discovered by Slovaknative l. Frivaldszkyin 1865 in the Earadla Cavenear Aggtelek, a partof the Domica-Baradla caveslstem. Anong aquatic crustaceans Niphargus tatrensisis worth mentioning. lt is distributed inthe Oribatd nite Pantelozetes€vari.us (body tength O.5 ntu). majorityof Slovakcaves with standingor flowing Photo: G. CsiAtu'rova and P, LuptAiik waters.As a symbolofstygobitic animats it js abun-

702 dantespecially in the St)r(River (Domica)where the individualsare notablefor their unusualsize of up to 4 centimeters.Rich communities of cladocerans (Cladocera)penetrate into the DomicaCave by the flow of the DomickyBrook especially during hea!ry rainfalls.However, in the undergroundwaters in Slovakiano obligatecave species has beendiscov- eredyet. Ofthe relatedcopepods (Copepoda), three speciespopulating the StyxRiver may be classified as phreatobitic:Acantocycrops venustus, Diacyclops lanEuidoidesand Microcyclopsrubellus. Bathynella natansbelonging to the Bathynellaegroup of crus- taceansis a phreatobiteliving in the microspaces of submergedsediments of the undergroundriver' /sopodMeso.lscus gEniger Photo:). Stankovia beds.lt maymarginally occur in lakessuch as in the BielaHall of the Dobsinsk6lce Cave. The pale and blind mllllpede{Diplopoda) of the genusIyphloiulus was detectedin the Domicaand Gombaseckdcaves thus representingone of the mostvaluable discoveries of cavefauna in Slovakia in recentyears. lt is olr largesttroglobite with a bodylength of 2.6 cm and 147 pairsof leg.The miL lipedeBrcchychaeteuma bradae is a typicalinhabit- ant of the DomicaCave. The Dobsinsk6 lce Cave is inhabitedbyAiiorhiscosoma sphinx, a cavernicolous milliDedeendemic to the WesternCaroathian caves. Millipedesare importantconsumers of decaying organicmaterials deposited in caves. Collsmbolans(Collembola) are amongthe most abundantarthroDods of the subterraneanhabitats. sq€obnt c/usraceanN pha/gustanens s. PhoroiJ. slankovid Severalspecies in ourcaves exhibit a highdegree of adaptationto that environment.They preferably feed on fung coveringthe surfaceof decayingorganic remains,or eventhe surfaceof the speleothems. HypoEastturacassaeEranulata dobsinensis is an inhabitantof the inversedentrance moss vegetation of the DobSinsk6lce Cave. lt is a glacialrelict surviving he.efrom the lastPleistocene gaciation. Several troglobiticforms of Collembolamay be recognized within the generaDeuteraphorura, Pseudosine//a F{r@d,'La l)]edr Barh'+lla nM6(body[.nErn L5 mm) ry'ofo r Hr-dec

hoElabitic mi ipede of the gen!s Typhloiulus,Domica Cave. ltaElobitic @llenbolan Pseudosinellaaggte ekiensis (body,ength 2 mnl,

103 and Anhopalites.Deutercphorurc krctochvili is BeetleE(Coleoptera) are among the mostremar- a typicalcollembolan of the Dob5insk6lce Cave. In kablecavernicolous arthropods. E, csiki and E. the cavesof the SlovakKarst the speciesis replaced Bokorprovided the first informationon the cave bVa closelyrelated undescribed one. Deuteraphorufa beetlesof the SlovakKarst at the begnningof the schoenviszlryiocurring in the Gombaseck6Cave is lastcentury. In cavesthe genera Trechus, quedius, a troglobiteas welland an endemicspecies of the Athetaand Choleva ate the mostoften representa- Slovakand Aggtelek Karsl Pseudosinelaagtelekien- tivesand the majorityofthem may be classified as sisis an endemic species distributed inthesame area troglophiles.Inthe Domicacave lrechus austriacus whereit isoften abundant. Inkarst regions of the cen- and ouediusmesomelinus create rather dense tralWestern Carpathians it is replacedby the closely populations.The genus Duvaiius covers the only relatedP pacnithat livesin the Doblinsk6lce Cave, obligatecave beetles in ourcountry. The species of too.The genus Arrhopaiites includes several obligate thegenus are endemics of theWestern Carpathian c€veforms in ourterritory Eutroglophilous A.pygmae mountainranges preferably inhabiting the superfi- usis a frequentspeciesofthe cave communities. 0n cialunderground environment (deeper parts ofthe the otherhand, troglobitic A. aggte,ekiensisis distri- stonydebris), in cavesthey are rarer. Interestingly, butedin a limitednumber of undistulbedcaves where two differentspecies live together in the Jasovska theymay be rather numerous, as in the oertova die€ Cavenear Domica and in Kr6snohorsk6Cave. lt isan endemicspecies of the SlovakParadise, Murenska Plateauand SlovakKarst. During biospeleological researchin recentyeals some unknown collembolan specieshave been dscovered such as a newspecies of the genusMesaphorura in the KrSsnohorskiCave orthat of Neelidesand Megalothorax in theJasovske Cave.Such discoveries confirm how limited our knowl- edgeof suchsmall livingcreatures populating subterraneanhabitats still is. Staphylhid beete Quedi6 msmeli.us, Oomie CrE PlDto: P tuptidiik

Co enbota on su.faceof a watetpoot. Kt6shohotsk' c.ave (body le gth ltoCtobitb catabid tuetle ol the ge.us Duvalius, (.dsnohoEld CaE, of snar/erMegalothorax sa o 5 hn). F,hoto:.1. stahkoia

Caue- D. hunEaicusand D. bokori.Dlptsran fllss (Diptera),buttertlles (Lepidoptera) and caddbflle3 (Trichoptera)represent an importantpart of the parietalfauna with seasonal emergence in caves (subtroglophiles).Of dipterans Triphleba anticola is a stablecave dweller ofthe JasovskdCave asso- ciatedwith the batguano accumulations. The but- terffiesScoliopterr4 libathix andTriphosa dubitata arepart of the typicalfauna of the entrancecom- munities,being numerous especially during the TtoCtobiticcoltenbolah Anhoparites aggterekiensis{body lencrh 2 mm),

104 regardedthem as an inseparablepart of caves.In conditionswith little knowledgethere were a lot of legendsand reportsabout them, Nowadays we knowthatbats lookfor caves as one ofthe possjble sheltersduring their resting period, either duringthe dayin the warm months, or in the winterseason dur- ingrrlDemat or (winterresting stage). They arc one of the fewgroups of warm-bloodanimals that stay duringthe winterin a stageof truewinter sleep in Aude.ft ScoliopteryxlibaMr inhabiart of the caveenttancT. whichblood circulation and respiration are several timesslower and body temperature drops to 10 - "C. 1 Batsusually fall intowinter rest in November Amphlblans(Amphibia) are regutar guests of the asdo butterflies and beetles, their main food source. DomicaCave - salamandersand frogspassively Incaves they a re rather protected against their natu- floodedinside the caveduring heavier rains. The ral enemies(manens, weasets, owls). Bats usually alarminglycoloured Salamandrc salamandra hibet- catchinsects during flight. orientation in the dark natesin numerousaggregations in the entrance spacesis enabled by echorocat or, namelythetype passageof the Gombaseckdcave. Of mammals oforientationbased on the reflection ofsupersound (lvammalia)small forms of rodents,dormice, wea- wavesthat are emittedin regularintervals and selsand martensare regularguests of the caves aresubsequently received. Thus the baGare able withan unusualsense for orientationin complete to distinguishbarriers or preyduring flight in the darkness.The martens are importantpredators of darkness.Species of thefamily Rhinolophidae emit batsin caves.Bats are the bestknown and most signalsby nose, those ofthe familyVespenilionidae, popularcave mammals. From ancient times Deoole on the otherhand, by mouth. The reflected signals arereceived by the ears.The sounds are situated, dependingon the particularspecies, within the extentof 15 - 115 kHz,so theycan only parfly be heardby the humanear. Echolocatjon signals emit- ted bybats are usually adapted to flightconditions and spacestructures that considerablyimpede theiridentification using "BAT" detectors - oevtces enablinganalysis ofthe batsounds.

Danubecated .ewt - IlituG dobrogtcls, Odne Cave.fto0oj p Llpnatt

- Common frog Ra.a temporatiat Doni@ Caw ronSJeSgedbar ri.l - rxods v6per| ionis rna,el, exte/ra,pa€site of

105 Guano pile in the Domica Cave.Photo: A. No6ko@

Gteater hoseshoe bat in detait, Photo: P. LuotAeik

individuals.The caveis one of the northern-most distributionsites ofthis batsoecies in Eurooe.In the neighbouringCertova diera Cave, part ofthe Domica system,the southernbent'wing bal Minioptetus schreibersiiwas very abundant in the past.lt istypi- Auruhn coloty af the EtearethoAeshoe bat - Rhinolophus feriumequnum. Pholoi P Llpbaik calofthis speciesthat in moreextensive karst areas it inhabitsonly a few caveswith favourableambi- ent conditions.lt is an extraordinarysocial species Consideringthe caveanimals it is impossibleto livingin coloniesduring the wholeyear round. In omitthe surfaceparasites of batsthat aresucking the above'mentionedcave it createdcolonies with theirblood - ticks,bugs, fleas and tinywingless 300 - 9,000 individuals.Thick guano deposits are fliesof the familyNycteribidae. Sulface mammal the traces of their activityin the cave. However, speciesare the otherhosts of theseparasites. Bat since 1989 hibernationof this specieshave not tick /xodesvesperti,ionis is the exceptionsince it beendetected, the possibleconsequence of instal is unableto liveoutside the cavesso that it is an lationof an unsuitablegate on the caveentrance obligateparasite of bats. Similarlyto other cave in the precedingyears. The JasovskdCave hosts animalsit hasstrikingly elongated legs. a highnumber of bat species.lt servesas the most The DomicaCave is well-knownfor its bat colo importanthibernation place for the greaterhorse- nies. In the cave we may observelarger guano shoe bat Rh,noioohusfenumeouinum in Slovakia pilesand guanopots - smallerspheroidal cavities with250 individuals.The Dobsinsk6 lce Cave repre' evolvingon the sintersas a resultof the actionof sentsone ofthe mostimportant hibernation sites of acidicbat excrements.In the Domicacave system the prevailinglycold adapted forest species Myot s 16 bat specieswere registered. The most important myotisa^d M. brardtii in CentralEurope with 422 is the lvlediterraneanhorseshoe bat Rhinolophus individualscounted in 1999. In glaciatedpafts of euryalepopulating the caveduring the wholeyear the cavea populationofthe northernbat Eptesicus and establishinglarge coloniesof up to 2,000 nilisonidominates.

106 BASICMETHODS OF CAVE FAUNA isdeposited.The mesh is putovera funnelfixed into RESEARCH a stand,a vesselwithconservation liquid is situated underthe funnel.With progressivedrying of the Caveinvertebrates are most often collected substratethefauna tries to escapeby movingdown directlyby visualsearching using pincers, brush towardsthe mesh and fallingthrough it into the ves. or gxtausto.,the deviceby whichwe areable to selwhere it issubsequently fixed. sucksmaller individuals into a vesselwith fixation Aquaticinvertebrates are collectedwith tweezers. iiquid.Baits (pieces of meat,cheese, fruits), usually Smallerforms may be co ectedby filtering of water depositedon the placefor longertime, are used to throughthe planktonicnet. The device consists of attractfauna. The majority of smalleranimals are softsilk cloth put on a circularframe with a handle. Thenet progressivelynarrows down into the neckof a smallplastic vial. In the vadosezone the water is filteredwith such a netfrom water pools and perce latingwater.Inlargerlakesand undergroundstreams theplanktonic net is usedjn thesame way as in the catchingof waterfauna in abovegroundhabitats. In suchmicrohabitats the densityof the faunais ratherlow' so collectingwith the nethas to bedone repeatedlyin different hydrological conditions during thet€ar lt isalso necessary to watch the surface of cavebottoms and walls for the aOuatic tuuna. Baits areused for attraction and subsequent collection of thewater fauna. During collecting we should keep in mindthe ethicalside of the investigationsand the Cottectionof terest at [email protected]]s,Dohica Cave,photo: p. Lupteeik aspectof the faunaconservation. Thus, it is highly desirableto modirymethods to preventremarl€ble lossesinthefauna populationsand to prevent distur, banceof theexisting equilibrium between particular componentsof the subterranean slstem. Occasionalcollecting of invertebratesin a cave cannotin factseriously threaten the subterranean communitiessince the c€ve is onlya windo{into an ammensesystem of crevices inthe limestone bedrock. Thus,the organisms found in the ca\r'e represent just a minutefraction of whatreally lives in the wnote karstmassif. Massive death of the c€vefauna may be causedby natural disturbances. For example, during hea\,yrains thousands of animalsare flooded out of the caveswhere they die in the surfacewaters. calection of aquattcfauha, Dobainske be cave.Photo: p t:u aajk Explorationsof bats are carriedout by three basicmethods. The first one is the directcounting. Howeverthis methodis difficultto usein batsDe- fixedand transported in vialswith ethyl€lcohol or ciesor particularindividuals preferring inaccessible formaldehyde,beetles in vesselswith wood shav- or hardlyaccessible spaces or crevicesfor resting. ingssaturated by ethylene acetate. Pitfall trapping Thefurther method is identificationofspecies using 'BAT" is anothercollecting method - a plasticvessel with detectorsbased on the characteristicsuoer- fixationliquid is duginto the softercave sediment soundsignals of the particularspecies. N4arginally, (loam,smaller stony debris) so thatthe sediment is as the additionalmethod, collecting into the nets levelledwith the vessel upper margin. For co ecting is sometimesused. The method is, however,tess of tinyanimal forms dwelling in bat guanoor rot- suitablesince it canstress the bats and may appar- ten wooda photo€clectorapparatus is used.The entlyinfluence the behaviour ofthe lnvestigated bat deviceconsists of the mesh on which study material colonyin the cave.

707 CONSERVATIONOF THE CAVEBIOTA waters. Similarly,negative effects are obserued when soent carbideor batteriesare deoositedin Thedistribution of the subterraneanorganisms is caves.Excessive organic substances originating in in fact not limitedto caves,it also coversepikarst thecattlefarmingor inthe rinse ofchemicals during (the systemof crevicesin disconnectedrock under wintersalting of roadsmay threaten water biota of surfacesoil layers),submerged spaces in the the subterraneanhabitats. Construction of damsin ohreaticzone, as well as micro-soacesbetween the watershedcauses changes in the profileof the particlesof softer sedimentsof the subterranean riverflows and in the accumulationof sediments. running waters (interstitialmicro-habitats). Since Interstitialmicro-environments with uniquefauna the resourcesof the subterraneansystem originate maybe completelydestroyed by fine clayparticles on the surfacefrom the oroductionofthe olantsand originatingin damssince they fill up minutespaces animals.the conseruationshould also coversurface betweensand and gravelparticles. Populations of habitatsdirectly connected with the cavesystems. caveinvertebrates may be threatenedby intensive Conservationofthe cavebiota results mainlyfrom collectionusing pitfall traps, especially if they are the fact that manyspecies of organismsare "living left in the cavefor longerperiods. fossils".They disappeared from the surfacewaters Dropin populationsof particularbat speciesis andterrestrial habitats millions years ago surviving causedespecially by disturbanceduring hiberna- in caves,underground waters and other types of tion and nursingof the young,local vandalism, subterraneanhabitats thus protectedagainst the lossof habitatby closingof cavesand minesor by harsh weatheroutside. The other reasonfor the changeof the cavemicroclimate by the openingof protectionis the uniqueness,rarity and increased new entrances.change of the landscapesurface vulnerabilityof the cavernicolousfauna, especially with decreaseof plant diversityand subsequent endemicforms and troglobites. Thefollowingspecies decreaseof insectsthat serveas the foodbasis has deservespecial protection in our caves:palpigrade negativeeffects upon bat populations.Moreover, Eukoeneniaspe/aea, pseudoscorpion Neobisium batsmay bethreatened directly byexcessive collect- s/ovacum,millipede of the genusTyphloiulus and ingfor scientificpurposes. collembolansPseudosinella aggtelekiensis, P pacl- Severalinternational Conventions can be aoolied ti, Arrhopalitesaggtelekiensis and A. irtermedius to subterraneanspecies and habitats,for example, since they representunique forms with very lim- the RioConvention (13 June,1992) for biodiversity ited distribution.The vulnerability of theseanimals protection.In Europethe EonnConvention (23lune, resultsfrom their lowecological flexibility and small 1979)for theconservation of wildmigratory species distributionranges. Thus, they are threatened even is relevantto troglophilousbats. Law no. 543/2002 by localchange of the ecologicalconditions or by of the NationalCouncil of the SlovakRepublic on pollutionof the environment. the conservationof natureand the landscapealso Directdestruction ofthe caveenvironment in lime- aooliesto all caveanimals. Based on this law it is stonequarries is the most seriousdanger for the forbiddento interruotDrotected animals in their subterraneanorganisms. In this way,the localityof naturaldevelopment, especially to kill,injure, catch the first discoveryof the subterraneancarabid bee- or movethem. Moreover,it is forbiddento destroy tle Duvaliusbokori (Leontina Cave near Gombasek) and to harm their biotopesand dwellings.Many almosttotally disappeared. Chemical pollution is the formsof the cavefauna are directly included in the mostdrastic for the waterfauna since water repre- listof protectedanimals. However, we should realize sentsa mediumof fasttransportfor pollution agents. that orderingof the givenspecies on the list of the Theseare mainlypesticides applied in agriculture, orotectedfauna does not havemuch imDortance if pollutionby petrolsubstances during accidents on we are not ableto guaranteeprotection of the par' water-coursesand pollutionof watersby heavymet- ticulararea, the wholesystem with its components alsor aggressivecoloration substances used during whichis populatedby this faunaand on whichit is investigationsof the hydrologyof the underground essentiallvdeoendent.

108 CAVES OFTHE WORLD HERITAGE IN SLOVAKIA

STATENATURE CONSERVANCY OF THESLOVAI( REPUBLIC SLOVAKCAVES ADMII{ISTRATION, LIFIOVSI(V MIKUIAS Edltors:DoG Rl{Dr lo[bf Jakiil, Dlsq I Rl{Dr.Pavel B€lla, PhD.

Author$Rl{Dr. Pavel E€lla, PiD. - SlovakCales Admlnlstr€tlon, Uptovski MikulSS RNor.fudovlt Ga6l- Slov€kCav€6 Admlnlstr8tion, Llptot6ld MlkuliiS Mdr.Dadrnar Havlarov6 - slo\€k Cdv€sMmlnlsuatlon, Llpbvsky Mlkul6a Ing.Joaf Hlae6d- slovakca\,€s Admlnlsuation, llpto6k, Mlkul6S Doc.Rt{Dr, .rozef jak6l, DEc, - GeographlcalInsttute of SqS,Btatislava Doc,RNDr. tubomlr Kov6d,csc. - Facultyof llatutal scl€nc€s,P. J. Saf6rlkUntu€rslty, Ko6lce Ing lrarcel Lall(ovL,CSc, - Facultyof NaturdlScl6nce5, M. B€l Unlw6rty.Bansk6 Stlavnlca PhDr.Mad6n Soldlq PhD.- ArcheologcalInsd$le of SAS,SplSskS Nor6 Ves Rl{Or.J5n Zellnka - SlovakCav€ Admlnistratlon,Llpto/sld MlkulSS

Revisver$RNDr. v6clav Cllek, CSc., Prof. PhDr Vliclav FumSnek, Drsc., Doc.RNDi. zdenko Hochmdh, csc., RNDr.vladlmlr K0d€1, csc., Doc.RNDr. Mllan L'apin, csc., RNDr.Ladlslav No\otny, Ing MlkulSSRozloznlk

Ceer pagBphotogapk: GombasecMCa!€. Photo: M. Rengevld Hruaovsk6cave. Photo: l. stanloue

lssuedbry O S'Hs NatureCorcervancy ofthe SlovakRepubllc, Slo/ak Cav6 Admlnlstatton,UptovBld Mlkul6s in 20Oa In KnEn6cemrum, Prcdm6tsk6 51, 010 0l Zlllna Redacuon:Mgr. Bohuslav Kortman T€nslatlon:Ing FbterGatlk, D,oc. RNDr. Lubomii yor/64, (S6. corecuons:Martln c. styan Graphlc$Ing. J6n f€s6k Prlnt PolWIafpdnr, Preao,

|sB Cra-AO-aOA4-308-4 CONTENS

Foreword(lozef Hlav66) 5 lntroduction(JozefJak6l, Pavel Bella)...... 9 SlovakKarst National Park - RemarkableLandscape onthe Surface and Ljnderground (Pavel Bella, JozefJak6l)...... lL SolubleRocks-TheEnvironmentforKarstandCaveDevelopment(LudovitGa6l),...... !7 Water- CreativeAgent of Karstand Caves (JozefJakSl, Pavel Bella) 25 TheSurface l\4orpholog/ of Karst Landscape (Jozef Jak6l)...... Passages,Labyrinths and Verticals underthe Earth's Surface (Pavel Bella)...... 47 TheStone Beauty of Caves(Ludovit Ga6l) 67 CaveMicroclimate (J6n Zelinka)...... a7 CaveLife(LubomirKov66) ...... 95 CaveSettlement (l\4ari6n Soj6k)...... 109 Historyof Knowingand Discoveringthe Caves (l\4arcel Lalkovii)...... 123 Thelvost lmportant Caves and Abysses of theWorld Heritage (Pavel Bella) ...... 139 Protectionof Karst and Caves (JozefJak6l, [udovit Ga6l, Dagmar Haviarov6) ...... SelectedLiterature L67