Original Scientific Article UDK 591.9:551.442(497.571) Received: 21-3-2012 SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES Slavko POLAK Zavod Znanje, OE Notranjska Museum Postojna, SI-6230 Postojna, Kolodvorska c. 3, Slovenia E-mail: slavko.polak@notranjski-muzej.si Jana BEDEK & Roman OZIMEC Croatian Biospeleological Society (CBSS), HR-10 000 Zagreb, Demetrova 1, Croatia Valerija ZAKSEK University of Ljubljana, Biotechnical Faculty, Department of Biology, SI-1000 Ljubljana, Vecna Pot 111, Slovenia ABSTRACT The subterranean fauna of twelve Istrian caves, six in Slovenia and six in Croatia, is presented in this paper. Caves in the carbonate region of White Istria (Cicarija) host the richest troglobiotic fauna in comparison to caves in the carbonate littoral region of Red Istria and the intermediate flysh region of Grey Istria. Caves in the contact zone of limestone and flysch situated in Grey Istria are inhabited by a surprisingly high number of troglobiotic taxa as well. Some subterranean taxa were found relatively far from their known distributional areas. In addition, several new taxa have been discovered for science. We briefly describe the general ecological conditions within the caves and estimate potential threats to their faunas. Istrian caves are inhabited by numerous endemic troglobiotic species, in this respect ranking this region close to the most diverse Dinaric region. Key words: subterranean fauna, troglobionts, cave conservation, Istria FAUNA SOTTERRANEA DI DODICI GROTTE ISTRIANE SINTESI L'articolo presenta la fauna sotterranea di dodici grotte istriane, sei delle quali presenti in Slovenia e sei in Croazia. Le grotte della regione carbonatica dell'Istria Bianca (Ciceria) ospitano la fauna di troglobi piu ricca in confronto alle grotte della regione carbonatica costiera dell'Istria Rossa e della regione flyschoide intermedia dell'Istria Grigia. Pe-raltro, anche nelle grotte della zona di confine fra calcare e flysch, situate nell'Istria Grigia, e stato trovato un numero sorprendentemente alto di taxa di troglobi. Alcuni taxa sotterranei sono stati trovati relativamente lontano dalle loro aree di distribuzione conosciute. Nell'articolo vengono inoltre segnalati alcuni taxa nuovi per la scienza. Gli autori descrivono brevemente le condizioni ecologiche generali all'interno delle grotte, e valutano i fattori che potrebbero nuocere alla fauna locale. Le grotte istriane ospitano numerose specie endemiche di troglobi, al pari della ben diversa regione dinarica. Parole chiave: fauna sotterranea, troglobi, tutela delle grotte, Istria Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 INTRODUCTION The Istrian Peninsula consists of three main ge-omorphological units extending from (in order from northeast to southwest) the inner, partly mountainous limestone region known as White Istria; the external, low limestone plateau with typical red soils called Red Istria; and a hydrological barrier flysch zone in between known as Grey Istria (Krebs, 1907). Up to now, over 2000 caves have been registered in Istria, mostly in limestone. There are also caves which have developed in the contact area between flysch and limestone. The Istrian Karst is relatively well isolated from the main Di-naric Karst, and for this reason caves in Istria are very interesting for biological research. Although biospeleo-logical exploration in Istria began early in the second half of the 19th century, the subterranean fauna of the area is not well known and has not been systematically explored and evaluated. Unfortunately, caves, springs and their fauna are under strong pressure from human activity (Sket, 1999; Culver & Pipan, 2009; Ozimec et al., 2009; Polak & Pipan, 2011). Geopolitically, the Istrian peninsula is divided among Croatia, Slovenia and Italy, a fact which necessitates international cooperation. Within the Karst Underground Protection project (KUP), biospeleolo-gical surveys, identification of potential threats to the Fig. 1: Map of Istria showing locations of the12 caves investigated. Sl. 1: Zemljevid Istre z lokacijami 12 raziskovanih jam. subterranean environment and accompanying popularization activities have been carried out (Ozimec et al., 2010, 2011). This project was financed by the OP IPA Slovenia - Croatia 2007-2013 program and led by the Istrian Region, with Natura Histrica as the Croatian partner and the Karst Research Institute ZRC SAZU from Postojna as the Slovenian partner. The aim of the speleobiological research within the framework of the KUP was to evaluate species richness and the ecological conditions of the subterranean habitats in Istria through a systematic survey of selected caves. In this paper we present an overview of the faunistic findings within twelve selected caves from Slovenian and Croatian Istria. Caves under study In total, twelve caves were selected for investigation, six in Slovenia and six in Croatia (Fig. 1). Six of them were in Čičarija (White Istria), three in the contact zone of limestone and flysch (Grey Istria) and three caves were in the littoral region of Red Istria. Some of the selected caves provide terrestrial and aquatic habitats (springs, subterranean rivers and lakes), whereas others feature only terrestrial habitats. The caves differ in terms of accessibility and use: some are accessed through an entrance shaft and are more difficult of access to people, while others are horizontal and regularly visited. Two of these caves have been used for tourism or by the military. In the following, we briefly present the caves being investigated. Cadastre Number (Cad. No.) refers to the corresponding Slovene cave cadastre. Number (Pecina br.) corresponds to the number of Istrian caves (Malez, 1960) and number No. VG (Grotte della Venezia Giulia) refers to the old Italian cave cadastre (Boegan, 1930). 1. Polina Peč (Cad. No. 938), Poljane, Obrov, Matar-sko podolje, Čičarija, Slovenia Synonymy: Caverna di Pogliane presso Castelnuovo, Polina pecina, No. 1105 VG (Müller, 1930). With its entrance at the bottom of a small doline, this cave is easily accessible. It is 365 m long and 40 m deep, consisting of a large descending entrance hall, a narrow passage and a huge inner chamber. The temperature of the entrance hall is strongly influenced by cold air, especially in winter when ice is regularly deposited to the very bottom. 2. Račiška Pečina (Cad. No. 942), Račice, Starod, Matarsko podolje, Čičarija, Slovenia Synonymy: Pečina pri Račicah, Grotta di Racizze, No. 613 VG, presso Castelnuovo (Müller, 1930). The cave is horizontal and 304 m long by 29 m deep. Between World War I and II and until the 1980s, the cave was used as a fuel storage place for military purposes. For this reason, some parts of the cave have been devastated; the floor has been artificially levelled and the cave is partly polluted with oil. In the inner part, Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 bones and traces of a cave bear (Ursus spelaeus) have been discovered (Mihevc, 2003); due to these discoveries, the cave has been closed for conservation purposes since 2007. 3. Dimnice (Cad. No.735), Markovscina, Materija, Matarsko podolje, Cicarija, Slovenia Synonymy: Grotte de Markovsina ou grotte d'Innice, Höhle bei Markovscina (Müller, 1905), Grotta Dimnice nei dintorni di Markovscina (Müller, 1906), Rauchgrotte bei Markovscina (Perko, 1909), Grotte Dimnice ou grotte qui fume, a Markovsina (Jeannel, 1911), Grotte di Marcossina detta Grotta del Fume, No. 626 VG, Grotta Dimnizze presso Slivje (Müller, 1928), Dimnica jama (Rauchgrotte) bei Slivia (Stammer, 1932). The total length of the passages is 6020 m; the cave is 134 m deep. The Dimnice cave is the only tourist cave included in our study. Down the 39 m deep entrance pit, the cave extends into horizontal passages and chambers of impressive dimensions, some of them being 20 m wide and high. Relatively small water current flows through Dimnice. 4. Medvedjak (Cad. No. 881), Markovscina, Materija, Matarsko podolje, Cicarija, Slovenia Synonymy: Medvedova jama, Grotta dell' Orso presso Markovsina (Perko, 1909), Medvedova jama bei Markovsina (Müller, 1913), Grotta dell'orso (medvedova jama) dist. Marcousina (Müller, 1928), Grotta ai pie-di del Monte Medvediak, No. 70 VG, Grotta dell 'orso presso Marcossina, tra Cosinae Castelnuovo (Müller, 1930). The total length of the cave is 1091 m with a depth of 129 m. The entrance section of the cave is a 45 m deep bell-shaped pit with a huge hall beneath. From this point, more or less horizontal galleries extend in different directions. In the upper parts there are many speleothe-mes, whereas the lower galleries are mostly muddy. Jama Pod Krogom (Cad. No. 4524) - Spilja pod Kru-gom (Bedek et al., 2006)., Mlini, Socerga, Cicarija, Slovenia-Croatia 5. The Jama Pod Krogom spring cave is situated in the contact zone where the Paleocene limestone thrusts over the Eocene flysch. The cave is 570 m long and ascends for 25 m. The entrance opens on a vertical rock cliff, right on the Slovene - Croatian border. During rainy periods, the whole cave is flooded and water flows out at the entrance. During dry periods the water is caught in the cave lakes and siphons several meters above the level of the permanent spring in the village of Mlini (Ha-bic et al., 1982). 6. Kubik (Cad. No. 4524), Brezovica pri Gradinu, Koprska brda, Slovenia Synonymy: Kobiak. The Kubik cave is a 292 m long horizontal cave of small dimensions. The entrance opens into a small pit. In its farthest and lowest section where a small stream and water pools occur, the cave is 10 m below the surface. The cave lies in on the contact between flysch san- dstones and nummulite, turbidite limestone originated by small stream erosion. Rabakova Špilja, Roč, Ročko polje, White Istria, Croatia Synonymy: Kerbauc pečina, Ročka Špilja, Grotta di Rabaco dei Bencici, No. 1193 VG, Pečina br. 4 (Malez, 1960). 7. Rabakova Špilja is 143 m long and 22 m deep (Malez, 1960). The cave was formed by the erosion of nummulite, turbidite limestones caused by periodic water flow. It is a complex cave system consisting of two morphologically different channels with periodic water flow in the lowest part. 8. Radota Jama, Radota brdo, Rakitovec, Brest, Bu-zet, White Istria, Croatia Synonymy: Grotta di Garnchino (Radetik), No. 298 VG, Pečina br. 42 (Malez, 1960). 8. Radota Jama is located on the southwestern side of the Radota hill in Čičarija. It represents a simple cave system, 268 m long and 170 m deep (Malez, 1960). In the eastern part of the main cave chamber, a steep, partly vertical 150 m long channel starts, with an intermittent water pool on the bottom. The channel is completely covered with sinter. 9. Piskovica, Jakliči, Gologorica, Pazin, Grey Istria, Croatia Piskovica is mostly a horizontal, 1036 m long passage, representing a simple, hydrologically active cave system. Two dozen meters before the end of the cave at its deepest point of 38 m, there are collapsed blocks with a water current flowing through. Piskovica cave was formed by water flow erosion of flysch sediments located between limestone layers. The cave has a unique and attractive morphology resembling a straight rectangular tunnel (Jekič & Zlokica, 1988). Markova Jama, Štancija Špin, Tar, Poreč, Red Istria, Croatia 10. Markova Jama is a complex 291 m long and 82 m deep pit system with two entrances, one of them closed at the moment. The cave was discovered in 1975 (Legovič, 1985). The deepest point at 82 m is 18 m below sea level; at this point a 3 m deep lake is located. 11. Jama Kod Buriči, Kanfanar, Red Istria, Croatia Jama Kod Buriči is a complex, 100 m long and 127 m deep pit, with a narrow (0.5 x 0.7 m) entrance. A long inner chamber extends into several muddy pits. Jama Kod Buriči is hydrologically active during heavy rains (Legovič, 1999). 12. Pincinova Jama, Tar, Poreč, Red Istria, Croatia. The entrance of the cave Pincinova Jama is located at the bottom of a small doline in the vicinity of the village of Monfaber near Poreč. The cave is 100 m long, with the deepest point at 85 m (Legovič, 1985). It is a simple pit with an underground water pool. The lake is, on average, 22 m deep. Since its discovery in 1976, many speleological research studies have been carried out within the cave. Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 MATERIALS AND METHODS At the beginning of our research, published information on the caves with respect to their geomorphology, physical properties and faunas was compiled and studied. The speleological data, including the cave maps of Slovene caves were obtained from the Slovene Cave Cadastre (IZRK ZRC SAZU and the Slovene Caving Association). The maps and data for the Croatian caves were obtained from published reports (Malez, 1960; Le-govic, 1985, 1999; Jekic & Zlokolica, 1988). Older cave catalogues (Bertarelli & Boegan, 1926; Garibaldi, 1926; Boegan, 1930) were also studied. The published fauni-stic records were taken from the cave fauna catalogues (Hamann, 1896; Wolf, 19341938) and from numerous papers and revisions of particular animal taxa. Field work in the selected caves was carried out during between two to four visits (at least two per cave) in different seasons. Measurements of environmental parameters are irrelevant for this study and are not presented here. The fauna was studied by systematic visual inspection of the cave floor, walls, ceiling and water micro-habitats. Sampling with baited pit-fall traps was not applied, except for one occasion in the Medvedjak cave to collect beetle specimens for photographic documentation and molecular studies. Most of the recorded species were photographed. Selected specimens were collected and preserved for further investigation. The biological samples from Slovenia are deposited in the Zoological Collection of the Department of Biology (Biotechnical Faculty, University of Ljubljana) and in the Notranjska Museum Postojna (Postojna). The material from Croatia is deposited in the Collection of the Croatian Biospe-leological Society (HBSD). The collected fauna were mostly identified by the authors although some species were delivered to other experts. We used classical classifications of subterranean environments and their associated fauna (e.g., Schiner, 1854; Camacho, 1992; Sket, 2008; Culver & Pipan, 2009, Novak et al., 2012). Troglobionts (stygobionts in the case of aquatic taxa) complete their life cycle in a completely dark, humid/water and thermally stable hypogean environment. Most of these clearly show tro-glomorphism. Troglophiles alternate between epigean and hypogean habitats or live permanently in subterranean habitats, and show some moderate adaptations to subterranean conditions, such a reduced eyes and adaptations to compensate for the lack of visual orientation. Some among these do not complete their life cycle underground, while others do. Trogloxenes are taxa which enter caves for shelter or feeding opportunities, but which exhibit no morphological adaptation to the hypogean environment and do not complete their life cycle there. Some edaphobionts living in soil are also common in the entrance sections of caves. Most of our field work focused on the troglobionts. Troglophiles and particularly trogloxenes were not sampled systematically. Small aquatic invertebrates (Turbellaria, Oligochaeta etc.), microscopic crustacean fauna (Copepoda), micro-organisms and fungi were not sampled and are not considered in this study. Owing to the lack of experts in some taxa and the absence of adult stages, some taxa have not been determined to the species level. The lists of the subterranean faunas of the three regions - White, Grey and Red Istria - were tested for differences in the presence or absence of species using the Cochan Q test (Sokal & Rohlf, 1995), where 1 indicates the presence and 0 the absence of a species. RESULTS Our findings on the subterranean faunas, as well as the faunistic records from literature about these studied caves are summarised in Table 1. 1. Polina Peč The fauna was investigated on 19.10.2010, 4.3.2011 and 22.6.2011. The cave is the type locality of the trechine beetle Anophthalmus schmidti istriensis (Müller, 1909) and the troglobiotic pselaphine beetle Mahaerites novissimus (Nonveiller & Pavicevic, 2001). Apart from the original descriptions, the only published faunistic data from Polina Peč were reported by Polak (1997). During our investigations the presence of the trechine Typhlotrechus bilimeki istrus (Fig. 5) and the sphodrine Laemostenus cavicola cavicola were stated as common; however, surprisingly, no leptodirine Leptodirus hochenwartii reticulatus was found. Some forgotten pit-fall traps left by collectors were found. So far, 15 troglobionts have been recorded from this cave (Tabs. 1, 2). Among other invertebrates, our discovery of the cave centipede Eupolybothrus obrovensis is significant, since only a couple of Slovenian localities have so far been identified (Matic & Darabantu, 1968). In this cave the troglomorphic diplurian Plusiocampa cf. nivea (Fig. 7) is common. A small number of the woodlice Titanethes dahli and Alpioniscus cf. strasseri were also found. 2. Račiška Pečina Before the military use of the cave, some biologists (J. Stussiner, J. Müller, K. Strasser) visited the cave and found the trechine beetle Typhlotrechus bilimeki istrus and the oniscoid Mesoniscus graniger (Verhoeff, 1933). The only published data on the fauna of Polina Peč were reported by Polak (1997). Our investigations on 20.10.2010 and 4.3.2011 resulted in a list of 16 troglobionts (Tabs. 1, 2). This cave is the second known locality for the leptodiri-ne beetle Prospelaeobates vrezeci (Fig. 3), previously recorded only from the type locality: the Medvedjak cave. In total, five beetle species, three crustacean, two spider and four false scorpion species were found. We found the cave spider Stalita taenaria (Fig. 8) only in the deepest part of the cave with permanently dripping water. Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Tab. 1: Checklist of the subterranean fauna of twelve caves in Istria. White ¡stria: 1 Polina Peč, 2 Račiška Pečina, 3 Dimnice, 4 Medvedjak, 5 Jama Pod Krogom, 7 Rabakova Špilja, 8 Radota Jama; Grey Istria: 6 Kubik, 9 Piskovica; Red Istria: 10 Markova Jama, 11 Jama Kod Burici, 12 Pincinova Jama. Legend: Tb - troglobiont, Tf - troglophile, Tx - trogloxene, Ed - edaphobiont, Par - parasite; * published data only. Tab. 1: Seznam podzemeljske favne dvanajstih jam v Istri. Bela Istra: 1 Polina peč, 2 Račiška pečina, 3 Dimnice, 4 Medvedjak, 5 Jama pod Krogom, 7 Rabakova špilja, 8 Radota jama; Siva Istra: 6 Kubik, 9 Piskovica; Rdeča Istra: 10 Markova jama, 11 Jama kod Burici, 12 Pincinova jama. Legenda: Tb - troglobiont, Tf - troglofil, Tx - trogloksen, Ed - edafobiont, Par - parazit; * podatek samo iz literature. Classis / Ordo Familia (Subfamilia) Genus, Species (Subspecies) Status White Istria Grey Istria Red Istria Zospeum isselianum Pollonera, 1887 Tb 1, 3, 6 6 Gastropoda / Carychiidae Z. kusceri (A. J. Wagner, 1912) Tb 2, 3* Pulmonata Z. spelaeum schmidti (Frauenfeld, 1854) Tb 4, 7, 8, 9 10 Zonitidae Oxychilus (Oxychilus) sp. Tf 9 10, 11 Polychaeta / Canalipalpata Serpulidae Marifugia cavatica Absolon & Hrabe, 1930 Tb 5 Clitellata / Haplotaxida Haplotaxidae Haplotaxis cf. gordioides (Hartmann, 1921) Tf 9 Clitellata / Hirudinea Erpobdelidae Dina krasensis Sket, 1968 Tf 9 Stalita taenaria Schiödte, 1848 Tb 2, 3*, 4 Dysderidae Stalita sp. Tb 10, 11 Mesostalita nocturna (Roewer, 1931) Tb 8 Nesticus cf. celullans (Clerk, 1757) Tf 1 9 Nesticidae N. eremita Simon, 1879 Tf 7 6, 9 10, 11, 12 Nesticus sp. Tf 5 Metidae Meta menardi (Latreille, 1804) Tf 1, 2 Arachnida / Araneae Metellina sp. Tf 2 Agelenidae Tegenaria sp. Tf 2, 6 9 Troglohyphantes excavatus Fage, 1919 Tf 1 T. cf. brignolii Deeleman-Reinhold, 1978 Tf 2, 3*, 7 Troglohyphantes sp. Tb 8 Linyphiidae Walckenaeria mitrata (Menge, 1868) Tx 9 Porrhomma cf. microps (Roewer, 1931) Tf 9 Leptyphantes sp. Tf 10, 11 Diplocephalus cf. crassiloba (Simon, 1884) Tx 10 Chthonius spelaeophilus histricus Beier, 1931 Tb 8 , Chthoniidae Chthonius sp. Tb 9 11 Arachnida / Troglochthonius doratodactylus Helversen, 1968 Tb 2, 8 11 Pseudoscorpiones Neobisium spelaeum istriacum (Müller, 1931) Tb 1, 2, 3, 4 Neobisiidae N. reimoseri reimoseri (Beier, 1929) Tb 2, 7, 8 N. reimoseri histricum Beier, 1939 Tb 3* Neobisium sp. Tb 9 Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Classis / Ordo Familia (Subfamilia) Genus, Species (Subspecies) Status White Istria Grey Istria Red Istria Arachnida / Phalangiidae Leiobunum rupestre (Herbst, 1799) Tf 1 Opiliones Trogulidae Trogulus banaticus Avram, 1971 Tf 5 6, 9 Arachnida / Eukoeneniidae Eukoenenia spelea (Peyerimhoff, 1902) Tb 4 Palpigradi Eukoenenia sp. 7, 8 Arachnida / Ixodea Ixodidae Eschatocephalus vespertilionis (Koch, 1844) Par 3, 7, 8 6 Arachnida / Mesostigmata Parasitidae Parastitus sp. Ed 8 Eugamasus cf. inferus Willmann, 1941 Ed 3 Rhagidiidae Rhagidia sp. Ed 3 9 Arachnida / Oribatida Galumnidae Liocarus sp. Ed 8 Galumna sp. Ed 8 Belbidae Belba sp. Ed 7 Arachnida / Prostigmata Eupodidae Linopodes sp. Ed 9 Hadzia fragilis S. Karaman, 1932 Tb 12* Niphargus stygius (Schiödte, 1847) Tb 2, 4 N. krameri (Schellenberg, 1935) Tb 3*, 5, 7 6, 9 Malacostraca / Niphargidae N. steueri steueri Schellenberg, 1935 Tb 5 12 Amphipoda N. stochi G. Karaman, 1994 Tb 5 N. longicaudatus A. Costa, 1851 Tb 9 N. cf. longicaudatus A. Costa, 1851 Tb 10 N. hebereri (Schellenberg, 1933) Tb 12* Titanethes dahli Verhoeff, 1926 Tb 1, 2, 3, 4, 5, 8 Alpioniscus strasseri (Verhoeff, 1927) Tb 6, 9 10, 11, 12 Trichoniscidae (Trichoniscinae) A. cf. strasseri (Verhoeff, 1927) Tb 1, 2, 5, 7 Androniscus stygius (Nemec, 1897) Tb 1, 4 A. roseus (C. Koch, 1838) Tf 6, 9 Trichoniscus sp. Tx 6 Hyloniscus sp. Tx 3 Malacostraca / Trichoniscidae (Haplophthalminae) Haplophthalmus sp. Tx 6 Isopoda Trichoniscidae (Thaumatoniscellinae) Thaumatoniscellus speluncae Karaman, Bedek & Horvatovic, 2009 Tb 4 Mesoniscidae Mesoniscus graniger (Frivaldsky, 1865) Tf 2* Ligiidae Ligidium sp. Tx 3 Trachelipodidae Trachelipus ratzeburgii (Brandt, 1833) Tx 9 Trachelipus sp. Tx 7 12 Agnaridae Protracheoniscus sp. Tx 6 Philosciidae Chaetophiloscia sp. Tx 12 Sphaeromatidae Monolistra bericum hadzii Sket, 1959 Tb 7 6 Monolistra sp. nov. Tb 9 12 Cirolanidae Sphaeromides virei virei Brian, 1923 Tb 5 12 Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Classis / Ordo Familia (Subfamilia) Genus, Species (Subspecies) Status White Istria Grey Istria Red Istria Malacostraca / Atyidae Troglocaris planinensis Birstejn, 1948 Tb 5 12 Decapoda Troglocaris sp. nov. Tb 12 Eupolybothrus obrovensis (Verhoeff, 1930) Tb 1, 3*, 4*, 8 Eupolybothrus sp. Tx 7 Chilopoda / Lithobiomorpha Lithobiidae Lithobius cf. lapidicola Meinert, 1872 Tf 7 9 L. cf. erythrocephalus C. L. Koch, 1847 Tx 12 Lithobius sp. 1 Tb 9 Lithobius sp. 2 Tx 2 Lithobiidae indet. Tx 6 Chilopoda / Cryptopidae Cryptops sp. Ed 2* 12 Scolopendromorpha Scolopendridae Scolopendra cingulata Latreille, 1789 Tx 7 Chilopoda / Goephilomorpha Linotaeniidae Strigamia sp. Ed 9 Trachysphaeridae Trachysphaera noduligera (Verhoeff, 1906) Tf 7 Diplopoda / Glomerida T. cf. noduligera (Verhoeff, 1906) Tf 8 Glomeridellidae Typhloglomeris cf. fiumarana Verhoeff, 1899 Tb 9 Macrosternodesmidae Verhoeffodesmus gracilipes Strasser, 1959 Tb 7 9 Diplopoda / Chordumatidae Melogena broelemanni Verhoeff, 1897 Tx 7* Chordeumatida Chordeumatidae Chordeumatidae indet. Tx 1 Diplopoda / Polydesmida Polydesmidae Brachydesmus inferus concavus Attems, 1898 Tb 3 B. subterraneus Heller, 1858 Tf 3, 7 9 12 Brachydesmus sp. Tf 4 Typhloiulus illyricus Verhoeff, 1929 Tb 7 9 12 Diplopoda / Julida Julidae Julus sp. Tx 8 9 Julidae indet. Tx 6 Entognatha / Diplura Japygidae Japyx cf. solifuga Halliday, 1864 Ed 12 Campodeidae Plusiocampa cf. nivea (Joseph, 1882) Tb 8 , 11 Tomoceridae Tritomurus scutellatus Frauenfeld, 1854 Tb 1, 2, 4 Paronellidae Troglopedetes pallidus Absolon, 1907 Tb 2, 4, 7, 8 6, 9 10, 11, 12 Collembola / Oncopoduridae Oncopodura sp. Tb 1, 3, 4, 7, 8 6 10, 11 Entomobryomorpha Pseudosinella sp. Tb 3 Entomobrydae Heteromurus nitidus (Templeton, 1835) Tf 7, 8 10, 11, 12 Isotomidae Folsomia sp. Tx 9 Cyphoderidae Cyphoderus sp. Tx 11 Collembola / Poduromorpha Onychiuridae Onychiurus giganteus (Absolon, 1901) Tb 8 , Onychiurus sp. Tb 9 Collembola / Symphypleona Sminthuridae Arrhopalites sp. Tx 9 Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Classis / Ordo Familia (Subfamilia) Genus, Species (Subspecies) Status White Istria Grey Istria Red Istria Collembola / Neelipleona Neelidae Neelus sp. Tf 4, 8 9 Insecta / Coleoptera Carabidae (Trechinae) Typhlotrechus bilimeki istrus (Müller, 1926) Tb 1, 2, 3, 4, 7 Anophthalmus schmidti istriensis Müller, 1909 Tb 1* A. spectabilis istrianus (Ganglbauer, 1913) Tb 3 Carabidae (Sphodrinae) Laemostenus cavicola cavicola Schaum, 1858 Tf 3, L. cavicola romualdi J. Müller, 1905 Tf 10, 11, 12 L. elongatus (Dejean, 1828) Tx 1* 2* 4* Staphylinidae (Pselaphinae) Machaerites novissimus Nonveiller & Pavicevic, 2001 Tb 1 M. kastavensis Pavicevic & Ozimec, 2009 Tb 7, 8 Machaerites sp. nov. Tb 9 Pauperobythus globuliventris Nonveiller, Pavicevic & Ozimec 2002 Tb 10 Bryaxis sp. Tx 9 Amaurops sp. Ed 7 Staphylinidae (Staphylininae) Atheta spelaea (Erichson, 1839) Tf 8 10, 11 Omalium validum Kraatz, 1857 Tx 1* Staphylinidae indet. Tx 5 6 Cholevidae (Leptodirinae) Oryotus schmidti subdentatus Müller, 1904 Tb 3* Prospelaeobates vrezeci Giachino & Etonti 1996 Tb 2, 4 Bathysciotes khevenhuelleri (Miller, 1852) Tb 2, 7, 8 6, 9 Leptodirus hochenwartii reticulatus Müller, 1905 Tb 1*, 2, 3 Cholevidae (Cholevinae) Choleva sp. Tf 5 9 11 Cholevidae (Platypsellinae) Leptinus testaceus Müller, 1817 Tf 7 Curculionidae Otiorhynchus (Troglorhynchus) cf. anophthalmoides anophthalmoides (Reitter, 1914) Tf 4 Absoloniella sp. n. Ed/ Tb? 7 Insecta / Orthoptera Rhaphidophoridae Troglophilus cavicola (Kollar, 1833) Tf 1, 2, 3, 4 T. neglectus Krauss,1879 Tf 1, 2, 3, 4, 5, 7, 8 6, 9 10, 11, 12* Insecta / Trichoptera Limnephilidae Micropterna sequax McLachlan, 1875 Tf 3* Stenophylax sp. Tf 8 Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Classis / Ordo Familia (Subfamilia) Genus, Species (Subspecies) Status White Istria Grey Istria Red Istria Tineidae Tineidae indet. Tx 8 Insecta / Geometridae Triphosa dubitata (Linnaeus, 1758) Tf 1, 2, 3, 4, 7 Lepidoptera Scoliopteryx libatrix (Linnaeus, 1758) Tf 7 6 Noctuidae Hypena obsitalis (Hübner, 1813) Tf 5 Amphipyra effusa (Boisduval, 1828) Tf 5 Limonia nubeculosa Meigen, 1804 Tf 6 Insecta / Diptera Limoniidae Limonia sp. Tf 1, 2, 3, 4, 5 9 10 Culicidae Culex sp. Tx 7, 8 9 Tipulidae Tipulidae indet. Tx 8 Mycetophilidae Speolepta leptogaster (Winnertz, 1863) Tf 1 Amphibia / Caudata Proteidae Proteus sp. nov. Tb 12 Bufonidae Bufo bufo (Linnaeus, 1758) Tx 10 Amphibia / Anura Hylidae Hyla arborea (Linnaeus, 1758) Tx 10 Ranidae Rana dalmatina Fitzinger, 1838 Tx 10 Aves / Strigiformes Strigidae Stryx aluco Linnaeus, 1758 Tx 1* Rhinolophidae Rhinolophus ferrumequinum (Schreber, 1774) Tf 1, 2, 3*, 4, 7, 8 10 R. hipposideros (Bechstein, 1800) Tf 1, 2, 3, 4, 7, 8 6 10 Mammalia / Chiroptera R. euryale Blasius, 1853 Tf 7* Barbastella barbastellus (Schreber, 1774) Tf 1* 3* Myotis myotis (Borkhausen, 1797) Tf 8 10 Vespertilionidae M. myotis / M. blythii Tf 3* M. blythii (Tomes, 1857) Tf 3* M. capaccinii (Bonaparte, 1837) Tf 3* M. daubentonii (Kuhl, 1817) Tf 3 Mammalia / Rodentia Gliridae Glis glis (Linnaeus, 1766) Tx 2, 7 Mammalia / Carnivora Mustelidae Martes foina Erxleben, 1777 Tx 1, 2* Our discovery of the pseudo-scorpion Troglochthonius doratodactylus (Fig. 9) was surprising, since there are no published records of this genus in Slovenia. In the cave, numerous troglophiles and trogloxenes can be found. In autumn (20.10.2010), 13 greater (Rhinolophus ferru-mequinum) and 42 lesser horseshoe bats (Rhinolophus hipposideros) hibernated in the cave. 3. Dimnice In the past, many naturalists and speleobiologists visited the cave and made a significant contribution to the knowledge about the Dimnice subterranean fauna. The cave is the type locality for Neobisium spe- laeum istriacum, Anophthalmus spectabilis istrianus and Oryotus schmidti subdentatus. We visited Dimnice twice (20.10.2010 and 6.3.2011) and added to the list some further troglobionts, now amounting 17 taxa (Tabs. 1, 2). We found a large population of the lepto-dirine beetle Leptoditus hochenwartii reticulatus (Fig. 2). The bat fauna of Dimnice has been regularly monitored since 1991 (Presetnik et al., 2009). During our study we spotted a colony of Daubenton's bats (Myotis daubentonii, Fig. 18) overwintering above the water current far from the tourist path. During our permanent temperature measurements we noticed significant temperature fluctuations. In winter the temperature Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 dropped below 0 oC, causing ice accumulation at the bottom of the entrance pits. 4. Medvedjak Since its description in 1904, several early cavers and cave beetle collectors (e.g., A. Perko, G. Ravasini, J. Müller) have reported on its beetle fauna. Bat observations have also been published (Presetnik et al., 2009). A new leptodirine genus and species Prospelaeobates vrezeci (Giachino & Etonti, 1996) (Fig. 3) was described from this cave. During our investigation we recorded a rich subterranean fauna, counting 15 troglobionts (Tabs. 1, 2). Our finding of the tiny terrestrial isopod Thauma-toniscellus speluncae (Fig. 14) is the first record in Slovenia and the third known locality, previously recorded only on Mt. Ucka in Istria (Croatia); its closest relatives live in Romania (Bedek et al., 2006, 2011; Karaman et al., 2009). 5. Jama Pod Krogom We visited the cave twice, on 21.10.2010 and 9.9.2011. Since the cave is active and regularly flooded, the terrestrial subterranean fauna is scarce and consists mainly of troglophiles (Tab. 1). Our finding of the aesti-vating noctuid moths Hypena obsitalis and Amphipyra effusa in the entrance chamber is important, because the first one is rarely found in caves, and Amphipyra was recorded for the first time in caves, thus showing its sub-troglophilic habits in Slovenia. The cave is one of the most important habitats for the cave tubeworm Marifugia cavatica and the cave cirolanid Sphaeromides virei (Fig. 12) in south-western Slovenia. Additionally, two species of the amphipod genus Niphargus, the cave shrimp Troglocaris planinensis and two species of terre- strial isopods - seven troglobionts in total - were recorded in this cave (Tab. 2). 6. Kubik There are no published data on this cave's fauna, even though it has been recently visited by biologists. We visited the cave on 21.10.2010 and 29.6.2011. The subterranean fauna found in the cave is listed in Table 1. Seven troblobiotic species have been recorded so far (Tab. 2). In the cave the leptodirine beetle Bathysciotes khevenhuelleri, the isopode Alpioniscus strasseri, two species of Collembola and the cave gastropod Zospeum isselianum have been recorded. Among the stygobionts, the amphipod Niphargus krameri and the isopod Mo-nolistra bericum hadzii (Fig. 13), both endemic to Istria, live in the cave stream. A recent molecular study of N. krameri showed the existence of two distinct species in Istria (Fiser, 2011). Despite some old garbage in the cave, the subterranean stream seems not to have been seriously affected by pollution. 7. Rabakova Spiija This cave is the type locality and up to now had been the only locality of the troglomorphic diplopod Verhoef-fodesmus gracilipes (Strasser, 1959) (Fig. 11). The cave has also been visited by the Slovene biospeleologists J. Bole (Bole, 1974), F. Velkovrh, B. Sket and N. Mrsic (Mrsic, 1994) and by the Dutch arachnologist C.R. Dee-leman-Reinhold in 1968 (Deeleman-Reinhold, 1978). During our research on 6.7.2010, 26.11.2010, 3.3.2011 and 26.6.2011 at least 13 troglobionts both terrestrial and aquatic, were recorded (Tabs. 1, 2). The small troglomorphic curculionid beetle of the genus Absoloniella was found in Croatia for the first time. The specimen Region Cave Habitat No. troglobionts White Istria Polina peč t 15 Račiška pečina t 16 Dimnice a, t 17 Medvedjak t 15 Rabakova špilja t 13 (14) Radota jama t 15 Jama pod Krogom a, t 7 Grey Istria Piskovica a, t 15 Kubik a, t 7 Red Istria Markova jama t 7 Jama kod Burici t 7 Pincinova jama a, t 11 Tab. 2: Number of troglobiotic taxa found in the given Istrian caves. Prevailing habitat of the cave: a - aquatic, t - terrestrial. Tab. 2: Število troglobiontov, ugotovljenih v raziskovanih istrskih jamah. Prevladujoč habitat jame: a - vodni, t -kopenski. Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 from Rabakova Spilja is significantly smaller and different in shape, compared to the closely related Absol-oniella reitteri (J. Müller, 1912) known from caves near Trieste. Since the cave lies in the middle of the village and is easily accessible, it is endangered by organic and inorganic pollution. 8. Radota Jama Before our surveys on 27.11.2010 and 27.6.2011, the speleobiologists J. Bole and B. Drovenik investigated the cave in 1968 (Bole, 1974) and R. Ozimec in 2001(unpu-bl.). Fifteen troglobiotic taxa have been recorded so far (Tabs. 1, 2). The cave represents one of the few known localities of the cave centipede Eupolybothrus obroven-sis, the palpigrade Eukoenenia sp. (unpubl.), the spider Mesostalita nocturna, the false scorpion Troglochthoni-us doratodactylus and the beetle Machaerites kastaven-sis. A large wintering colony of the greater mouse-eared bat (Myotis myotis) was found in the cave. Because of its remote location and difficulty of access, the cave has not been damaged or polluted. 9. Piskovica Before our project surveys on 29.11.2010, 3.3.2011 and 27.6.2011, R. Ozimec provided the only intensive studies of the cave fauna in 1998 and 2001 (unpubl.). It turned out that both aquatic and terrestrial subterranean faunas are extremely rich and unique. Among the collected specimens, new taxa are in the process of description. Fifteen troglobionts have been found in Piskovica so far (Tabs. 1, 2). The cave is the second known locality of the millipede Verhoeffodesmus gracilipes. Additionally, the millipede Typhloglomeris cf. fiumarana (Fig. 10) recorded in the cave, and known to date only from the Liburnian Karst, represents a significant addition to the subterranean fauna of Istria. The cave functions as a periodic sinkhole. Organic and inorganic debris are accumulated in the inner parts of the cave by intensive water flow during the rainy season. A serious threat to Piskovica cave is the intention of local inhabitants to exploit the cave for tourism. 10. Markova Jama Speleobiologically, the cave was first explored in 1997 by members of the Croatian Biospeleological Society. The cave is known as an important site of a large summer maternity colony of the greater mouse-eared bat (Myotis myotis), with over 1000 individuals. In the cave, a new genus and species of the troglobiotic pselaphinae beetle Pauperobythus globuliventris (Fig. 4) have been found (Nonveiller et al., 2002). Despite our careful research on 28.11.2010 and 29.6.2011 only seven troglobionts were recorded (Tabs. 1, 2). Markova Jama has been protected as a natural geomorpholo-gical monument since 1986. Nevertheless, the cave is endangered by organic waste disposal; sheep carcasses were found in the pit before and during our research. The cave fauna is vulnerable, especially the maternity colony of bats, which remains in the cave during spring and summer. For that reason, visits to the cave should be organized only during the period between October and March. 11. Jama Kod Burici The first speleobiological survey was carried out by members of the Croatian Biospeleological Society in May 2001. On that occasion, the false scorpion Tro-glochthonius doratodactylus was recorded in Croatia for the first time (Ozimec, 2002). During our visits on 29.11.2010 and 28.06.2011, additional species were recorded (Tabs. 1, 2), but only seven among them were troglobionts. Sheep carcasses were found at the bottom of the pit. This put a stop to further exploration. In addition, the cave is critically endangered due to organic waste, especially dead goats and sheep. Besides, the cave fauna is potentially endangered by underground water pollution because of the industry nearby. 12. Pincinova Jama Since its discovery in 1976, many speleological studies have been carried out in the cave. The endemic Istrian cave salamander Proteus sp. nov (Goricki & Tron-telj, 2006) and other important stygobionts were found there (Rada, 1980; Sket, 1994; Zaksek et al. 2009). In our visits on 28.6.2011 and 10.9.2011, eight stygobionts were recorded; besides the cave salamander, these were three amphipod species, two aquatic isopod species and two species of cave shrimp (Tab. 1). Since the dry part of the cave is short, only three terrestrial troglobionts were found. Since 1986, Pincinova Jama has received protection as a geomorphological monument. The cave is critically endangered by organic and inorganic pollution. Deposits of garbage are located less than 100 m in front of the cave entrance. At the same distance there is an old quarry, which is still periodically active. Recently the cave was closed off by gates, which are often breached by illegal visitors. A broken gate was also detected during our last visit. The caves in the carbonate region of White Istria (Cicarija) hosted 44 troglobiotic species, i.e., many more in comparison to the caves in the carbonate littoral region of Red Istria (19 troglobionts) and intermediate flysch Grey Istria (18 troglobionts). According to the Cochran test, the presence of species of subterranean fauna in White Istria differed significantly from both Grey Istria (Q test = 16.20, p < 0.001) and Red Istria (Q test = 14.69, p < 0.001), while there were no difference between Grey and Red Istria (Q test = 0.04; p = 0.835). DISCUSSION AND CONCLUSIONS Faunistic investigations of twelve caves in Istria allowed us to estimate and briefly evaluate the species Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 richness of the subterranean fauna in Istria and to provide some environmental and threat information for these caves. The most important finding was that the division of the Istrian peninsula into three main geomor-phological units - the inland White, the Red littoral and the mainly noncarbonate intermediate region of Grey Istria - accurately reflects the distribution of subterranean species as well as species richness. Caves located in the easternmost part of White Istria, in Matarsko Podolje and Čičarija, are biotically the most diverse. In each of these caves at least 15 troglobiotic taxa were found (Tab. 2). With respect to troglobiotic fauna, some of these caves (Polina Peč, Račiška Pečina, Dimnice, Medvedjak and Radota Jama) can be listed among the richest in the north-western Dinaric region. Given that the troglobiont lists are not complete, since the caves have not been investigated for microfauna and some taxa (e.g. copepods, Acarina), the number of troglobiotic species is undoubtedly greater. In contrast with Čičarija, the caves situated in the littoral region of Red Istria (Markova Jama and Jama Kod Burici) are inhabited by troglobiotic fauna only half as rich (Tab. 2). The profusion of troglobiotic fauna in White Istria reflects the close contact of White Istria with the rest of the Dinaric Karst, which is known to be the richest region in the world for subterranean fauna (Sket et a!., 2004; Culver et a!., 2006; Zagmajster et a!., 2008). Most of these troglobionts seem to be endemic to this area. Similar levels of troglobiont richness in the Red and Grey Istria regions and their composition similarity indicate that the flysch areas are also of considerable biospeleological importance. On the other hand, the noncarbonate belt of Grey Istria represents a physical barrier between the littoral Karst and the main Dinaric Karst, resulting in the endemic species and genera of the Red Istria region. The similarities between the Grey and the Red Is-trian subterranean faunas, on the one hand, and their divergence from those of White Istria, on the other, denote the very important role of Grey Istria in geological isolation and in preventing gene flow between the faunas. Other factors could influence differences in species richness among the regions. The Karst in Čičarija has been more fragmented on account of the orography (Otoničar, 2007) and is therefore more extensive in comparison to the more or less flat and unfractured littoral Karst of Red Istria. The climate of the mountainous White Istria region is more humid, providing many more Subsurface Subterranean Habitats (SSHs), like taluses and similar habitats with soil on the surface, in comparison with the more arid lowlands of Red Istria. Humidity in caves is essential for the existence of most terrestrial troglobionts, while, on the other hand, an arid climate is one of the main reasons forcing animals to seek refuge in the subterranean environment. As result of our investigations, some subterranean taxa were discovered far from their known localities, indicating that their areas are considerably expanded. The new species (subspecies) discovered belong to the crustacean genera Mono!istra and Trog!ocaris (the Istra phylogroup; Zakšek et a!., 2009), the false scorpion Ch-thonius and the Machaerites, Abso!onie!!a and Bathy-sciotes beetles. Slovenian fauna has been enriched by the finding of the recently described oniscoid genus and species Thaumatonisce!!us speluncae (Karaman et a!., 2009) in the Medvedjak cave, and the false scorpion genus Trog!ochthonius doratodacty!us in the Račiška Pečina cave. In Slovenia, the latter species had previously been found by the Italian biospeleologist Fulvio Gasparo in Štefakova Pečina (No. 1142) near Materija and in Pečina Pod Medvejkom (No. 949) near Sežana (unpub!., courtesy F. Gasparo). New localities have been found for some other troglobionts in Croatia, such as the millipedes Verhoeffodesmus graci!ipes and Typh-!og!omeris cf. fiumarana that significantly extends their known distribution. We evaluated the potential threats to these caves and their faunas. In large caves, such as the Dimnice cave, occasional visitors do not threaten the cave fauna. On the other hand, organized hikes into smaller caves, such as Polina Peč, Markova Jama and particularly Piskovica, can seriously disturb bat colonies and directly damage the subterranean habitat. The main threat to caves is the disposal of garbage and organic waste in caves. Caves near human settlements, such as Rabakova Špilja, Markova Jama and Jama Kod Burici, are especially threatened. Aquatic cave fauna is endangered by pollution of the underground water draining from the surface. So far, subterranean waters in Jama Pod Krogom, Kubik, Piskovica, Dimnice and Pincinova Jama remain sufficiently clean to host a rich stygobiotic fauna, including the endemic Istrian cave salamander in Pincinova Jama. This cave deserves special attention to avoid potential pollution from a quarry and rubbish dump in the near vicinity. ACKNOWLEDGEMENTS The long term field study was carried out in the scope of the Karst Underground Protection (KUP) project, funded by the Slovenia-Croatia Operational Programme 2007-2013 (IPA). We are grateful to our colleagues, speleobiologists Marko Lukic (Zagreb), Cene Fišer (Ljubljana) and Lubomir Kovač (Košice, Slovakia) for determinations of some taxa. We wish to thank Silvio Legovic-Kawasaki, who allowed us to stay in his spe-leo-house in Baredine (Croatia) and for valuable information and help, as well as Franci Malečkar and Janko Brajnik (JK Dimnice) for giving us access to the Dimnice Cave (Slovenia). We are grateful to Mitja Prelovšek (IZRK ZRC SAZU) and the caver Davor (Speleoklub Buzet) for technical support during difficult caving in Medvedjak and Radota Jama, as well as to the group of Croatian speleodivers (Branko Jalžic, Vedran Jalžic and Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Alan Kovacevic) who were responsible for safety sampling in Pincinova Jama. We would also like to express our gratitude to Tone Novak for helpful editorial comments and significant improvement of the manuscript, Franc Janzekovic for help with statistics and to Michelle Gadpaille for improving the manuscript linguistically. Finally, we are grateful to the local inhabitants for their kind hospitality. PODZEMELJSKA FAVNA DVANAJSTIH JAM V ISTRI Slavko POLAK Zavod Znanje, OE Notranjski muzej Postojna, SI-6230 Postojna, Kolodvorska c. 3 E-mail: slavko.polak@notranjski-muzej.si Jana BEDEK & Roman OZIMEC Hrvaško biospeleološko društvo (HSBD), HR-10000 Zagreb, Demetrova 1, Hrvaška Valerija ZAKŠEK Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo, SI-1000 Ljubljana, Večna pot 111 POVZETEK V prispevku so zbrani literaturni podatki o podzemeljski favni dvanajstih jam v Istri in rezultati lastne natančne inventarizacije teh jam. Raziskava je bila opravljena v okviru projekta Karst Underground Protection v letih 2010 in 2011. Izbrane jame so v različnih predelih Istre in se razlikujejo med seboj po okoljskih dejavnikih, po tem, da je v njih le kopenska ali tudi vodna favna ter po različni stopnji ogroženosti jam in favne zaradi človeške dejavnosti. Za raziskavo smo izbrali naslednje jame v Sloveniji: Polina peč, Račiška pečina, Medvedjak, Dimnice, Jama pod Krogom in Kubik ter na Hrvaškem: Radota špilja, Rabakova jama, Markova jama, Jama kod Burici, Pincinova jama in Piskovica. Izkazalo se je, da so glede števila troglobiontov - vrst, specializiranih za življenje v podzemlju - jame na območju Cičarije (Bela Istra) več kot dvakrat bogatejše (44 vrst) od jam v Sivi Istri (18) in litoralnem območju Rdeče Istre (19). V posameznih jamah v Cičariji živi po 15 vrst troglobiontov ali več; te vrste so večinoma ozko sorodne s favno bližnjega severozahodno dinarskega območja. V dveh jamah Rdeče Istre je bilo najdenih po 7 troglobiontov. Jama pod Krogom in Pincinova jama sta pomembni zlasti kot habitat številnih vodnih troglobiontov - stigobiontov. Jame, nastale na kontaktu apnencev in fliša v nekraški Sivi Istri, so se izkazale za nepričakovano bogate s troglobionti. Nekatera odkritja so bila presenetljiva in predstavljajo pomemben prispevek k poznavanju razširjenosti nekaterih vrst in rodov podzemeljskih živali. V prispevku prvič objavljamo podatek o prisotnosti podzemeljskega paščipalca Tro-glochthonius doratodactylus v Sloveniji. Prav tako je bil podzemeljski enakonožni rak - mokrica Thaumatoniscellus speluncae - v Sloveniji najden prvič. Najdba podzemeljskih dvojnonog Vehoeffodesmus gracilipes in Typhloglomeris cf. fiumarana v jami Piskovica predstavljata znatno povečanje znanih območij njune razširjenosti. Majhen hrošč ril- Slavko POLAK et aL: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 čkar iz rodu Absoloniella sp. je bil prvič najden na ozemlju Hrvaške. Med podzemeljskimi raki, pajkovci in hrošči je bilo nekaj taksonov že prepoznanih kot novih za znanost; njihovi znanstveni opisi so v pripravi. Za raziskane jame smo ovrednotili ekološke razmere ter ocenili sedanje in potencialne vire ogrožanja jam ter njihove favne. Ugotovili smo, da so nekatere jame organsko in anorgansko onesnažene zaradi nelegalnega odlaganja smeti in kadavrov domačih živali, potencialno pa tudi zaradi kamnolomov v bližini. Zaradi možnosti onesnaževanja podtalnih voda je še potencialno ogrožena vodna favna jam. Med potencialne grožnje favni štejemo tudi nenadzorovano nabiranje favne v jamah in v nekaterih primerih prepogost obisk jam. Ključne besede: podzemeljsko živalstvo, troglobionti, varstvo jam, Istra REFERENCES Bedek, J., S. Gottstein Matočec, B. Jalžic, R. Ozimec & V. Štamol (2006): Katalog tipskih špiljskih lokaliteta faune Hrvatske (Catalogue of Cave Type Localities of Croatian Fauna). Nat. Croat., 15 (Suppl. 1), 11-54. Bedek, J., S. Taiti & S. Gottstein (2011): Catalogue and atlas of cave-dwelling terrestrial isopods (Crustacea: Oniscidea) from Croatia. Nat. Croat., 20(2), 237-354. Bertarelli, L. V. & E. Boegan (1926): Duemilla Grotte. TCI ed., Milano. Boegan, E. (1930): Catastro delle Grote Italiane, Fasc. 1. Grotte della Venezia Giulia, Trieste. Bole, J. (1974): Rod Zospeum Bourguignat 1856 (Gastropoda, Ellobiidae) v Jugoslaviji. Razprave SAZU, Cl. IV, 17(5), 249-291. Camacho, A. I. (1992): A classification of the aquatic and terrestrial subterranean environment and their associated fauna. In: Camacho, A. I. (ed.): The natural history of biospeleology. Monografias, Museo nacional de ciencias naturales, Madrid, pp. 59-103. Culver, D. C, L. Deharveng, A. Bedos, J. J. Lewis, M. Maden, J. R. Reddelli, B. Sket, P. Trontelj & D. White (2006): The mid-latitude biodiversity ridge in terrestrial cave fauna. Ecography, 29, 120-128. Culver, D. C. & T. Pipan (2009): The biology of caves and other subterranean habitats. Oxford University Press, New York, 254 p. Deeleman-Reinhold, C. L. (1978): Revision of the cave-dwelling and related spiders of the genus Troglohy-phantes Joseph (Linyphiidae), with special reference to the Yugoslav species. Razprave SAZU, Cl. IV, 23(6), 1-221. Fišer, Ž. (2011): Znotrajvrstna raznolikost istrske slepe postranice Niphargus krameri. Diplomsko delo. Biotehniška fakulteta, Ljubljana, 33 p. Garibaldi, C. I. (1926): Catastro delle cavita naturali sotterranee della Venezia Giulia. L'Instituto Geografico Militare di Firenze, Firenze. Giachino, P. M. & M. Etonti (1996): Prospelaeo-bates gen. nove due sp.n. di Leptodirinae delle Isole del Quarnero e dell'Istria (Coleoptera: Cholevidae). Acta Entomol. Slov., 4(2), 63-71. Goricki, S. & P. Trontelj (2006): Structure and evolution of the mitochondrial control region and flanking sequences in the European cave salamander Proteus an-guinus. Gene, 378, 31-41. Habic, P., R. Gospodaric, A. Mihevc & F. Sustersic (1982): Movraska in Smokavska vala in Jama pod Krogom. Acta Carsologica, 11, 77-97. Hamann, O. (1896): Europäische Höhlenfauna. Eine Darstellung der in den Höhlen Europas lebenden Tierwelt mit besonderer Berücksichtung der Höhlenfauna Krains. Costenoble, Jena. Jeannel, R. (1911): Biospeologica XIX. Révision des Bathysciinae (Coléoptères Silphides). Morphologie, distribution géographique, systématique. Arch. Zool. Expérimentale Gén., (5)7, 1-641. Jekic, M. & M. Zlokolica (1988): Pecina Piskovica, Bilten SpeleoBih, 12, 69-78. Karaman, I. M., J. Bedek & M. Horvatovic (2009): Thaumatoniscellus speluncae n. sp. (Isopoda: Oniscidea: Trichoniscidae), a new troglobitic oniscid species from Croatia. Zootaxa, 2158, 57-64. Krebs, N. (1907): Die Halbinsel Istrien Landeskundliche Studie, Geograph. Abhandl. hg. v. A. Penck IXj2, Leipzig. Legovic, S. (1985): Nova istrazivanja Markove jame. Bilten SD Proteus, 4, 3-6. Legovic, S. (1999): Jama kod Burici, Kanfanar, Istra. Speleozin, 7(11), 12-17. Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Malez, M. (1960): Pečine Cicarije i Učke u Istri. Acta Geologica, 2, 164-260. Matic, Z. & C. Darabantu (1968): Contributions à la connaissance des Chilopodes de Yougoslavie. Razprave SAZU, Cl. IV, 11, 201-227. Mihevc, A. (2003): Sledovi jamskega medveda v Račiški pečini. Naše jame, 45, 48-55. Mršic, N. (1994): The Diplopoda (Myriapoda) of Croatia (Dvojnonoge (Diplopoda: Myriapoda) Hrvatske). Razprave SAZU, Cl. IV, 35(12), 219-296. Müller, J. [G.] (1905): Vier neue Höhlenkäfer aus dem österreichischen Litorale. Wien. Ent. Zeitg., 24, 32 p. Müller, J. [G.] (1906): Nuovi coleoteri cavernicoli del Litorale. Il. Tourista, 11(1904), 12-15. Müller, J. [G.] (1913): Beiträge zur Kenntnis der Höhlenfauna der Ostalpen und der Balkanhalbinsel, 2. Revision der blinden Trechus-Arten. Denkschr. Akad. Wiss. Wien, 90, 1124. Müller, J. [G.] (1928): Il genere Orotrechus Mülle., Jeannel. Contributo ala conoscenza della fauna cavernicola friulana. Boll. Soc. Entomol. Ital., 60(6), 92-98. Müller, J. [G.] (1930): I Coleoteri cavernicoli Italiani. Elenco geografico delle Grotte con indicazione delle specie e varietà dei Coleoteri cavernicoli finora trovati in Italia. Le Grotte d'Italia, 4, 65-85. Nonveiller, G. & D. Pavicevic (2001): Description d'une sous-espèce nouvelle et de six espèces nouvelles du genre Machaerites Miller, 1855 de Slovénie et de Croatie (Coleoptera, Pselaphinae, Bythinini). Nouv. Rev. d'Entomol., 18(4), 317-333. Nonveiller, G., D. Pavicevic & R. Ozimec (2002): Description d'un nouveau Pselaphidae cavernicole d'Istrie (Croatie): Pauperobythus globuliventris n. gen., n. sp. (Coleoptera, Pselaphinae, Bythinini). Bull Soc Entomol Fr., 107(1), 13-18. Novak, T., M. Perc, S. Lipovšek & F. Janžekovič (2012): Duality of terrestrial subterranean fauna. Int. J. Speleol., 41(2), 181-188. Otoničar, B. (2007): Upper cretaceous to paleo-gene forbulge unconformity associated with foreland and basin evolution (Kras, Matarsko podolje and Istria; SW Slovenia and NW Croatia). Acta Carsologica, 36(1), 101-120. Ozimec, R. (2002): Review of genus Troglochtho-nius Beier, 1939 (Arachnida, Pseudoscorpiones, Chtho-niidae), Abstracts of 16th International Symposium of Biospeleology. 815 September 2002, Verona, Italy, pp. 56-57. Ozimec, R., J. Bedek, S. Gottstein, B. Jalžic, R. Slap-nik, V. Štamol, H. Bilandžija, T. Dražina, E. Kletečki, A. Komerički, M. Lukc & M. Pavlek (2009): Crvena knjiga špiljske faune Hrvatske (Red book of Croatian cave dwelling fauna). Ministarstvo kulture & DZZP, Zagreb, pp.1-371. Ozimec, R., S. Polak, J. Bedek & V. Zakšek (2010): Biospeleological component of the project KUP (Karst Underground Protection) in Istra Peninsula. In: Moškrič, A. & P. Trontelj (eds.): Abstract book. 20th International Conference on Subterranean Biology. 29 Aug - 3 Sept 2010, Postojna, Slovenia, pp. 89-90. Ozimec, R., S. Polak, J. Bedek & V. Zakšek (2011): Importance of biospeleological research for protection of cave fauna and their habitats - example based o the project Karst underground protection on the Istrian peninsula. In: Prelovšek, M. & N. Zupan Hajna (eds.): Pressures and Protection of the Underground Karst -Cases from Slovenia and Croatia (Pritiski in varovanje podzemnega krasa - Primeri iz Slovenije in Hrvaške; Pritisci i zaštita podzemnog krša - Primjeri iz Slovenije i Hrvatske). Karst Research Instutute, Postojna, pp. 160-168. Perko, G. A. (1909): Speleologia. Grotta dell'Orso. Il Tourista, 12/13, 1905/06, 61-82. Polak, S. & T. Pipan (2011): Subterranean habitats and fauna, their threats and conservation. In: Prelovšek, M. & N. Zupan Hajna (eds.): Pressures and Protection of the Underground Karst - Cases from Slovenia and Croatia (Pritiski in varovanje podzemnega krasa - Primeri iz Slovenije in Hrvaške; Pritisci i zaštita podzemnog krša - Primjeri iz Slovenije i Hrvatske). Karst Research Instutute, Postojna, pp. 23-32. Polak, S. (1997): Prispevek k poznavanju podzemeljske favne nekaterih jamskih objektov Matarskega podolja, jugozahodna Slovenija. In: Bedjanič, M. (ur.): Raziskovalni tabor študentov biologije Podgrad '96. ZOTKS, Gibanje znanost mladini, pp. 45-54. Presetnik, P., K. Koselj & M. Zagmajster (ur.) (2009): Atlas netopirjev (Chiroptera) Slovenije. Atlas faunae et florae Slovenia 2. Center za kartografijo favne in flore, Miklavž na Dravskem polju, 151 p. Rada, T. (1980): Čovječja ribica u Pincinovoj jami kod Poreča. Priroda, 48(7/8), 179-181. Schiner, J. R. (1854): Fauna der Adelsberger-, Lue-gger-, and Magdalenen Grotte. In: Schmidt, A. (ed.): Die Grotten und Höhlen von Adelsberg, Lueg, Planina, und Laas. Braumüller, Vienna, Austria, pp. 231-272. Sket, B. (1994): Distribution patterns of some subterranean Crustacea in the territory of the former Yugoslavia. Hydrobiologia, 287, 65-75. Sket, B. (1999): The nature of biodiversity in hypo-gean waters and how it is endangered. Biodiversity Con-serv., 8, 1319-1338. Sket, B. (2008): Can we agree on an ecological classification of subterranean animals? J. Nat. Hist., 42(2122), 1549-1563. Sket, B., K. Paragamian & P. Trontelj (2004): A census of the obligate subterranean fauna of the Balkan Peninsula. In: Griffiths, H. I., B. Kryštufek & J. M. Reed (eds.): Balkan Biodiversity: pattern and process in the European hotspot. Kluwer Academic Publisher, Dordrecht, pp.309-322. Sokal, R. R. & F. J. Rolph (1995): Biometry: The principles and practice of statistics in biological research. 3rd edition. W.H. Freeman, New York. Slavko POLAK et al.: SUBTERRANEAN FAUNA OF TWELVE ISTRIAN CAVES, 7-24 Stammer, H. J. (1932): Die Fauna des Timavo. Ein Beitrag zur Kenntnis der Höhlengewässer des Süß- und Brackwassers im Karst. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 63, pp. 521-656. Strasser, K. (1959): Verhoeffodesmus n. g., ein Hö-hlendiplopode aus Istrien (Polydesmoidea). Acta Carso-logica, 2, 99-106. Verhoeff, K. W. (1933): Arthropoden aus sudostalpinen Höhlen, gesammelt von Karl Strasser. Mitteilungen über Höhlen- und karstforschung, 4, 1-21. Wolf, B. (1934-1938): Animalium cavernarum cata-logus. Junk, s'Gravenhage. Zagmajster, M., D. C. Culver & B. Sket (2008): Species richness patterns of obligate subterranean beetles in a global biodiversity hotspot-effect of scale and sampling intensity. Diversity Distrib., 14, 95-105. Zaksek, V., S. Gottstein, D. Franjevic, B. Sket & P. Trontelj (2009): The limits of cryptic diversity in groundwater: phylogeography of the cave shrimp Troglocaris anophthalmus (Crustacea: Decapoda: Atyidae). Mol. Ecol., 18, 931-946. PLATE I: Some representatives of the Istrian subterranean fauna photographed during the study. TABLA I: Nekateri predstavniki istrskega podzemeljskega živalstva, fotografirani med raziskavo Photograph credits / Avtorji fotografij: RO - Roman Ozimec, SP - Slavko Polak, JB - Jana Bedek. Fig. 2 / Sl. 2: Leptodirus hochenwartii reticulatus (SP) Fig. 3 / Sl. 3: Prospelaeobates vrezeci (SP) Fig. 4 / Sl. 4: Pauperobythus globuliventris (RO) Fig. 5 / Sl. 5: Typhlotrechus bilimeki istrus (SP) Fig. 6 / Sl. 6: Onychiurus giganteus (RO) Fig. 7 / Sl. 7: Plusiocampa cf. nivea (SP) Fig. 8 / Sl. 8: Stalita taenaria (RO) Fig. 9 / Sl. 9: Troglochthonius doratodactylus (SP) PLATE II: Some representatives of the Istrian subterranean fauna photographed during the study. Photographs credits as in Plate I. TABLA II: Nekateri predstavniki istrskega podzemeljskega živalstva, fotografirani med raziskavo. Avtorji fotografij kot na Tabli I. Fig. 10 / Sl. 10: Typhloglomeris cf. fiumarana (RO) Fig. 11 / Sl. 11: Verhoeffodesmus gracilipes (RO) Fig. 12 / Sl. 12: Sphaeromides virei virei (SP) Fig. 13 / Sl. 13: Monolistra bericum hadzii (JB) Fig. 14 / Sl. 14: Thaumatoniscellus speluncae (JB) Fig. 15 / Sl. 15: Zospeum spelaeum schmidti (SP) Fig. 16 / Sl. 16: Triphosa dubitata (SP) Fig. 17 / Sl. 17: Myotis daubentonii (SP)