Acta carsologica

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Acta carsologica

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Acta carsologica
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Acta Carsologica
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Krasoslovni zbornik
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Vol. 40, no. 3 (2011)

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KARST LANDFORMS IN AN INTERIOR LA Y ERED DEPOSIT WITHIN THE COPRATES CHASMA, MARS KRAKE OBLIKE NA PLASTNATIH USEDLINAH V COPRATES CHASMI, MARS Davide BAIONI 1 Nadja ZUPAN HAJNA 2 & Forese Carlo WEZEL 3 Izvleek UDK 551.435.8:523.43 Davide Baioni, Nadja Zupan Hajna & Forese Carlo We z el: Krake oblike na plastnatih usedlinah v Coprates Chasmi, Mars Coprates Chasma tvori del osrednjega dela sistema kanjonov Valles Marineris. V njenem najzahodnejem delu, na dnu pod severno steno, se nahaja gmota plastnatega materiala, ki kae znailno morfologijo dome. Mineralna sestava dome in njene okolice je bila doloena z analizami CRISM (Compact Recon naissance Imaging Spectrometer for Mars) podatkov (image HRL00003752). Na analizirani enoti je jasen zapis kizerita, to je evaporitnega minerala, ki ga najdemo tudi na Zemlji. Z analizami MRO HiRISE (Mars Reconnaissance Orbiter High Resolution Imaging Science Experiment images) smo preiskovali povrinske oblike na domi in mone procese, ki so bili vpleteni v njihov nastanek in oblikovanje. Analize so pokazale, da opazovane oblike jasno kaejo na prisotnost procesov raztapljanja, ki so tudi oblikovale tam prisotne krake oblike. Re zultati naih opazovanj tudi nakazujejo, da je na povrju dome v preteklosti morala obstajati tekoa voda in to zadosti dolgo, da so lahko nastale korozijske oblike; ter da preiskovane krake oblike kaejo stareje erozijske starosti ali kraji as delovanja vode, kot enake oblike, ki so bile preiskovane na podobni kizeritni domi v Tithonium Chasmi, drugem jarku sistema Vales Marineris. Kljune besede: evaporitne usedline, kras, Coprates Chasma, Valles Marineris, Mars. 1 Dipartimento di Scienze della Terra, Universit degli studi di Parma, Campus Universitario, Via delle Scienze 43100 Parma (PR), Tel. e Fax 0521/905323, e-mail: davide.baioni@unipr.it 2 Karst Research Institute ZRC SAZU, Titov trg 2, 6230 Postojna, Slovenia, e-mail: zupan@zrc-sazu.si 3 Planetary Geology Research Group, University of Urbino Carlo Bo" Italy, e-mail: forese-carlo.wezel@uniurb.it Received/Prejeto: 29.9.2011 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 473, POSTOJNA 2011 Abstract UDC 551.435.8:523.43 Davide Baioni, Nadja Zupan Hajna & Forese Carlo We z el: Karst landforms in an interior layered deposit within the Co prates Chasma, Mars e Coprates Chasma forms part of the backbone of the Valles Marineris canyon system. In the westernmost part of the chasma in an embayment on the northern wall a mound of layered material rises from the chasma oor and displays a characteristic dome-shaped morphology. e mineralogical characteristics of the dome and its surroundings have been de termined by analysis of the CRISM (Compact Reconnaissance Imaging Spectrometer for Mars) data (image HRL00003752). e unit shows the clear signatures of kieserite, an evaporite mineral also found on Earth. rough analysis of the M.RO. HiRISE (Mars Reconnaissance Orbiter High Resolution Imag ing Science Experiment images) we have investigated the dome landforms and the possible processes involved in their forma tion and shaping in great detail. e analysis shows that the landforms observed clearly indicate the presence of solutional processes that made those karst landforms. e results of our observation also suggest that liquid water must have existed on the dome in the past for long enough for the solution features to be formed, and that the karst landforms investigated exhib it an older erosional age or shorter than the same landforms studied in a similar kieserite dome located within Tithonium Chasma, another graben of the Valles Marineris system. Keywords: evaporite deposits, karst, Coprates Chasma, Valles Marineris, Mars.

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ACTA CARSOLOGICA 40/3 2011 474 Layered deposits of Mars were rst observed on Mari ner 9 images and have been reported as thick deposits with internal layering, and a relative high albedo (Luc chitta et al. 1994). Signicant deposits occur in almost all deep canyons of the Valles Marineris system where they are called interior layered deposits (ILD). e ILDs have been widely studied in the last few decades, but still they remain among the most puzzling and controversial of the canyons' features (Carr 2006). Recently, observations from the OMEGA spectrometer, on board Mars Express have shown the presence of hydrate minerals, in particu lar sulphates, in association with ILDs. e Coprates Chasma (CC) is part of the Valle Mar ineris structure (Fig. 1A), a ri system that belongs to the arsis radial pattern of fractures (Carr 1981). Locat ed next to the Martian equator, the Coprates Chasma ex tends approximately eastwest for roughly 966 km (600 miles). In the Coprates Chasma, the ILD of Late Hes perian through Middle Amazonian age (Lucchitta et al. 1994), indicated by OMEGA data to consist of magne sium sulphate (Bibring et al. 2006), are not found other than at the western end of the trough near its conuence with the Melas Chasma. A dome-shaped ILD rises from the chasma oor in the central part of an embayment lo cated in the westernmost part of the Coprates Chasma and is the topic of this paper. e objective of this work was, following a previous study on an ILD located in the eastern part of Tithonium Chasma (called East Tithonium Dome, ETD) that high lighted the presence of several karst landforms (Baioni et al. 2009; Baioni & W ezel 2010), to investigate the pres ence of karst landforms in the ILD located in north Co prates Chasma, with the aim of obtaining more informa tion about the morphological history and processes that aected the ILDs. To this end, a morphological analysis of the available Mars Reconnaissance Orbiter (MRO) High Resolution Imaging Science Experiment (HiRISE) images was performed with the aim of describing surface features and landforms, and, where possible, morphoge netic processes. e morphological features of the struc ture were investigated through integrated analysis of HiRISE, High Resolution Stereo Camera (HRSC), Mars Orbiter Camera (MOC), and ermal Emission Imaging System (THEMIS) data. e morphometric characteris tics of the structure were measured using a topograph ic map (50-m contour interval) built from HRSC and Mars Orbiter Laser Altimeter (MOLA) data. INTRODUCTION STUD Y AREA SETTING GEOLOG Y AND MINERALOG Y e Coprates Chasma (Fig. 1B) extending east from the Melas Chasma, consists of several parallel sections sepa rated by at-topped or sharp-crested ridges depending on the width of the divides (Carr 2006). It forms part of the backbone of the Valles Marineris canyon system. e Coprates Chasma is the longest and deepest of the Valles Marineris system troughs. e trough is about 1,000 km long, and ranges in depth from 6 km in the eastern sec tion to over 10.5 km in the west, with an average depth of 8 km. e geology is dominated by Noachian wall rock and various trough oor materials of Middle Hespe rian age (Schultz 1998; Jernsletten 2004). e main canyon has a oor that displays only a small regional slope, which is relatively free of landslide debris and layered sediments compared with other Valles Mari neris canyons, although it is partly blocked at its nar rowest point. At the canyon's western end are some of the highest walls within the canyon system. ey are roughly 10,000 m high, extending from under -4,500 m to over 5,000 m in elevation (Carr 2006).e at oor is at an elevation 9 km below the surrounding plains. Canyon walls are characterized by spur and gully mor phology and distinct competent layers. Recent studies (Jernsletten 2004) observing the apparent longevity of the spur and gully morphology and the topographic patterns that indicate the absence of ground ice conclude that these morphologies were formed by the action of liquid water and not by dry mastwasting above an ice-rich crust. Others studies indicate the possible presence of ice and/or snow at tropical to mid latitude regions of Mars in the Amazzonian age due to relative planet low obliquity periods (Head et al. 2005; Pacici et al. 2009). Distinct competent layers are observed in the slopes of the eastern Coprates Chasma where the stratigraphy consists of alternating thin strong layers and thicker se quences of relatively weak layers. e strong, competent, layers maintain steeper slopes and play a major role in controlling the overall shape and geomorphology of the chasma slopes. is dierent combination of layers may have a large impact on chasma and slope morphology, D AVIDE BAIONI, N ADJA ZUPAN HAJNA & FORESE CARLO WEZEL

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ACTA CARSOLOGICA 40/3 2011 475 as well as the locations and sizes of landslides (Beyer & McEwen 2005). Along some of the walls, particularly the north walls, the downward branching spurs are truncated near their bases by fault scarp to form triangular facets. Alignment of the facets results in continuous linear es carpments hundreds of kilometers long providing com pelling evidence that they are fault scarps (Carr 2006). e faults dening the northern boundary of the trough are more continuous and recently active compared with those dening the southern boundary (Schultz 1998; Peulvast et al. 2001). In the westernmost part of the Coprates Chasma, next to the borderline with Melas Chasma, an embay ment on the northern wall can be observed. e embay ment displays the southern boundary formed by linear embankment trending approximately 110. Along this boundary, the base of the embayment is located 1 km above the oor of the chasma (Fueten et al. 2010). In the central part of the embayment, a mound of layered material, referred to as the North Coprates Dome (NCD) rises from the chasma oor and displays a char acteristic dome-shaped morphology (Fig. 1C). e term dome is used from the morphological point of view and is related to the shape of the mound. It does not imply any interpretation or meaning about the processes that built the studied relief. e NCD is approximately 12 km long and 8 km wide and its base is around 2,700 m above the Coprates oor. Following the terminology of Malin and Edgett (2000), that divide into three categories the layered de posits within the Valles Marineris, the NCD can be con sidered as composed of primarily layered units, which means that it is constituted mainly by light-to intermedi ate-toned "layered" units. e mineralogical characteristics of the NCD and its surroundings have been previously determined by KARST LANDFORMS IN AN INTERIOR LA Y ERED DEPOSIT W ITHIN THE COPRATES CHASMA, MARS Fig. 1: (A) V alles M arineris, M ars. L ocation of Coprates Chas ma (CC), M ars (black box). I m age taken from NASA Planetary Photo Journal Collection web site (www.nasaimages.org ID 110192). (B) I mage of Coprates Chasma, showing the interior layered deposit (the North Co prates D ome; NCD) at the west ern end of the chasma, marked by the black box. I mage NASA/ USGS/ESA/DLR/FU Berlin (G. Neukum) taken from G oogle M ars. (C) I mage of the North Co prates D ome viewed from southeast. I mage NASA/USGS/ESA/ DLR/FU Berlin (G.Neukum) taken from G oogle M ars. (D) CRIS M image HRL00003752 of the NCD acquired at Ls = 153.

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ACTA CARSOLOGICA 40/3 2011 476 NCD MORPHOLOGICAL CHARACTERISTICS e dome, which shows an estimated thickness of about 2 km, rises from the chasma oor to an absolute altitude of about 200 m that is also the maximum elevation at which the deposits can be identied within the embay ment, approximately 3.4 km below the elevation of the plateau (Fueten et al. 2010). e dome shows an elongate elliptical plane shape. Its longest axis displays a NNE SSW -elongated trend with a length of about 17 km, con sidering as part of the deposits the layers that from its southern edge can be seen extending southwards for sev eral kilometers. Its width varies from 10 to 15 km. e crestal region of the NCD is the central part of the struc ture and consists of a nearly horizontal summit plateau that is about 5 km long and has a maximum width of about 1 km. It displays a very gentle slope toward south and is surrounded by steeply sloping anks. e NCD has a ank topography that varies in its dierent sectors, generally displaying higher gradient values in the western ank. In fact, the ank slopes are similar in the central and northern part of the dome, while they are very dierent in the southern part. In the central part of the dome, the anks have comparatively steep slopes, ranging from 30% on the western side to 26% on the eastern side. e northern part of the dome has anks with slope angles that range from 29% on the western sides to about 23% on the eastern sides. e asymmetric anks of the southern part of the dome have the maximum dierence in slope angles, ranging from 24% on the eastern side to 49% on the western side. Here the western ank displays the highest slope values of the entire NCD. e base of the NCD and its transition to the surrounding chasma oor and wallrock cannot be recognized. As suggested by a previous study (Fueten et al. 2010) the base of the deposit and the underlying basement can only be estimated assuming that it dips uniformly from north to south. In fact, the borderline between the base of the NCD and the chasma oor is obscured on almost all sides of the dome. On the south ern, eastern and western sides the margins of the NCD are covered by fan deposits, rock debris or thin material, while on the northern and northeastern sides they seem visible and linear. In particular, on the northeastern part the NCD margin appears as a continuous straight and vertical wall that rises from the chasma oor. analysis of the CRISM (Murchie et al. 2007) image HRL00003752 (Fig. 1D). In particular, a recent study (Fueten et al. 2010) shows that clear signatures of kieser ite (MgSO 4 .H 2 O) can be observed both in two large re gions on the oor (in the SE and SW sides of the NCD), and in a discrete number of locations on the slope of the NCD (to the SE and from the SW to the NW ), while sig natures of polyhydrates sulphates can be detected on the southern slope of the NCD and on the chasma oor to the north of the northern slopes. Studies on a similar dome-shaped layered deposit (ETD) located within the Tithonium Chasma showed that the kieserite displayed on the surface could be the result of alteration processes and that the dome probably consists of the same salts that exist on Earth changing into kieserite, such as carnalite, kainite and halite (Son nefeld 1984; Baioni & W ezel 2008, 2010 and reference there in). KARST LANDFORMS Typical karst corrosional surfaces formed by sheet wash and channeled water ow can be observed along the NCD anks. Similar corrosional features have been de tected also in other ILDs within the Tithonium Chasma in the northern part of the Valles Marineris canyon sys tem (Baioni et al. 2009; Baioni & W ezel 2010). Dierent solution features were detected on the slopes of the NCD: such as karren and various depres sions. K ARREN FEATURES Karren is complex group of smallto medium-sized karst landforms of great shape variety; some of karren features can dene the genetic factors of dissolution processes in volved in their formation (Ford & W illiams 2007; Gins 2009). In the southwestern region of the dome slope, below the edge of leveled terrace (cuesta like shape), complex giant solutional channels (like wandkarren) D AVIDE BAIONI, N ADJA ZUPAN HAJNA & FORESE CARLO WEZEL

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ACTA CARSOLOGICA 40/3 2011 477 are present down the slope; these channels are evidence of water ow down the slope in the past. at they are not just erosional gullies but also corrosional features is evidenced by sharpened edges of the ridges between runnels and also the well expressed sharp edge of ter race rim at their beginning (Fig. 2A). Sharpened edges of karren forms are characteristic for dissolution by snow melting (Gins 2009). Leveled surfaces controlled by bedding planes are present on the western slope of the dome steps (Fig. 2B). e surfaces of the steps and slopes between them are mostly smoothed, as if they were dissolved by melting snow/ice waters like in the Alps (Gins et al. 2009); bowl shaped shallow depressions are present on them in some places. Also some juvenile karren forms (Figs. 2C & 2D) can be detected between single steps, some times starting just below the step rim or below open ings along bedding planes; perhaps as the evidence of short-lived outow (Fig. 2D). From extant features we can assume that there was not enough available water to shape more evident relief or that they were already eroded by some other processes. D EPRESSIONS Many closed depressions of various sizes and shapes exist on the dome anks (Fig. 3). e process of their formation is not easy to be determined. W e have no evi dence if they are solution or collapse dolines or some solution pans, but their shape is not like of wind scal lops at all. e depressions on the eastern and western anks are either bowl-shaped or rounded (Figs. 3A to 3D), they have diameters up to 50 m, display asymmetrical walls (Figs. 3A & 3B) and concave-up (Figs. 3A & 3C) or have at oor geometry (Figs. 3B, 3D & 3E). In some cases, like on the western ank, the oors are subdi vided into smaller and shallower depressions (Fig. 3E) that have diameters ranging from one to a few meters, highlighting a second generation of solution depres sions. ese multiple forms indicate minimum two episodes of water appearance (ice melting) on the dome surface and at least two-stage dissolution of evaporite rock during depression formation. On the upper part of the southern ank, where the incline of the slope is higher, depressions have an elongated shape with a wide Fig. 2: (A) complex giant solutional channels (like wandkarren) on the down slope part of the south-western ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north toward the le of image). (B) steps like levelled surfaces on the upper part of the western ank of the NCD (MRO H iRISE image PSP_001456_1695, north toward the upper le of image). (C) Juvenile karren (white dashed arrows) on the down slope part of the western ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north toward the upper le of image). (D) Juvenile karren (white arrow) on the middle part of the western ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north toward the upper le of image). I mages from H iRISE website (http://hirise.lpl.arizona.edu). KARST LANDFORMS IN AN INTERIOR LA Y ERED DEPOSIT W ITHIN THE COPRATES CHASMA, MARS

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ACTA CARSOLOGICA 40/3 2011 478 Fig. 3: D olines. (A) Roundedshaped doline on the western ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north at the top of image). (B) Roundedshaped doline with asymmetrical walls and at oor geometry on the south-eastern ank of the North Coprates D ome (MRO H iR I SE image PSP_001456_1695, north at the top of image). (C) Bowl-shaped doline with asym metrical walls and concave-up oor geometry on the south-east ern ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north at the top of image). (D) D oline with an elongated shape on the upper part of the southern ank of the North Coprates D ome (MRO H iRISE image PSP_001456_1695, north at the top of image). (E) D oline on the western ank of the North Coprates D ome, highlighting a second generation of solution depressions (MRO H iRISE im age PSP_001456_1695, north at the top of image). I mages from H iRISE website (http://hirise.lpl. arizona.edu). top and a narrow bottom (Fig. 3D). ese landforms strongly resemble similar features found on the Earth displaying deep morphological similarities with dolines that develop in all kinds of evaporite and mountain karst terrains, but here were most likely formed like huge solution pans or bevels by sheet wash water ow during ice melt. Polygonal-like karst ( W illiams 1972; W hite 1988; Ford & W illiams 2007) can be observed on the parts of the NCD that have lower slope angles, such as the down slope region of the southeastern part of the dome (Fig. 4A). Here depressions entirely pock some parts of the surface and occupy most of its area (Fig. 4B). e depressions appear to be spaced farther apart, display well dened shapes with sharp divides (Figs. 4B & 4C) and have diameters that range from 30 m to more than 100 m. is area viewed from above, has an irregular egg-box-like topography and the divides between ad jacent depressions form a cellular mesh pattern (Fig. 4B) just as typically happens in evaporite terrains on Earth where the inherently high solubility of evaporite rocks make densely-packed depressions ( W arren 2006; Ford & W illiams 2007). Because of their rounded shape these can be interpreted as dolines of polygenetic origin ( W illiams 1972; W hite 1988; Ford & W illiams 2007). But we have no evidence that the water really penetrates into evaporite rock, so they can also be formed like huge solution pans by standing or very slowly owing water down the inclined slope. Against the hypothesis that they are formed as wind scallops is the fact that many of them are rounded and not elongated, especial ly those close to the slopes of the dome above the pla teau. Depressions at the edges of the dolines eld were most likely inuenced by aeolian process, because they D AVIDE BAIONI, N ADJA ZUPAN HAJNA & FORESE CARLO WEZEL

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ACTA CARSOLOGICA 40/3 2011 479 DISCUSSION AND CONCLUSIONS e presence of the described karst features on the NCD suggests their formation by laminar/sheet wash water ow which was partly canalized in runnels or wall kar ren. e karst features on the NCD, which were most likely formed by water during snow or ice melting, can be well correlated with the forms typical for high moun tain karst regions on Earth, such as, decantation runnels, solutional bevels, tritkarren, and solution ripples (Gins et al. 2009). e depressions observed do not display any evidence that they functioned as typical solution dolines where more rock mass is removed from their centers than from around their sides by the dissolution of inltration water. In fact, many of them look more like solution pans or bevels (ausgleichschen) or closed corrosional terraces formed by melting water owing along the surface without any or little penetration into the bedrock (kieserite). e necessary liquid water for the solutional pro cesses observed has been provided by melting ice or snow, which on steeper slopes can be canalized. e melting of ice or snow, which can be formed during pe riods of ice-snow-rich deposition from the atmosphere that may occur as the result of changes in the obliquity of Mars (Mustard et al. 2001; Laskar et al. 2004, Paci ci et al. 2009), probably occurred gradually rather than rapidly, and had to persist long enough to shape the karst landforms observed. e landforms observed on the NCD allow us to establish that the source of the water might be the melting of ice or snow rather than other dierent processes such us, rain or atmospheric humid ity. In particular, on the down slope part in the western ank of the NCD, the presence of few lobate morpholo gies and some gully systems seem to indicate melting of Fig. 4: Polygonal like karst landscape. (A) L ocation of the polygonal like karst area on the down slope region of the south-eastern part of the North Coprates D ome (MRO H iRISE image PSP_001865_1695, north at the top of image). (B) Perspe ctive view of polygonal like karst (MRO H iRISE image PSP_001865_1695, north at the top of image). (C) Particu lar of the depressions displaying well dened shape with sharp divides (MRO H iRISE image PSP_001865_1695, north at the upper le of image). I mages from H iRISE website (http://hirise.lpl. arizona.edu). are elongated in the wind blow direction and can be compared to the direction of the dunes in the vicinity. Elongated shape of depressions can be also interpreted as hydroaeolian landforms (Maire et al. 2009) formed by horizontal solution accelerated by wind deection. at means that during water existence on the surface of the NCD slopes strong and permanent winds facili tate laminar ow in direction of wind blow and solu tion forms are elongated along it. KARST LANDFORMS IN AN INTERIOR LA Y ERED DEPOSIT W ITHIN THE COPRATES CHASMA, MARS

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ACTA CARSOLOGICA 40/3 2011 480 REFERENCES Baioni, D. & F.C. W ezel, 2008: Similarities of a Martian dome with terrestrial salt domes.Italian Journal of Geosciences, 127, 3, 453. Baioni, D., Zupan Hajna, N. & F.C. W ezel, 2009: Karst landforms in a Martian evaporitic dome. Acta Carsologica, 38, 1, 9. Baioni, D., & F.C. W ezel, 2010: Morphology and origin of an evaporitic dome in the eastern Tithonium Chasma, Mars.Planetary and Space Science, 58, 847. Beyer, R.A. & A.S. McEven, 2005: Layering stratigraphy of eastern Coprates and northern Capri Chasmata, Mars. Icarus, 179, 1, 1. ice or snow as the main processes involved in their for mation and shaping. e analysis carried out also highlights that along the NCD slopes there is no evidence of deep depressions or collapse dolines and almost no outow evidences (just a few juvenile runnels below bedding planes), which are the only possible surface evidences of ground water existence in some stage of NCD development. In our observation of the NCD surface we did not notice any deep depressions with steep slopes, cave entrances, va dose shas or outow caves, as we found in a previous study in a similar evaporite dome located in the Titho nium Chasma (Baioni et al. 2009) and which represent ed the strong evidences of well developed and karstied underground and quite long-lasting or multiphase water existence. Karst surface and underground features are much more abundant on the ETD than on the NCD. From these facts can be anticipate that: a) less water (ice) available on the NCD; b) melting period was shorter or very fast; or c) karst features were already eroded by ae olian processes (the NCD is more exposed to the wind or karstication was older than on the ETD and erosion went on for longer time). e characteristics of evaporite karst and its rapid formation on the Earth (Klimchouk 2004), lead us to think that liquid water existed on the NCD and persisted long enough for solution features to form but that there probably was only one episode during which water was available, but still perhaps in few events according to presence of some secondary depressions in older ones. e karstication was probably short-lived. Aer this period of karstication, there was no more water avail able and hence no additional karstication of the NCD. e existence of liquid water that owed gradually for a period along the NCD surface also seems to be con rmed by the landforms found at the foot of the southwestern ank. Here, systems of gullies running from the alcove sources to the fan deposits located on the base of the NCD can be clearly observed, displaying strong evi dence for the presence of running liquid water. e karst landforms observed on the NCD oen ap pear to be partly reworked or modied by wind erosion. It has been observed that wind erosion does not aect the landform found on the dome surface in the same way. In fact, the landforms located in the down slope part of the south-western region appear less aected by wind ero sion than the landforms located in all other parts. is might be explained by the dierent exposition to wind action. In fact, the south-western part of the NCD ap pears to be more protected by the wind action than by the topographic context. e characteristics of the karst landforms and the presence of landforms with wind-related modications might suggest that they are not of a very young erosional age. In particular, comparing (these landform) with the others found in a similar kieserite dome in the Titho nium Chasma investigated in a previous study (Baioni et al. 2009), they exhibit an older erosional age. It may be due either to the dierent exposition to the wind, or to the dierent time that they have been exposed to the wind, or, more probably, to both factors. e analysis carried out in this study suggests that: i) e karst landforms might be consistent with re sponses to climatic change and the presence of enough liquid water to form the observed landforms. ii) ere was only a single geologically short "coolwet episode" with available water during the NCD his tory, probably occurring in the Amazonian period, aer which there was no water. iii) e karst landforms observed in the NCD seem to be more aected and modied by wind erosion than those found and reported in a similar evaporite dome lo cated in the Tithinium Chasma (Baioni et al. 2009), ex hibiting an older erosional age. It suggests that the ILDs located in Tithonium Chasma and in the NCD might have dierent ages and that the NCD might be older than the ETD. D AVIDE BAIONI, N ADJA ZUPAN HAJNA & FORESE CARLO WEZEL

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ACTA CARSOLOGICA 40/3 2011 481 Bibring, J.P., Langevin, Y ., Mustard, J.F., Poulet, F., Arvid son, R., Gendrin, A., Gondet, B., Mangold, N., Pi net, P., Forget, F., & the OMEGA team, 2006: Global Mineralogical and Aqueous Mars History Derived from OMEGA/Mars Express Data.Science, 312, 400. Carr, M.H., 1981: e Surface of M ars. Y ale Univ. Press, pp. 232, New Haven. Carr, M.H., 2006: e surface of M ars. Cambridge Uni versity Press, pp. 307, New Y ork. Ford, D.C. & P.W W illiams, 2007: K arst H ydrogeology and G eomorphology.W iley & Sons Ltd, pp. 562, W est Sussex, England. Fueten, F., Racher, H., Stesky, R., MacKinnon, P., Hauber, E., McGuire, P.C., Zegers, T. & K. Gwinner, 2010: Structural analysis of interior layered deposits in Northern Coprates Chasma, Mars.Earth and Plan etary Science Letters, 294, 343. Gins, A., 2009: Karreneld landscapes and karren land formsIn Gins et al. (eds.) K arst rock features kar ren: sculpturing. ZRC Publishing, pp. 13, Postoj naLjubljana. Gins, A., Knez, M., Slabe, T. & W Dreybrodt, 2009: K arst rock features: karren sculpturing. ZRC Pub lishing, pp. 561, Postojna Ljubljana. Head, J.W ., Neukum, G., Jaumann R., Hiesinger H., Hau ber E., Carr, M., Masson P., Foing, B., Homann, H., Kreslavsky, M., W erner, S., Milkovich, S., Van Gasselt, S. & e HRSC Co-Investigator Team, 2005: Tropical to mid-latitude snow and ice accu mulation, ow and glaciatio on Mars.Nature, 434, 7031, 346. Jernsletten, J. A., 2004: A topographic test for the exis tence of ground ice in the walls of Coprates Chas ma, Mars.Journal of Geophysical Research, 109, E12004. Klimchouk, A., 2004: Evaporite karst.In Gunn, J. (ed.) Encyclopedia of caves and karst science. Fitzroy Dearborn, pp. 343, New Y ork. Laskar, J., Correia, A.C.M., Gastineau, M., Joutel, F., Lev rard, B. & P. Robutel, 2004: Long term evolution and chaotic diusion of the insolation quantities of Mars. Icarus 170, 343. Lucchitta, B.K., Isbell, N.K. & A. Howington-Kraus, 1994: Topography of Valles Marineris: implications for erosional and structural history.Journal of Geophysical Research, 99, 3783. Malin, M.C. & K.S. Edgett, 2000: Sedimentary rocks of early Mars.Science, 290, 1927. Maire, R., Jaillet, S. & F., Hobla, 2009: Karren in Pa tagonia, a natural laboratory for hydroaeolian dis solution.In Gins et al. (eds.) K arst rock features karren: sculpturing. ZRC Publishing, pp. 329, PostojnaLjubljana. Murchie, S., Arvidson, R., Bedini, P., Beisser, K., Bibring, J.-P., Bishop, J., Boldt, J., Cavender, P., Choo, T., Clancy, R.T., Darlington, E.H., Des Marais, D., Es piritu, R., Fort, D., Green, R., Guinness, E., Hayes, J., Hash, C., Heernan, K., Hemmler, J., Heyler, G., Humm, D., Hutcheson, J., Izenberg, N., Lee, R., Lees, J., Lohr, D., Malaret, E., Martin, T., McGovern, J.A., McGuire, P., Morris, R., Mustard, J., Pelkey, S., Rhodes, E., Robinson, M., Roush, T., Schaefer, E., Seagrave, G., Seelos, F., Silverglate, P., Slavney, S., Smith, M., Shyong, W .-J., Strohbehn, K., Taylor, H., ompson, P., Tossman, B., W irzburger, M. & M. W ol, 2007: CRISM (Compact Reconnaissance Imaging Spectrometer for Mars) on MRO (Mars Reconnaissance Orbiter).Journal of Geophysical Research, 112. Mustard, J.F., Cooper, C.D. & M.K. Ritin, 2001: Evi dence for recent climate change on Mars from the identication of youthful near surface ground ice.Nature, 412, 411. Pacici, A., Komatsu, G. & M. Pondrelli, 2009: Geologi cal evolution of Ares Vallis on Mars: Formation by multiple events of catastrophic ooding, glacial and periglacial processes.Icarus, 202, 1. Peulvast, J.P., Mege, D., Chiciak, J., Costard, F. & P.L. Masson, 2001: Morphology, evolution and tectonics of Valles Marineris wallslopes Mars.Geomorphol ogy, 37, 329. Schultz, R.A., 1998: Multiple-process origin of Valles Marineris basins and troughs, Mars.Planetary and Space Science, 46, 827. Sonnefeld, P., 1984: Brines and Evaporites. Academic Press, pp. 613, London. W arren, J.K., 2006: Evaporites: Sediments, Resources and H ydrocarbons. Springer-Verlag, New Y ork. W illiams, P.W ., 1972: Morphometric analysis of polygo nal karst in New Guinea.Geological Society of America Bulletin, 83, 761. W hite, W .B., 1988: Geomorphology and Hydrology of Karst Terrain. Oxford University Press, pp.464, New Y ork. KARST LANDFORMS IN AN INTERIOR LA Y ERED DEPOSIT W ITHIN THE COPRATES CHASMA, MARS



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DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA PROPADANJE RNEGA APNENCA Z DRENOVEGA GRIA NA OBJEKTIH KULTURNE DEDIINE LJUBLJANA, SLOVENIA Sabina KRAMAR 1 Ana MLADENOVIb 2 Helmut PRISTACZ 3 & Breda MIRTI 4 Izvleek UDK 552.541:725.94(497.4Lj.) Sabina Kramar, Ana Mladenovi, Helmut Pristacz & Breda Mirti: Propadanje rnega apnenca z Drenovega gria na ob jektih kulturne dediine (Ljubljana, Slovenia) rni apnenec z Drenovega gria (Osrednja Slovenija) velja zaradi svoje tipine barve za enega izmed najlepih sloven skih naravnih kamnov. Apnenec smo raziskali s keminega, mineralokega in petrozikalnega vidika. Nadalje podajamo oblike propadanja preiskovanega apnenca na primeru dveh spomenikov, izmed katerih je eden izpostavljen notranjemu in drugi zunanjemu okolju. Na obeh objektih smo doloili tevilne oblike propadanja, kot so zrnato razpadanje, luenje, drobljenje, eorescenca, skorje in prisotnost mikroorganizmov. Vzorce apnenca smo nato preiskali z optino mikroskopijo, vrstinim elektronskim mikroskopom z elektronsko dispe rzno spektroskopijo, rentgensko prakovno difrakcijsko analizo, meritvami odprte poroznosti in kapilarnega dviga ter meto dami ivosrebrove porozimetije in plinske sorpcije z uporabo argona. Kjub temu, da ima apnenec izredno nizke vrednosti poroznosti in kapilarnosti, so na obeh objektih opazne tevilne oblike propadanja, ki so vezane prav na transport in precipitacijo topnih soli v apnencu. Kljune besede: apnenec, propadanje, topne soli, naravni ka men, spomeniki. 1 Institute for the Protection of Cultural Heritage of Slovenia, Conservation Centre, Restoration Centre, SI-1000 Ljubljana, Slovenia, e-mail: sabina.kramar@rescen.si 2 Slovenian National Building and Civil Engineering Institute, Dimieva 12, SI-1000 Ljubljana, Slovenia, e-mail: ana.mladenovic@zag.si 3 University of Vienna, Institute of Mineralogy and Crystallography, Althanstrasse 14, 1090 Vienna, Austria, e-mail: helmut.pristacz@univie.ac.at 4 University of Ljubljana, Faculty of Natural Sciences and Engineering, Department of Geology, SI-1000 Ljubljana, Slovenia, e-mail: breda.mirtic@guest.arnes.si Received/Prejeto: 2.4.2010 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 483, POSTOJNA 2011 Abstract UDC 552.541:725.94(497.4Lj.) Sabina Kramar, Ana Mladenovi, Helmut Pristacz & Breda Mirti: Deterioration of the black Drenov Gri limestone on historical monuments (Ljubljana, Slovenia) e black limestone from Drenov Gri quarry (Central Slove nia) is considered one of the most beautiful Slovenian natural stones due to its typical color. e limestone was character ized from mineralogical, chemical, and petrophysical points of view. Furthermore, deterioration phenomena of the limestone from two monuments exposed to indoor and outdoor environ ments were studied. In situ investigation of two monuments by means of monument mapping has identied several types of deterioration phenomena, such as granular disintegration, aking, crumbling, eorescences, crusts, and the presence of microorganisms. Samples were characterized using Opti cal Microscopy (OM), Scanning Electron Microscopy (SEM) coupled with Energy Dispersive Spectroscopy (EDS), X-Ray Powder Diraction Analysis (XRD), porosity accessible to wa ter under vacuum, capillary absorption, Mercury porosimetry (MIP), and Ar-sorption. Although very low values of porosity of the fresh stone as well as slow capillary kinetics were de termined, both monuments showed severe deterioration as a consequence of the transport and precipitation of soluble salts within the stone. Keywords: limestone, deterioration, soluble salts, natural stone, historical monuments.

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ACTA CARSOLOGICA 40/3 2011 484 S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI Limestone has been one of the most common stones used for buildings or monuments since prehistoric times. It is widely known, that all stones eventually change due to their interaction with the various envi ronmental conditions. us, similarly as the weather ing processes occur at the Earth's natural surfaces, they also aect man-made monuments or buildings. Lime stone weathering and erosion results in a large variety of karstic forms that have been an object of interest for geologists. On the other hand, the urban atmosphere is creating special environment that aects exposed stone surfaces, usually enhancing the process of decay sev eral times compared to that in natural rural environs (W inkler 1997). Monuments that currently exist in natural environments, or those that were removed from outdoor exposure before air pollution set in, are wellpreserved aer even millennia of outdoor exposure. In contrast, many of those exposed even for a few decades in industrial environments have decayed beyond recog nition (Gauri & Bandyopadhyay 1999). Many Slovenian monuments and modern buildings are built of various limestones, the consequence depen dent on large access of that natural stone in the region, as around 35% of Slovenian territory consists of lime stone (Gams 1974). e properties, behavior and decay of limestones as building materials have been intensively studied over the last decades, by means of dierent ap proaches (Cardell et al. 2003; Benavente et al. 2004; Cardell et al. 2008; Cultrone et al. 2008). Nevertheless, these studies have been mainly concerned with porous limestones, whereas detailed studies of the properties of compact limestones, comparable to Slovenian lime stones, are still rare. Moreover, among the studies, which are focused on the characterization the limestones as building materials or estimating durability (Mladenovi et al. 1999; Skobe & Mirti 2005; Kramar et al. 2010c), there are only few studies related to the weathering phe nomena of limestone on monuments from Slovenian Region and thus the behavior of the limestone in built environment (Kramar et al. 2010a; Kramar et al. 2010b; Kramar et al. 2011). e limestone known as black Drenov Gri limestone is considered one of the most beautiful Slovenian natural stones due to its typical black color interwoven with white veins. It used to be exploited in three major quarries west of Ljubljana which are all at a standstill now. e Triassic well stratied limestone occurs in 10 to 80 cm thick beds which alternate with thin sheets of marls. Some fragments of fossils of bivalvia, gastropoda, algae, foraminifera, os tracods, and chorals are also found (Ramov 2000). Black limestones are normally micritic limestones relatively rich in carbonaceous and bituminous matter, which is supposed to be responsible for their dark color ( W inkler 1997; Marinoni et al. 2007; Marszaek 2007). As the lime stone is dense, resulting in a good polishing eect, it was popularly considered as marble. e Drenov Gri lime stone was widely used particularly in baroque architecture not only in Ljubljana but also in other regions of Slove nia. Many inner and outdoor architectural elements and monuments, especially portals of houses and altars, were made of this limestone (Ramov 2000). e extensive use of black limestones was also reported to be characteristic for the baroque period in other parts of European coun tries (Marinoni et al. 2002; Zehnder 2006; Marinoni et al. 2007; Marszaek 2007). However, when exposed to climat ic inuences, chromatic weathering and salt weathering are recognized as the main deterioration phenomena of these limestones. As many Slovenian monuments built of the inves tigated black limestone are also extensively deteriorated, recognition of limestone properties and understanding of deterioration factors and processes is necessary for their successful maintenance, protection, and proper restora tion/conservation interventions. us, the mineral com position and microstructure of the black Drenov Gri limestone has been studied in order to determine the in trinsic parameters that aect its durability and behavior when exposed to the dierent environmental conditions. e deterioration patterns and related weathering mech anisms of two selected baroque monuments exposed to indoor and outdoor environments have been character ized and discussed. INTRODUCTION EXPERIMENTAL SAMPLING Samples of fresh limestone were collected from the main quarry in Drenov Gri near Ljubljana (Fig. 1a), which had a primary role in past centuries in supplying building material to central parts of Slovenia (Ramov 2000). e samples of weathered black limestone were collected from the two monuments, which have been

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ACTA CARSOLOGICA 40/3 2011 485 DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA exposed to environmental conditions since the baroque period. Sampling was carried out from: the Church of St. James, Ljubljana (Fig. 1b). e altar, made of 18 dierent natural stones with prevailing black limestone, was constructed during the years 1709 (samples JAL). bljana (Fig.1c), constructed during the years 1708 (samples SEP). Information regarding samples of both unweath ered and weathered limestone is provided in Tab. 1. METHODS Detailed registration of the weathering forms was made by means of in situ monument mapping. e applied mapping method was based on a classication scheme proposed by Fitzner & Heinrichs (2002). Polished thin sections of samples of the limestone from the quarry were studied by optical microscopy us ing an Olympus BX-60 equipped with a digital camera (Olympus JVC3-CCD). e samples of both unweathered and weathered limestone were examined by a Scanning Electron Micro scope (JEOL 5500 LV) using back-scattering electrons. Some particular areas of the samples were analyzed for chemical composition using Energy Dispersive Spectros copy (EDS). e excitation voltage was 20 kV, and the pressure was between 10 and 20 Pa. e mineral composi tion of both the unweathered limestone and the weath ering products was determined by X-ray powder dif fraction using a Philips PW3710 X-ray diracto meter equipped with Cu K radiation and a secondary graphite monochromator. e samples were milled in an agate mortar to a particle size of less than 50 m. e data were collected at 40 kV and a current of 30 mA in the range from 2 to 70 2. Aerwards, each sample of limestone was treated in or der to extract its acid-insolu ble residue. About 400 mg of each sample was crushed and dissolved in 20 ml of dilute HCl (1:10) (Marszaek 2007). e residue was then washed with distilled water in order to remove all traces of HCl. e acid-insoluble fraction was then analyzed using Xray powder diraction. All the samples of the unweathered limestone were Fig. 1: (a) One of the three historical quarries of the black limestone in D renov G ri (Central Slo venia). (b) e detail of the side altar in the Church of St. James, L jubljana (Slovenia). (c) Portal with two H ercules at Semenie, L jubljana (Slovenia). T ab. 1: Summary of the investigated samples, related weathering forms and mineralogy as deter mined by X-ray powder difraction and SEM-EDS. Unweathered samples Limestone Location Investigated Samples Primary mineralogy Drenov Gri quarry DG1, DG2, DG3, DG4, DG5 calcite, dolomite, quartz, clinochlore, pyrite, muscovite/illite Samples from monuments Weathering Type Location Investigated Samples Weathering products aking, sublorescence altar JAL23, JAL24, JAL111 gypsum eorescence altar JAL12, JAL13, JAL15, JAL77 hexahydrite, pentahydrite, starkeyite, gypsum white crust altar JAL88, JAL161, JAL 167 gypsum crumbling altar JAL102, JAL153 gypsum splitting altar JAL122, JAL 159 gypsum bleaching altar JAL125 / black crust portal SEP8, SEP11, SEP12 gypsum detachment of crust portal SEP2, SEP4, SEP5 gypsum aking portal SEP7 gypsum bleaching, crumbling portal SEP1, SEP6 / microorganisms portal SEP6, SEP9, SEP10 /

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ACTA CARSOLOGICA 40/3 2011 486 analyzed for their major chemical composition in an ac credited commercial Canadian laboratory (Acme Ana lytical Laboratories, Vancouver, BC, Canada), using dif ferent analytical methods. According to the results of the reports, SiO 2 Al 2 O 3 Fe 2 O 3 MgO, CaO, Na 2 O, K 2 O, TiO 2 and P 2 O 5 were measured aer fusion with a mixture of lithium metaborate/tetraborate and dissolution in nitric acid by inductively coupled plasma emission spectros copy (ICP-ES). e total carbon content was obtained by combustion in an oxygen current (LECO method) and the CO 2 and volatile contents were measured by preci sion scale weighing aer calcination at 1100 C (LOI). e accuracy and precision of the sample analyses were assessed using the reference material CCRMR SO-18 CSC. e total porosity (N t ) of the samples of unweath ered limestone (50 50 50 mm) was measured by water uptake under a vacuum, according to the RILEM recom mendations: RILEM I.1 Norm (RILEM 1980). e water absorption coecient (A), expressed in g m s /2 was measured according to the RILEM II.6 Norm (RILEM 1980). e pore systems of the samples of unweathered limestone were further characterized by means of mercu ry intrusion porosimetry and gas sorption. Small blocks, approximately 2 cm 3 in size, were dried in an oven for 24 hours at 60 C and analyzed on a Micromeritics Auto pore IV model 9500 porosimeter. Adsorption and des orption isotherms of argon were obtained at C on a Micromeritics ASAP 2020 Analyzer. Ar adsorption measurements of samples with a surface area of less than 5 m 2 g are more accurate than using N 2 which usually yields excessively high values (Sing et al. 1985). Prior to measurement, samples were heated at 250 C for 8 h, and outgassed to 10 Torr. Gas adsorption analysis in the rel ative pressure range of 0.05 to 0.3 was used to determine the total specic area, the BET surface area of the sam ples (Brunauer et al. 1938, Gregg & Sing 1982, Adam son & Gast 1997). e pore size distribution curves, the pore volume, and the mean pore size of the rock samples were determined from argon desorption data by the BJH method (Barret et al. 1951). In order to determine the microorganisms, the samples were carefully scratched from the limestone surface and put in sterile asks containing liquid media BG11. Samples were incubated under uorescent light at 25 C and relative humidity of 88% for three months in a climatic chamber (Binder, KB WF 240). Aer the incu bation, a drop of the culture medium was put on a thin section and observed under an optical microscope using a Carl Zeiss model Jenavert. RESULTS AND DISCUSSION L IMESTONE PROPERTIES Petrographical characteriz ation e limestone is determined as micritic and biomi critic according to Folk (1962) and mudstone and pack stone according to Dunhams classication of carbonate rocks (1962). Fragments of fossils are also present (os tracods, bivalvia, foraminifera), and recrystalized bival via bioclasts are especially abundant. Observation with a polarizing light microscope evidences veins, macro scopically exhibited as white color, as well as the pres ence of stylolithes and laminas occurring due to patch ing of limestone during the diagenesis (Fig. 2a). Samples are interwoven by veins of dierent generations, which could be up to 150 m thick. e veins are composed mainly of sparitic calcite; some of them are partially sub stituted with dolomite, as seen in Fig. 2b. Some veins are lled with organic mater. Non-carbonate components include pyrite, clay minerals, quartz, and organic mat ter. Pyrite is represented as authigenic mineral or more frequently as framboidal pyrite. It is concentrated in clay veins and around fragments of fossils, where it could be concentrated in areas 100 m in length and 50 m in width. Some samples contain spots of authigenic quartz ranging in size from 5 to 200 m. Clay minerals could occur either as stylolithes together with organic matter or as veins. SEM-EDS analysis of clay minerals evidences the presence of K, Al, Si, Na, and Mg and is assumed to be sericite. e mineral composition was determined with Xray diraction and besides the main component, calcite, the presence of dolomite, quartz, pyrite, and illite/mus covite, indicating sericite, was also conrmed. Addition ally, the insoluble fraction revealed quartz, pyrite, and illite/muscovite as well as the presence of graphite and clinochlore. e chemical composition of limestone samples by ICP-ES is given in Tab. 2. e results are in accordance with X-ray powder analysis. e values of SiO 2 Al 2 O 3 Fe 2 O 3 Na 2 O, and K 2 O could be attributed to quartz (Si) and clay minerals (Si, Al, K, Fe, Mg). us, the value of SiO 2 is relatively high in samples where the presence of illite/muscovite and clinochlore were determined. Addi S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI

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ACTA CARSOLOGICA 40/3 2011 487 tionally, the presence of Al 2 O 3 is also enhanced in these two samples. A high fraction of Fe 2 O 3 and total sulfur (TOT/S) indicates the presence of pyrite. Mg is attribut ed mainly to dolomite and the concentration is enhanced in samples where dolomite was also evidenced by X-ray powder diraction. Petrophysical characteriz ation As water plays a fundamental role in stone dete rioration, the properties of the stone structure and wa ter transfer were measured. Porosity and pore size dis tribution inuence rock susceptibility to weathering, as nearly all weathering processes rely on the presence and transport of moisture to transport salts or other con taminants into stone and to enable chemical reactions to occur (Nicholson 2001; W arke et al. 2006). e results revealed that porosity of the limestone is very low, as the value of total porosity accessible to water under vacuum is 0.77 0.46%. On the other hand, mercury porosime try, inuenced by free porosity and therefore by the pore network connectivity, yielded values of 3.34 1.40%. e capillary curves of limestone illustrate a weak water absorption anity. e coecient of capillarity is rather low as well, being .38 0.31 gm s ,5 e obtained values are of the same order of magnitude as data pub lished for comparable limestones, where low values were also reported (Sahlin et al. 2000; Marinoni 2002; Kramar et al. 2010b; Miller et al. 2006). Samples that exhibited the highest values of porosity also showed high water ab sorption capillarity coecients. To investigate the mi croporosity (<0.1 m), the BET method is used to de tect nanoscale pores. W hen considering the damage due to salt crystallization, the critical pore radius where the crystallization pressure is eective ranges below 0.05 m (Steiger et al. 2005). e gas-physisorption method is thus suitable for investigation of this range of pore radii. Scherer et al. (1999) established that the maximum pres sure that salt crystallization can achieve is highly depen dent on the size of the pores, predicting that most of the damage occurs when salt-rich uids migrate from pores of larger size to pores of smaller size in the range be tween 4 nm and 50 nm. All samples share a similar pore size distribution with relative maxima located around 2 nm. Values of specic surfaces area vary from 0.4323 to 3.3318 m 2 /g and are higher in samples with higher poros ity. e samples of micritic limestone show a type II iso therm (Sing et al. 1985) indicating the non-microporous or macroporous nature of its pore system network; this is further supported by their very low surface area. In con trast, samples with abundant bioclasts exhibit hysteresis loops (i.e. contrasting shapes of the initial adsorption and nal desorption curves) and much higher surface areas. ese hysteresis loops can be further classied as H3 or H4 and are commonly interpreted as being due to aggre gates of plate-like particles giving rise to slit-shaped pores (e.g. Sing et al. 1985; Lowell et al. 2004). e presence of these kinds of particles in the biomicritic limestone re sults in a signicant increase in the micropore volume relative to the non-microporous micritic limestone. D ETERIORATION PHENOMENA ON THE MONUMENTS Weathering forms As a part of broader conservation/restoration proj ects, in situ investigation of the side altar of St. James Church and the portal at Semenie by means of monu Fig. 2: (a) M icritic limestone with stylolithe in sample DG1. T ransmission light, parallel polars. (b) D olomitized areas in calcitic veins of the limestone in sample DG1. SEM-BSE. T ab. 2: Bulk chemical composition of the limestone samples (percentages by mass). ELEMENT SiO 2 Al 2 O 3 Fe 2 O 3 MgO CaO Na 2 O K 2 O LOI TOT/C TOT/S SAMPLE % % % % % % % % % % DG1 1.83 0.72 0.36 0.82 54.19 0.04 0.13 41.17 12.20 0.08 DG2 1.17 0.75 0.23 0.72 53.96 0.03 0.16 42.80 12.90 0.07 DG3 0.61 0.19 0.25 0.94 56.36 0.02 0.04 41.50 13.91 0.11 DG4 1.52 0.33 0.14 1.14 53.5 0.03 0.08 43.2 12.6 <0.01 DG5 3.78 1.62 0.52 0.67 51.63 0.13 0.28 41.1 11.66 0.2 DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA

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ACTA CARSOLOGICA 40/3 2011 488 ment mapping has shown several types of deterioration phenomena (Kramar et al. 2010b). W eathering forms of the Drenov Gri limestone, documented according to the Fitzner classication (Fitzner & Heinrichs 2002), are giv en in Tabs. 3 & 4 for the altar and the portal, respectively. Examples of weathering forms are shown in Fig. 3. Indoor monument Back weathering and break out represent very fre quent weathering forms characterizing loss of stone material in the case of limestone exposed to the indoor environment. Especially in the lower part of the altar, back weathering is strikingly high in a range of capil lary rise. Loss of stone material is also represented by re lief in lower parts. Loose salt deposits, crusts, and color change are the main weathering forms characterizing deposits and discoloration Eorescence (salt crystalli zation on the surface) and suborescence (salt crystal lization under the surface) are frequent in the lower part of the altar in a range of capillary moisture levels and in areas with joint mortar and walls (Figs. 3a & 3b) as also observed in the case of Lesno Brdo limestone (Kramar et al. 2010b). A color change to grey occurs in the upper part of the monument. W hite crusts are present about 1 m from the ground. Crumbling, aking, and splitting are most abundant in a group characterizing stone detach ment. Depending on the orientation, the detachment of the limestone is expressed as splitting (not parallel) or aking (parallel to the surface). Flaking of the limestone represents one step before loss of material (Fig. 3b). e most intense detachment of the material occurs at about 0.5 m from the oor, although it is distributed to a smaller extent over the whole monument. In some ar eas ssures are notable, indicating salt crystallization un der the surface. Lower parts of the altar in particular are extensively damaged by salt crystallization, while in the upper parts the original color has been bleached. Flaking is leading to back weathering. Outdoor monument Black limestone exposed to the outdoor environ ment is aected by a wide range of weathering forms as well. On the exposed areas of the portal (ngers, gures of angels, and decorative balls) there is an intensive loss of stone material, expressed as back weathering and re lief. Color change (discoloration), crusts, and biological colonization are the main weathering forms character izing deposits and color change. e monument shows changes in the original black color, which turns to grey over the whole area. On sheltered areas of the monu T ab. 3: Weathering forms of limestone exposed to the indoor environment. Groups of weathering forms Main weathering forms Individual weathering form Terminology Denition Group 1Loss of stone material Back weathering Uniform loss of stone material parallel to the original stone surface Back weathering due to loss of scales Back weathering due to loss of pieces Back weathering due to loss of crumbs Relief Morfological change of the stone surface due to partial or selective weathering Rounding/notching Weathering out of stone components Alveolar weathering Break out Loss of compact stone fragment Break out due to natural cause Group 2Discoloration/ Deposits Discoloration Alteration of the original stone colour Bleaching Loose salt deposits Poorly adhesive deposits of salt aggregates Eorescences Sublorescences Crust Strongly adhesive depostis on the stone surface Light-coloured crust tracing the surface Group 3Detachment Crumbly disintegration Detachment of larger compact stone pieces of irregular shape Crumbling Flaking Detachment of small, stone pieces (akes) paralel to the stone surface Single akes Detachment of stone layers depending on stone structures Detachment of larger stone sheets or plates following the stone structure Splitting up Group 4Fissures/ Deformation Fissures Individual ssures or systems of ssures due to natural or constructional couses Fissures independent on stone structure S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI

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ACTA CARSOLOGICA 40/3 2011 489 ment, compact black crusts are presented (Fig. 3c). e presence of microorganisms is documented in areas of the monument which are exposed to humidity with out direct light (Fig. 3d). Crumbling, aking, and crust detachment are the most abundant types in a group of stone detachment. Crumbling is also seen on volutes on the upper parts of the monuments. Detachment of black crust is present in some areas. WEATHERING PRODUCTS AND MECHANISMS Indoor monument X-ray diraction analysis conrmed the pres ence of gypsum and soluble salts of the MgSO 4 .nH 2 O series, such as starkeyite (MgSO 4 .4H 2 O), pentahy drite (MgSO 4 .5H 2 O), and hexahydrite (MgSO 4 .6H 2 O) (Tab. 1). Gypsum occurs in both suborescence and eo rescence, whereas the absence of magnesium sulfates hy drates is obvious in suborescence and they are not pres ent on the outdoor monument. It is important to note that niter, forming the eorescence on another lime stone (Lesno Brdo Limestone) of the same monument, is absent here (Kramar et al. 2010a, Kramar et al. 2010b). W hite crust also consists of gypsum, and occurs in out door and indoor monuments. Black crust is composed of gypsum crystals which are orientated perpendicular to the limestone surface. Platy crystals are 50 m in size. Crusts exhibit denser crystal aggregation with respect to the crystals of the eores cence. e origin and growth of the sulfated crusts have been widely studied in past years (Camuo et al. 1983). W hite crust is composed of gypsum (Fig. 4a). For crust formation the amount of supplied solution must be high: the substrate is very humid or wet when substrate forms (Arnold & Zehnder 1985), suggesting a sucient water supply in that area of the altar, whereas outdoors it occurs in sheltered areas. A network of ssures occurs about 100 m below the limestone surface. e s sures are about 5 to 50 m thick. Fissures under the surface of the limestone are lled with gypsum. Crystals of gypsum in ssures are oriented parallel to the ssure wall and are not in contact with limestone, while on the surface they are orientated perpendicular to the surface. Eorescence consists of acicular and hair-like crystals, indicating so-called whisker growth (Arnold & Kueng 1985). e successive penetration of solutions oc curs through migration from the ground by capillarity, and they are then evaporated, with soluble salts precipi tation. W hisker growth indicates either a very low water content of the porous substrate, a very low water supply, or a very slow evaporation rate. In contrast to the whis ker growth, crust formation requires a comparatively high solution supply. Fluy eorescence preferentially develops on dense looking substrate (Arnold & Kueng 1985). Polymineral eorescence consists of magnesium sulfate hydrates and gypsum. Hexahydrite and epsomite are common hydrates on the Earths surface that occur at ambient temperature and moderate and elevated rela tive humidity (RH) (Chipera & Vaniman 2007). W ith the exception of hexahydrite-epsomite, transformations between the various species involve more than simple removal of water, requiring signicant crystal structure rearrangement and overcoming of activation energy bar Fig. 3: Examples of weathering forms of the black micritic limestone. (a) Crystallization of salts on the surface (eorescence), stone element on altar. e image is about 20 cm in width. (b) Sublfo rescence resulting in aking of the limestone. e presence of surface ssures of limestone indicates salt crystallization under the limestone surface. e image is about 10 cm in width. (c) Black crust, detail of the portal. e image is about 10 cm in width. (d) Presence of microroganisms, foot of the H ercules statue of the portal. e image is about 30 cm in width. DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA

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ACTA CARSOLOGICA 40/3 2011 490 mainly restricted to the upper 400 m of the limestone surface. e mechanisms of salt decay are based on the pressure exerted on the walls of the stone pores when the salt crystallizes with repeating cycles. e solutions can migrate inwards in the stone by capillarity. On the stones contaminated by salts, at relative humidities >75% the presence of water vapor until the salts dissolve ensures a weak supercial penetration by capillary suc tion, evaporation of the solution, and participation of the gypsum inside the pores and cracks underneath the supercial crystals (Chabas & Jeanette 2001). Crystalli zation of these salts and hydrationdehydration trans formations, depending mainly on relative humidity and temperature, lead to aking and alveolar weathering of the limestone. As joint mortars between the stone ele ments of the altar contain high quantities of soluble calcium and magnesium (Kramar et al. 2007), they are thus considered as a potential source of these damag ing salts. e deterioration occurs as a consequence of crystallization close to the surface of the stone. Any sub sequent decrease in the ambient RH at the exposed sur face will cause the dissolved salts to migrate towards the surface, as surface water evaporates to compensate for the change in RH (Colston et al. 2001). Slow evapora tion promotes salt crystallization inside the stone with a greater disruptive eect. e factors determining the occurrence of eorescence over cryptoorescence are kinetically driven (Colston et al. 2001). W hen the petro physical properties allow fast and long-distance transfer riers. Close to room temperature, epsomite is the stable form in the presence of liquid water. Under dry condi tions, epsomite can dehydrate to form hexahydrite and nally the monohydrate kieserite (Juling et al. 2000). W hile the occurrence of magnesium sulfate hydrates is limited to the bottom parts of the altar, gypsum appears to occur on higher parts of the altar as well. It is pres ent in ssures, and leads to the scaling and crumbling. e presence of gypsum also in the areas of the capillary range suggests that the solution migrates and water also penetrates through other mechanisms such as condensa tion. SEM micrographs of black limestone in transverse section show a highly fractured limestone surface, while the inner part of the limestone shows a low porosity and a compact structure. Humidity and temperature uctu ations cause cycles of salt precipitation and dissolution. During the one-year monitoring period, the RH ranged from a maximum of 100 % to a minimum of 40%, whereas the ambient temperature ranged from -0.6 C to 28.1 C (Kolenc 2006). As gypsum is less soluble it crystallizes rst, whereas magnesium sulfate hydrates stay longer in a solution. In our context, supersatura tion probably results from evaporation. Salt weather ing is primarily a mechanical process that disrupts the physical integrity of stone, whereas chemical weather ing processes degrade specic chemical elements within the crystal lattice of constituent minerals (W arke et al. 2006). Crystallization of salts below the stone surface (suborescence) is leading to aking. e damage is T ab. 4: Weathering forms of limestone exposed to the outdoor environment. Groups of Main weathering forms Individual weathering form weathering forms Terminology Denition Group 1Loss of stone material Back weathering Uniform loss of stone material parallel to the original stone surface Back weathering due to loss of scales Back weathering due to loss of pieces Back weathering due to loss of crumbs Relief Morfological change of the stone surface due to partial or selective weathering Rounding/notching Weathering out of stone components Group 2Discoloration/ Deposits Discoloration Alteration of the original stone colour Bleaching Crust Strongly adhesive depostis on the stone surface Dark-coloured crust tracing the surface Biological colonization Microbiological colonization Group 3Detachment Crumbly disintegration Detachment of larger compact stone pieces of irregular shape Crumbling Flaking Detachment of small, stone pieces (akes) paralel to the stone surface Single akes Detachment of crust with stone material Detachment of crusts with stone material sticking to the crust Detachment of a dark coloured crust tracing the stone surface Group 4Fissures/Deformation Fissures Individual ssures or systems of ssures due to natural or constructional couses Fissures independent on stone structure S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI

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ACTA CARSOLOGICA 40/3 2011 491 and the porosity allows a ow that compensates evapo ration, salts crystallize on the surface without causing signicant weathering. In contrast, when the capillary supply does not compensate evaporation, salts precipi tate inside the stone, under the surface, where loose ness of supercial crystals results (Lewin 1982; Chabas & Jeanette 2001). e lowering of the RH signicantly aects the decay, and a combination of a lower RH and a longer drying time produces the greatest rate of decay (Colston et al. 2001). As proved by SEM examination (Fig. 4b), in areas of veins with clay-mica group minerals (presence of K, Al, Si) disrupture of the limestone occurs. Due to water absorption/desorption clays are swellowing resulting in irreversible crumbling of the limestone. e phenomena are observable outdoors as well as indoors. Sometimes gypsum is evidenced in the area enriched with clay min erals. Fig. 4c shows an area of about 100 m of granular disintegration that is respon sible for the color change of the limestone. Outdoor monument On limestone exposed to the outdoor environment, decohesion between the grains is observed (Fig. 4d). A change of the original black color into grey is observed on both monuments. In the majority of cases the situa tion is seen due to the granu lar disintegration, whe reas in a small number of cases in the indoor environment it is due to the surface cover of a thin crust of salts. e surface is rough, with dis continuities characterized by a system of intraand intercrystalline microfractures in the rst 200 m. Marinoni (2007) reported that decohe sion and higher porosity are present on the outer layer of the samples, suggesting that thermal weathering may play a key role in black limestone discoloration. is was also suggested by Zehnder (2003), whereas other researchers ascribed this phenomenon as the oxidation of organic matter, with hydrocarbons be ing the least stable carbon pigments (W inkler 1997). e reasons for granular decohesion could be ascribed to mechanical weathering, in particular thermal cracking of the calcite crystals. Limestone can have good proper ties in pure water but deteriorates rapidly when freezing in a salt solution (Lindquist et al. 2007). Damage caused by freezing in salt solutions will be dependent on the nanopores, while the damage caused by freezing in pure water is more dependent on the micron-sized porosity (Lindquist et al. 2007). In the outdoor samples, the alter nation of gypsum and limestone occurs in bands of 10 to 20 m, suggesting that gypsum lled the areas between the laminas of the limestone (Fig. 4f). Dissolution of cal cite crystals under the gypsum crust, which is about 300 m thick, is evident. Delamination and disrupture in ar eas with clay minerals occur both outdoors and indoors, Fig. 4: SEM microphotographs of deteriorated limestone. (a) Crystallization of gypsum at the surface of limestone. Sample JAL176. (b) I rreversible ssures occur in veins of clay minerals; the presence of K, Al, and Si evidence the sericite. Na and Fe are present in minor amounts. Sample JAL122. (c) G ranular disintegration in She area about 100 m under the surface of the limestone, indoors. Sample JAL125. (d) H ighly fractured black limestone surface outdoors. G ranular disin tegration caused due to the anisotropic thermal expansion. Sample SEP1. (e) Biopitting. Sample SEP7. f) Alternation of layers of gypsum with limestone. Sample SEP3. DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA

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ACTA CARSOLOGICA 40/3 2011 492 similar as in the case of Lesno Brdo limestone (Kramar et al. 2010b). e results indicate that microorganisms which form green patina on the limestone surface are represent ed by unicellular cyanobacteria (Chroococcale) and uni cellular (Bracteacoccus) and lamentous (Ulotrichales) Chlorophyceae. It was reported (Ortega-Calvo et al. 1995) that cyanobacteria and Chlorophyceae comprise two of the most common groups of algae found on build ing stones, as endolithic habitats provide good growth conditions in otherwise harsh environments. Cyanobac teria and algae are normally pioneers in colonizing stone surfaces because they are autotrophic organisms (Pera za-Zurita et al. 2005). On the other hand, organic acids, which are produced in their metabolic processes, could contribute to the limestone dissolution. Indeed, biopit ting is evidenced from samples taken from the outdoors, as proved by SEM analysis (Fig. 4e). Although limestone consist mainly of calcite, it can show signicant variations in compositions, texture and structure, resulting in complex and contrasting weather ing behavior, as suggested by the results of the present study and supported with the observations of previously studied Lesno Brdo limestone (Kramar et al. 2010b). W ide range of identied weathering forms indicates that the primary cause of extensive weathering is mechanical rather than chemical weathering. Processes of mechani cal weathering are fully operative in polluted environ ments, but this is also believed for a natural environment (Gauri & Bandyopadhay 1999). Study indicates that weathering and deterioration of the limestone in urban environment is controlled in great extent by the environmental factors, one of the most important weathering factors being water and sol uble salt crystallization. Still, intrinsic properties of the limestone inuence the type of the weathering form; for instance veins lled with phyllosilicates will cause a dis ruption of the stone. As the latter was observed at mon uments made of Drenov Gri limestone and Lesno Brdo limestone (Kramar et al. 2010b), exposed to indoor and outdoor environment, the same behavior it can be ex pected in natural environments as well. Also, karst and related phenomena are sensitive to environmental con ditions (Faniran & Jeje 1983), where dissolution (relief, erosion) and secondary deposits (crust formation) are known. One characteristic of the natural environment pertinent to rock weathering is that lacks the industrial auent sulfur dioxide and nitrogen dioxide, which in uence the weathering signicant in the urban environ ment. CONCLUSIONS Besides calcite as the main mineral of the limestone, dolomite and non-carbonate components including py rite, clay minerals, quartz, and organic substances were determined. e micritic grains of the limestone and its very dense structure are consistent with small pore radii, low porosity, and small capillary kinetics. e small po rosity and weak water transfer kinetics mean that solu tion transport is low, increasing the possibility that high oversaturation ratios are reached below the stone surface, precipitating as suborescence. A great variety of weathering forms characterized by loss of stone material, discoloration/deposits, and detachment were identied on the monument. Besides color change, soluble salts formation was found to be the most important degradation of the investigated monu ments, outdoors as well as indoors. Salt causes dierent types of degradation under dif ferent environmental conditions. Gypsum crystallization leads to scaling and crumbling of the limestone as well as crust and eorescence formation, whereas magnesium sulfate hydrates occur only as eorescence indoors. Disrupture of the limestone in areas with clay minerals occurs both outdoors and indoors. e change of the original black color into grey is attributed to the granular integration and occasionally to the thin surface covering with gypsum (indoors). Granular decohesion is more obvious outside, where the changes in temperature are higher. ere is no evidence of microorganisms inside, whereas the presence of microorganisms (Cyanobacteria and Chlorophyceae) outdoors resulted in biopitting. Although the limestone yielded good physical char acteristics, the deterioration is relatively high in both in door and outdoor environments. Deterioration is mainly controlled by the environmental factors, such as water and crystallization of soluble salts, but is also inuenced by the intrinsic parameters, such as veins of phyllosili cates (mineral composition). Urban environment represent a specic environ ment where karstic processes are enhanced due to the presence of anthropogenic pollutants. However, the properties of the limestone, such as presence of specic mineral or porosity will inuence the behavior of the weathering in the similar manner also in the natural ru ral environs. S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI

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ACTA CARSOLOGICA 40/3 2011 493 is research has been supported nancially by the Slov enian Research Agency, under contract number 321105-000545. e authors also are grateful to Joe Drear for performing the necessary sampling works on the se lected monuments. Many thanks are due to Alexandre Franois from Laboratoire de recherche des monuments historiques, France, for the identication of microorgan isms. e Joef Stefan Institute is gratefully acknowl edged for providing experimental support. e photo graph shown in Fig. 1b is included by kind permission of Valentin Benedik. ACKNO WLEDGMENTS REREFENCES Adamson, A.W & A.P. Gast, 1997: Physical Chemistry of Surfaces .J. W iley & Sons, pp. 353, New Y ork. Arnold, A. & A. Kueng, 1985: Crystallization and habits of salt eorescences on walls, I., Methods of inves tigation and habits.In: Felix, G. (ed.) Proc 5 th I nt Congress D eterioration and Conservation of Stone 25..9.1985, Lausanne. Presses Polytechniques et Universitaires Remandes, 255, Lausanne. Arnold, A. & K. Zehnder, 1985: Crystallization and hab its of salts eorescences on walls II. Condition of crystallization.In: Felix, G. (ed.) Proc 5 th I nt Con gress D eterioration and Conservation of Stone 25..9.1985, Lausanne. Presses Polytechniques et Universitaires Remandes, 269, Lausanne. Barret, E.P., Joyner, L.G. & P.P. Halenda, 1951: e deter mination of pore volume and area distributions in porous substances. I. Computations from nitrogen isotherms.Journal of the American Chemical So ciety, 73, 373. Benavente, D., Garci del Cura, M.A., Fort, R. & S. Or dez, 2004: Durability estimation of porous build ing stones from pores structures and strength.En gineering Geology 74, 113. Brunauer, S., Emmett, P.H. & E. Teller, 1938: Adsorption of gasses in multimolecular layers.Journal of the American Chemical Society, 60, 309. Camuo, D., Del Monte, M. & C. Sabbioni, 1983: Origin and growth mechanisms of the sulphated crusts on urban limestone.W ater, Air, and Soil Pollution, 19, 351. Cardell, C., Celalieux, F., Roumpopoulos, K., Moropou lou, A., Auger, F. & R. Van Grieken, 2003: Salt in duced decay in calcareous stone monuments and buildings in a marine environment in SW France.Construction and Building Materials 17, 165. Cardell, C., Benavente, D. & J. Rodrguez-Gordillo, 2008: W eathering of limestone building material by mixed sulfate solutions. Characterization of stone micro structure, reaction products and decay forms.Ma terials Characterization, 59, 1371. Chabas, A. & D. Jeannette, 2001: W eathering of marbles and granites in marine environment: petrophysical properties and special role of atmospheric salts.Environmental Geology, 40, 359. Chipera, S.J. & D.T. Vaniman, 2007: Experimental stabil ity of magnesium sulfate hydrates that may be pres ent on Mars.Geochimica et Cosmochimica Acta, 71, 241. Colston, B.J., W att, D.S. & H.L. Munro, 2001: Envi ronmentally-induced stone decay: the cumula tive eects of crystallization-hydration cycles on a Lincolnshire oopelsparite limestone.Journal of Cultural Heritage, 4, 297. Cultrone, G., Rusoo, L.G., Calabr, C., Urosevic, M. & A. Pezzino, 2008: Inuence of pore system characteris tics on limestone vulnerability: a laboratory study.Environmental Geology, 54, 1271. Dunham, R.J., 1962: Classication of carbonate rocks ac cording to depositional texture, in Classication of Carbonate Rocks.American Association of Petro leum Geologist, Mem Serv, 1, 62. Faniran, A. & L. Jeje, 1983: H umid T ropical G eomorphol ogy: A Study of the gemorphological Processes and L andforms in Warm H umid Climates .Longman, pp. 414, London. Fitzner, B. & K. Heinrichs, 2002: Damage diagnosis on stone monuments-weathering forms, damage cat egories and damage indices.In: Prikryl, R. & H.A. Viles (eds.) Understanding and M anaging Stone D e cay, Proc I nt Conf Stone Weathering and Atmospheric Pollution Network (SWAPNE T 2001) 7..5.2001, Prague. Charles University in Prague, e Karoli num Press, 11, Prague. DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA

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ACTA CARSOLOGICA 40/3 2011 494 Folk, R.L., 1962: Spectra subdivision of limestone types, in Classication of Carbonate Rocks.American Association of Petroleum Geologist, Mem Serv, 1, 108. Gauri, K.L. & J.K. Bandyopadhyay, 1999: Carbonate Stone: Chemical, Behaviour, D urability, and Conser vation.J. W iley and Sons, pp. 284, New Y ork. Gams, I., 1974: K ras.Slovenska matica, pp. 395, Lju bljana. Gregg, S.J. & K.S.W Sing, 1982: Adsorption Surface Area and Porosity .Academic Press, pp. 303, London. Juling, H., Kirchner, D., Brggerhof, S., Linnow, K., Steiger, M., El Jarad, A. & G. Glker, 2000: Salt damage of porous materials: a combined theoretical and experimental approach.In: Kwiatkowski, D. & R. Lfvendahl (eds.) Proc 10th I nt Congress D eterio ration and Conservation of Stone, 1 27.6..7.2004, Stochholm. ICOMOS Sweden, 187, Stock holm. Kolenc, I., 2006: Sv. Jakob, Prikaz meritev temperature in vlage. Restavratorski center, Ljubljana. Kramar, S., 2006: Cerkev Sv. Jakoba Ljubljana: kapela sv. Franika Ksaverja ED 332 : poroilo o preiska vi malt in soli. Ljubljana: ZVKDS, Restavratorski center, Ljubljana. Kramar, S., Urosevic, M., Pristacz, H. & B. Mirti, 2010a: Assessment of limestone deterioration due to salt formation by micro-Raman spectrometry: applica tion to architectural heritage.Journal of Raman spectroscopy, 41, 11, 1441. Kramar, S., Mladenovi, A., Urosevic, M., Mauko, A., Pristacz, H. & B. Mirti, 2010b: Deterioration of Lesno Brdo limestone on monuments (Ljubljana, Slovenia).RMZ-Materials and Geoenvironment, 57, 1, 53. Kramar, S., Mladenovi, A., Kozamernik, M. & B. Mirti, 2010c: Durability evaluation of some Slovenian buildings limestones.RMZ-Materials and Geoen vironment, 57, 3, 331. Kramar, S., Mirti, B., Knller, K. & N. Rogan-muc, 2011: W eathering of the black limestone of histori cal monuments (Ljubljana, Slovenia): Oxygen and sulfur isotope composition of sulfate salts.Applied geochemistry, 26, 1632. Lewin, S.Z., 1982: e mechanism of masonry decay through crystallization.In: Baer, N.S. (ed.) Conser vation of H istoric Stone Buildings and M onuments .National Academic Press, pp. 120, W ashington DC. Lindquist, J.E., Malaga, K., Middendorf, B., Savukoski, M. & P. Ptursson, 2007: Frost resistance of natu ral Stone, the Importance of Microand NanoPorosity.[online] Available from: http://www.sgu. se/dokument/fou_extern/Lindquist-et-al_2007.pdf [Accesed 23.11.2008.] Lowell, S., Shields, J.E., omas, M.A. & M. ommes, 2004: Characterization of Porous Solids and Pow ders: Surface Area, Pore Size and D ensity .Springer, pp. 349, United States. Marinoni, N., Pavese, A., Bugini, R., & G. Di Silvestro, 2002: Black limestone used in Lombard architec ture.Journal of Cultural Heritage, 3, 241. Marinoni, N., Pavese, A., Riva, A., Cella, F. & T. Cerul li, 2007: Chromatic weathering of black limestone quarried in Varenna (Lake Como, Italy).Building and Environment, 42, 68. Marszaek, M. 2007: e mineralogical and chemical methods in investigations of decay of the Devonian Black marble from Dbnik (Southern Poland).In: Pikryl, R. & B.J. Smith (eds.) Building Stone D ecay: From D iagnosis to Conservation .Special Publications, 271. Geological Society, pp. 109. London. Miller, A., Dioniso, A. & M.F. Macedo, 2006: Primary bioreceptivity: A comparative study of dierent Portuguese lithotypes.International Biodeteriora tion and Biodegradation, 57, 136. Mladenovi, A., Mirti, B. & N. Viintin, 1999: ermal conductivity of selected types of Slovenian natural stone.RMZ-materials and Geoenvironment, 46, 1, 539-547. Nicholson, D.T., 2001: Pore properties as indicators of breakdown mechanisms in experimentally weath ered limestone.Earth Surfaces Processes and Landforms, 26, 819. Ortega-Calvo, J.J., Arino, X., Hernanerez-Marine, M. & C. Saiz-Jimenez, 1995: Factors aecting the weath ering and colonization of monuments by pho totrophic microorganisms.Science of Total Envi ronment, 167, 329. Peraza Zurita, Y ., Cultrone, E.G., Sanchez Castillo, S., Se bastian, E. & F.C. Bolivar, 2005: Microalgae associ ated with deteriorated stonework of the fountain of Bibatauin in Granada, Spain.International Biode terioration and Biodegradation, 55, 55. Ramov, A., 2000: Podpeki in rni ter pisani lesnobrdski apnenec skozi as .Mineral, pp. 115, Ljubljana. RILEM, 1980: Recommended tests to measure the dete rioration of stone and to assess the eectiveness of treatment methods.Material Structures, 14, 175 253. S ABINA KRAMAR, A NA MLADENOVIb, H ELMUT PRISTACZ & B REDA MIRTI

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ACTA CARSOLOGICA 40/3 2011 495 Sahlin T., Malaga-Starzec, K., Stigh, J. & R. Schouen borg, 2000: Physical properties and durability of fresh and impregnated limestone and sandstone from central Sweden used for thin stone oor ing and cladding.In: Fassina, V. (ed.) Proc 9 th I nt Congress D eterioration and Conservation of Stone 19..6.2000, Venice. Elsevier Science, 181, Venice. Scherer G.W ., 1999, Crystallization in pores.Cement and Concrete Research, 29, 1347. Sing, K.S.W., Everett, D.H., Haul, R.A.W., Moscou, L., Pierotti, R.A., Rouqurol, J. & T. Siemien iewska, 1985: Reporting physisorption data for gas/solid systems.Pure and Applied Chemis try, 57, 603 619. Skobe, S. & B. Mirti, 2005: Vpliv mineralne sestave in strukture na obstojnost apnencev kot naravnega kamna/e weathering durability of limestones as a function of their mineral composition and tex ture.RMZMaterials and Geoenvironment, 52, 4, 697. Steiger, M., 2005: Crystal growth in porous materials II: Inuence of crystal size on the crystallization pres sure.Journal of Crystal Growth, 282, 470. W arke, P.A., McKinley, J. & B.J. Smith, 2006: W eathering of building stone: approaches to assessment, predic tion and modeling.In: Kourkoulis, S.K. (ed.) Frac ture and Failure of Natural Building Stones Springer, pp. 3131, Dordrecht. W inkler, E.M., 1997: Stone in Architecture: Properties, D urability .Springer, pp. 313, Berlin. Zehnder, K. 2006: Greying of black polished limestone a case study to clarify the phenomenon.Zeitschri fur Kunsttechnologie und Konservirung. 2, 361 367. DETERIORATION OF THE BLACK DRENOV GRI LIMESTONE ON HISTORICAL MONUMENTS LJUBLJANA, SLOVENIA



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FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE NORTH ADRIATIC KARST FLORISTINA IN FUNKCIONALNA PRIMERJAVA KRAKIH PANIKOV IN KRAKIH TRAVNIKOV SEVERNOJADRANSKEGA KRASA Nataa PIPENBAHER 1 *, Mitja KALIGARI 1 & Sonja KORNIK 1 1 Biology Department, Faculty of Natural Sciences and Mathematics, University of Maribor, Koroka 160, SI-2000 Maribor, Slovenia, fax: +368 2 251 81 80, e-mails: natasa.pipenbaher@uni-mb.si, mitja.kaligaric@uni-mb.si, sonja.skornik@uni-mb.si Received/Prejeto: 6.2.2011 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 515, POSTOJNA 2011 Abstract UDC 633.2.03(497.472) Nataa Pipenbaher, Mitja Kaligari & Sonja kornik: Floris tic and Functional comparision of karst pastures and karst meadows from the North Adriatic Karst In the present study, we compared the species richness and the oristic and functional composition of two types of extensively managed, species rich dry grasslands (class Festuco-Brometea ) from the North Adriatic Karst: karst pastures (alliance Sat ureion subspicatae ) and karst meadows (alliance Scorzonerion villose ). Karst pastures are characterized by shallow rocky soils, high pH, and dry, warm conditions, whereas karst meadows have developed on deeper soil, with more humus and moisture and neutral to alkaline pH. e data set included a table with 100 phytosociological relevs of the studied grasslands and a matrix with 15 functional traits determined for 180 plant spe cies. W e found high species richness in these grasslands but no statistically signicant dierences in species richness between karst pastures and meadows. Dierences in oristic composi tion were analysed with Detrended Correspondence Analysis, which supported a clear division between the two vegetation types and indicated that species composition could best be explained in terms of soil humidity and nutrient availability. W e also detected several dierences in plant functional traits between meadows and pastures. Some of the traits indicate greater resource availability on karst meadows (in particular, high SLA, low LDMC). In contrast, karst pastures have more slow-growing species with a combination of traits that can be interpreted as an avoidance strategy in relation to distur bance (e.g., grazing) in low productive habitats (e.g., low SLA, high LDMC, early owering species and plants with rosette). A lower relative proportion of competitors (C) and ruderals (R), and a higher relative proportion of stress-tolerators (S) in karst pastures also suggested that these grasslands generally experience higher intensities of stress when compared to karst meadows, presumably owing to lower resource availability on Izvleek UDK 633.2.03(497.472) Nataa Pipenbaher, Mitja Kaligari & Sonja kornik: Floristina in funkcionalna primerjava krakih panikov in krakih travnikov severnojadranskega Krasa Predstavljamo rezultate raziskave, v kateri smo primerjali vrstno pestrost, oristino in funkcionalno sestavo med dvema tipoma suhih travi (razred Festuco-Brometea ) severnojadran skega Krasa krakimi paniki (zveza Satureion subspicatae ) in krakimi travniki (zveza Scorzonerion villosae ). Za krake panike so znailna plitka kamnita tla z bazino kemijsko reakcijo in suhe ter tople razmere, medtem ko so kraki travni ki razviti na globljih in bolj hranilnih tleh, z ve vlage, kemijska reakcija tal pa je nevtralna do zmerno kisla. Podatki so obsegali tabelo s 100 tocenolokimi popisi krakih panikov in travnikov ter matriko s 15 funkcionalnimi potezami, doloenimi za 180 rastlinskih vrst. Ugotovili smo visoko vrstno pestrost prouevanih travi, hkrati pa ni bilo najdenih statistino znailnih razlik v vrstni pestrosti med krakimi paniki in krakimi travniki. Razlike v oristini sestavi smo ugotavljali z DCA analizo, ki je potrdila jasne razlike med obema vegetacijskima tipoma in nakazala, da se vrstna sestava popisanih sestojev spreminja vzdol gradientov vlanosti in hranilnosti tal. Ugotovili smo tudi razlike v funkcionalnih potezah rastlin med krakimi travniki in paniki. Na osnovi nekaterih funkcionalnih potez (zlasti vijih SLA, nijih LDMC vrednosti) lahko sklepamo, da so kraki travniki bolje zaloeni s hranili. Kraki paniki imajo v primerjavi z njimi ve poasi rastoih vrst s kombinacijo potez (npr. nizke SLA in visoke LDMC vrednosti, zgodnje cvetenje, rozete), ki jih razlagamo kot strategija izo gibanja motnji (npr. pai) v slabo produktivnih habitatih. Tudi nija delea kompetitorskih vrst (C) in ruderalk (R) ter hkrati viji dele stres-toleratork (S) na krakih panikih potrjujejo, da so v teh habitatih bolj stresne razmere kot na krakih travni kih, kar je po vsej verjetnosti posledica slabe preskrbljenosti rastlin z vodo in hranilnimi snovmi zaradi kamnitih in plitkih

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ACTA CARSOLOGICA 40/3 2011 516 stony, shallow soil. W e could conclude that karst meadows and pastures dier signicantly in both oristic composition and functional trait means, owing to their distinctive land-use (disturbance) and environmental conditions. Keywords: dry grasslands, plant functional traits, FestucoBrometea SW Slovenia, DCA, CSR plant strategies. tal. Zakljuimo lahko, da obstajajo znailne razlike v oristini in funkconalni sestavi med krakimi travniki in paniki kot posledica razlik v rabi (konja in paa) ter razlik v okoljskih dejavnikih. Kljune besede: suha travia, funkcionalne poteze rastlin, Festuco-Brometea JZ Slovenija, DCA analiza, CSR strategije rastlin. INTRODUCTION e Karst grasslands of the North Adriatic Karst are known as one of the richest plant communities among grasslands (Kaligari et al. 2006). is diversity has been demonstrated, emphasized and partly explained on many occasions (Horvat & Horvati 1934; Ferlan & Giacomini 1955; Horvati 1973; Horvat et al. 1974; Poldini 1980; Piskernik 1988; Poldini 1989; Piskernik 1991; arni & Kaligari 1992; Kaligari 1994; Kaligari & Poldini 1996; Kaligari 1997; Kaligari & kornik 2002; Pipenbaher et al. 2008; kornik et al. 2010) in terms of the specic biogeographic position of these grasslands, being in the transitional area of the Mediterranean basin, the Dinaric mountains and Central Europe. e contingent of subMediterranean-Illyrian species is numerous; in addition there are other large interesting contingents of species: e.g., Mediterranean-Pontic or Mediterranean-Pannon ian species (Poldini 1989). As has been established by phytosociologists, the internal variability among grass lands is also very high. e North Adriatic grasslands, which all belong to the order Scorzoneretalia villosae Horvati 1975 (class Festuco-Brometea ), were therefore divided in to two separate alliances: Satureion subspica tae Horvat 1962 and Scorzonerion villosae Horvati 1949 (Kaligari 1997). e oristic dierences between them have already been described as substantial (Kaligari 1997), and the two alliances have been considered not as biogeographic parallels (as alliances sometimes are), but as ecological parallels (Kaligari & kornik 2002; Kaligari et al. 2006). e Satureion subspicatae is com post by sclerophyllous plants and represents extensive, semi-natural pasture, whereas Scorzonerion villosae is the more mesophilous alliance, representing extensive unfertilized hay meadows (Kaligari & Poldini 1996; Kaligari 1997). It should be emphasized that natural conditions on the karst terrain also pre-determine to a considerable ex tent also the potential land-use of both vegetation types: Scorzonerion villosae grasslands, which appear in the karst polje, sink-holes and within the karst valleys where natural soil accumulation and natural soil fertility are higher, were more prone to mowing. In contrast, the ma jor management form in the Satureion subspicatae alli ance was moderate grazing, as this has developed on the slopes, where because of erosion, soils are shallow and rocky and denitely not convenient for mowing. It has been well established that dierences in es sential constraints, e.g. weather, topography, soil con ditions and disturbance regime, lead to dierent plant communities. e species combination assemblage is based on ltering plants with particular sets of func tional traits from the available species pools (Lavorel et al. 1997; Lavorel & Garnier 2002, de Bello et al. 2005). It was emphasized by Keddy (1992), Lavorel et al. (1997) and especially by Diaz et al. (2002) that plant functional traits and their combinations rather than species iden tities are the units that are aected by ecological pro cesses. In order to gain insight into functional traits, Grimes CSR triangle theory should be considered as well (Pipenbaher et al. 2008). is scheme is built on the assertion that three major determinants of species exist: competition (C), abiotic stress (S) and disturbance (R). e position of any species can be displayed upon a tri angular ordination diagram (Grime 1974; Hodgson et al. 1999; Grime 2001). Each strategy is characterized by a distinct set of ecological, morphological and physiologi cal traits and is found in species occupying habitats of a particular kind (Grime et al 1988). In the Northern Adriatic Karst area the distinction between pastures and meadows based on plant species composition has been conrmed (Kaligari 1997), but the degree of functional dierentiation between the two grassland types has not yet been considered and docu mented. erefore, we would like to answer the follow ing questions: 1. To what degree do the karst pastures and meadows dier on the basis of their species compo sition? 2. To what degree if at all do they dier on the basis of most important plant functional traits? 3. W hat could be revealed by a oristic and functional compari son between karst meadows and karst pastures? 4. W hat kind of plant ecological strategies are characteristic of karst meadows and karst pastures? N ATAA PIPENBAHER, MITJA KALIGARI & SONJA KORNIK

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ACTA CARSOLOGICA 40/3 2011 517 STUD Y AREA Slovenia, situated at ca. 46N, 14E in the contact area between the Alps, the Dinarides, and the Mediterranean and Pannonia plains, has a relief consisting of plains, hilly regions, highlands, mountains etc. 40% of the land is underlain by carbonate rocks, mainly well karstied and dolomitised (W atts 2004). e position of the study area lies between the Adriatic Sea and the Pre-Dinaric region in Slovenia. It represents the most north-western part of the Balkan Peninsula. It consists of the limestone Karst plateau (Low Karst) and mountains (High Karst). e Karst Plateau is 100 m a.s.l. and is characterized by its geomorpho logical phenomena (rocks, karst poljes, dolinas, caves, etc.) (Mihevc 1997). e climate is sub-Mediterranean (Ogrin 1995). Precipitation varies from 1,356 mm and up to 1,781 mm on the High Karst. e strong bora wind causes desiccation and erosion. e mean annual tem perature is 8.5 C (ARSO 2009). Poldini (1989) charac terized the climate as transitional between Mediterra nean and continental pre-Alpine, with rainy cool winters and long, dry summers. Floristically the study area is a contact area be tween the Mediterranean and Euro-Siberian biogeo graphical zones, having characteristics of both zones. e natural vegetation is deciduous forest, constitut ed by various species of oak (Quzel & Medail 2003). Mixed deciduous thermophilous woodlands prevail at lower altitudes up to 8001,000 m. In this belt tree species such as Quercus pubescens Carpinus orienta lis Ostrya carpinifolia Fraxinus ornus and, rarely, Acer monspessulanum and Pistacia terebinthus constitute the sub-Mediterranean vegetation below the beech forest zone. Temperate deciduous trees such as beech nowa days represent the higher altitudinal belt above 800 1,000 m. e derivates of the deciduous forests are dry and semi-dry grasslands of the order Scorzoneretalia vil losae Horvati 1975 (class Festuco-Brometea ) (Kaligari 1997). Most of the non-forest vegetation belongs to pas tures (alliance Satureion subspicatae Horvat 1962). e association Carici-Centaureetum rupestris Horvat 1931 is the most widespread and common in the study area. It is characterized by shallow rocky soils, high pH, and dry, warm conditions. Productivity is very low. Due to hydric stress, the vegetation becomes almost completely dry in the summer (Kaligari 1997). Dry and semi-dry meadows (alliance Scorzonerion villosae Horvati 1949), association D anthonio-Scorzoneretum villosae Horvat et Horvati (1956) 1958 have developed on deeper soil, where there are more humus and moisture and neutral to alkaline pH (Kaligari et al. 2006). VEGETATION SURVE Y W e analysed 48 relevs of karst meadows association D anthonio-Scorzoneretum villosae (relevs from 1 (Tab. 1) are in book Kaligari 1997) and 52 relevs of karst pastures association Carici-Centaureetum rup estris Horvat 1931 (relevs from 10-62 (Tab. 4) are in Kaligari 1997). On each site, vascular plants were sam pled using a seven-point species cover-abundance scale (r, +, 1 to 5), according to the Braun-Blanquet method (Braun-Blanquet 1964, W estho & Van der Maarel 1973, Dierschke 1994) within 5 x 5 m quadrates (relevs). Plant species richness was noted as the number of species re corded in each relev. Taxonomic nomenclature follows Martini et al (2007); syntaxonomic nomenclature fol lows Kaligari (1997). P LANT FUNCTIONAL TRAITS In choosing key traits, we followed various literature sources (Hodgson et al. 1999; Kahmen et al. 2002; Cor nelissen et al 2003). W e selected 15 traits for each spe cies. Traits were chosen from our own database (protocol standardized by Cornelissen et al 2003). Information on species traits was also taken from two existing trait data bases: BiolFlor (Klotz et al. 2002) and LEDA (Kleyer et al. 2008). Species were characterized by basic traits as well as composite traits (such as C-S-R strategy). Owing to the variety of species sets, we focused on traits that were easy to measure. e traits selected were as follows: life form, life cycle, growth form, spinescence, hairiness, plant height, specic leaf area (SLA), leaf dry mat ter content (LDMC), owering start, owering end, owering period, leaf persistence, leaf anatomy, leaf form and CSR strategy. e CSR strategy scheme provides a system for classifying herbaceous plants as having strategies adapt ed to intense competition (C), abiotic stress (S) and frequent disturbance (R) (Grime 1974; McIntyre et al ., 1995; Lavorel et al 1997; W estoby 1998). To determine CSR functional types for 122 plant species recorded in 100 analysed vegetation relevs, we used a rapid method published by Hodgson et al. (1999). e list of traits with the description of classes in the matrix and the sources of information are presented in Tab. 1. Categorical traits were all transformed into binary variables, with one for each possible level of the factor. In this way the number of traits in the matrix in creased from 15 to 37. For plant abundance, data from the seven-point Braun-Blanquet ordination scale were rst converted into a 1 to 7 scale (Pyek & Pyek 1991). All plant species occurring only in one relev were re moved to exclude casual occurrences from the analysis. MATERIALS AND METHODS FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE ...

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ACTA CARSOLOGICA 40/3 2011 518 trait j for species i (Ricotta & Moretti 2010). is opera tion results in 37 new traits by 100 relevs matrix (matrix CWM). STATISTICAL ANAL Y SIS Detrended Correspondence Analysis (DCA, Hill & Gauch 1980) ordination was applied to the species-relevs ma trix. e intention was to provide a visual representation of dierences in oristic composition between the two vegetation types being studied. To test dierences in the specic plant functional traits between karst meadows and karst pastures, the CWM values of all 37 functional traits were analysed with a Student's ttest for independent samples (SPSS I nc. 2006). Only traits (n = 29, Tab. 2) with signicant dif ferences in their CWM values between karst meadows To assess the functional trait composition of relevs, we combined the species by relevs matrix with the cor responding species by traits matrix. is was done by calculating the community-weighted mean trait val ues (CWM) for each relev as the average of trait val ues weighted by the relative abundance of each species (Garnier et al. 2004; Lep et al 2006; Lavorel et al. 2008; Moretti et al. 2009; Ricotta & Moretti 2010). e metric is computed simply as where CWM jk is the community-weighted mean value of trait j at site k; p ik is the relative abundance of species i (i = 1, 2, ...., S) at site k; and x ij is the value of T ab. 1: Plant traits, recorded on 180 vascular plant species from karst meadows and karst pasture (SW Slovenia). Scales of measure ment were originally categorical (cat), continuous (cont) or binary (bin). Traits Abbreviation and description Data source Life form LF_Ge = geophytes; LF_He = hemicryptophytes Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Martini et al. 2007 Life cycle LC_Annu = annual; LC_Bien = biennial; LC_Pere = perennial Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Martini et al. 2007 Growth form GF_Tuss = tussocks; GF_Rose = rosette; GF_le_st = leafy stem; GF_ro_le = rosette and leafy stem Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Martini et al. 2007; Rothmaler 1995 Spinescence Spi_pres = present; Spine_0 = none Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Martini et al. 2007 Hairiness Hair_low = low; Hair_hig = high; Hair_0 = no Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Poldini 1991; Martini et al. 2007 Plant height Plan_hig = cm Own measurements Specic leaf area SLA = mm 2 /mg Own measurements; LEDA database (Kleyer et al. 2008) Leaf dry matter content LDMC = mg/g Own measurements; LEDA database (Kleyer et al. 2008) Flowering start Flo_star = months Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Poldini 1991; Martini et al 2007 Flowering end Flo_end = months Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987, Poldini 1991; Martini et al. 2007 Flowering period Flo_peri = months Hegi 1958, 1963, 1964, 1965, 1966, 1974, 1987; Poldini 1991; Martini et al. 2007 Leaf persistence LP_1 = spring green; LP_2 = summer green; LP_3 = overwintering green; LP_4 = persistent green Own measurements; BiolFLor (Klotz et al. 2002) Leaf anatomy LA_2 = scleromorphic; LA_3 = mesomorphic Own measurements; BiolFLor (Klotz et al. 2002) Leaf form Lea_for3 = grass-like; Lea_for4 = longleaf; Lea_for5 = simple; Lea_for7 = lobate; Lea_for8 = pinnatid; Lea_for10 = palmate; Lea_for11 = pinnate; Lea_for12 = bipinnate BiolFLor (Klotz et al. 2002) C-S-R strategy C = competitors; S = stress-tolerators; R = ruderals Own measurements N ATAA PIPENBAHER, MITJA KALIGARI & SONJA KORNIK

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ACTA CARSOLOGICA 40/3 2011 519 and karst pastures ( Student s t-test) were used for further analysis. e CWM matrix was further analysed by means of Principal Component Analysis (PCA) (Goodall 1954). Gradient length for the rst PCA axis of ordination was 0.22, indicating that linear ordination methods are suit able for the analysis. e ordination methods (PCA, DCA) and visual ization of their results were carried out using the Canoco and CanoDraw programs (ter Braak & milauer 2002). RESULTS S PECIES RICHNESS AND SPECIES COMPOSITION e total number of vascular plant species recorded in 100 relevs of these particular karst grasslands was 180, with 144 on karst meadows (mean = 34 + 5,6 s. d. per relev, N=48) and 134 (mean = 31 + 8.7 s.d. per relev, N = 52) on karst pastures. W e found no statistically signicant dif ferences in species richness between the grassland types (P = 0.100). All species are listed in Appendix 1. ere are 98 common species, 46 species exclusive to the karst meadows and 36 exclusive to the karst pastures. Dierences in oristic composition were rst analy sed with DCA analysis of the 180 species x 100 relevs matrix. Eigenvalues for rst two DCA axes are 0.530 and 0.271. e rst two axes together explain 23.8% of the variability in the species composition. DCA ordina tion is shown in Fig. 1. e DCA Axis 1 of this analysis showed evident divergence, which supports a clear divi sion between these two grassland types: karst pastures and karst meadows. e species with the lowest scores (-X) are those frequent in karst meadows, such as D acty lis glomerata, Ononis antiquorum, Peucedanum cervaria, Chrysopogon gryllus, G alium verum Species that are as sociated with relevs of karst pastures received the high est scores (+X) in the rst DCA: for example, G lobularia cordifolia, Anthyllis jacquenii, Satureja liburnica, Sesleria juncifolia, G enista sericea and Plantago argentea P LANT FUNCTIONAL TRAIT COMPOSITION In order to identify dierences in plant trait composition between karst meadows and pastures, the CWM ma trix was analyzed with PCA. e ordination graph is pre sented in Fig. 2, where only traits with signicant dier ences in their CWM values (n = 29, Tab. 2) between karst pastures and meadows are shown. Relevs of both veg Fig. 1: D CA ordination dia gram of matrix with 100 relevs x 180 species. Eigenvalues: Axis 1 = 0.530; Axis 2 = 0.171; 23.8% of variance in species explained by both axes. Shown species have the highest weight. L egend: karst meadows; karst pastures. Abbreviations of species explained in Appen dix 1. Relev numbers corre spond to those in the Appendix 1. FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE ...

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ACTA CARSOLOGICA 40/3 2011 520 Plant functional traits Species abbreviations Karst meadows Karst pastures p Life form LF_He 0.9923 0.9824 *** Life cycle LC_Annl LC_Bien LC_Pere 0.0343 0.0128 0.9528 0.0191 0.0027 0.9782 ** *** *** Growth form GF_Tuss GF_Rose GF_le_st GF_ro_le 0.3088 0.1164 0.4707 0.1003 0.2274 0.1461 0.5062 0.1203 *** * Hairiness Hair_low Hair_hig Hair_0 0.4355 0.2233 0.3411 0.3751 0.1700 0.4548 *** *** *** Plant height Plan_hig 39.23 25.76 *** Specic leaf area SLA 14.55 13.00 *** Leaf dry matter content LDMC 307.57 325.76 *** Flowering start Flo_star 5.5131 5.2079 *** Flowering end Flo_end 7.8051 7.3367 *** Flowering period Flo_peri 3.3085 3.1524 *** Leaf persistence LP_1 0.0115 0.0263 *** Leaf anatomy LA_2 LA_3 0.5001 0.4987 0.6730 0.3248 *** *** Leaf form Lea_for3 Lea_for5 Lea_for8 Lea_for10 Lea_for11 Lea_for12 0.2819 0.1675 0.0456 0.1239 0.0549 0.0399 0.2289 0.2408 0.0663 0.0728 0.0794 0.0258 *** *** ** *** ** C-S-R strategy C S R 0.5193 0.2684 0.2105 0.4360 0.3970 0.1657 *** *** *** < 0.05, ** < 0.01, *** < 0.001 T ab. 2: T raits with signicant diferences in their CWM values between karst meadows (N = 48) and karst pastures (N = 52) (Stu dents t-test). V alues in the table represent the mean of the CWM values of each trait in both groups of relevs. Fig. 2: PCA ordination diagram of CWM matrix with 100 relevs of karst meadows (N = 48) and pastures (N = 52) and plant func tional traits. Only traits (n = 29) with signicant diferences in their CWM values between karst meadows and karst pastures (Students t-test) are shown. Eigenvalues: Axis 1 = 0.86; Axis 2 = 0.14. L egend: karst meadows; karst pastures. Abbrevia tions of plant traits explained in T ab. 1. N ATAA PIPENBAHER, MITJA KALIGARI & SONJA KORNIK

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ACTA CARSOLOGICA 40/3 2011 521 DISCUSSION AND CONCLUSIONS S PECIES RICHNESS AND SPECIES COMPOSITION Calcareous dry grasslands ( Festuco-Brometea ) are known as particularly species-rich habitats (in vascular plants, bryophytes and animals), and they thus host a consider able proportion of Europes biodiversity (W illems 1990). ey are among the most species-rich plant communities in Europe in terms of the number of plant species they support per unit area (W allis DeVries et al. 2002). Our study also found high species richness in both karst meadows and pastures (on average of 34 and 31 species per 25 m 2 respectively), whereas basic compari sons between the two grassland types showed a rather uniform view of species richness. Many studies have ex amined the inuence of dierent management regimes on grassland vegetation (e.g., Diaz et al. 2001; del Pozo et al. 2006; kornik et al. 2010). However, only a few of them directly compare grazing and mowing. Stammel et al. (2003) found that plant species richness on mown straw meadows was signicantly higher than on pastures (in number of species per plot and per site). Hansson & Fogelfors (2000) also detected slightly more species on mown semi-natural grasslands in Sweden, whereas Gutser & Kuhn (1998) found no dierence between mown and grazed grasslands. In contrast, Schlpfer et al. (1998) found signicantly more species on grazed xeric grasslands than on mowed sites. In our case, the slightly higher species diversity of meadows is probably connect ed with their more favorable conditions for plant growth and/or establishment, owing to their more humid and nutrient-rich soil. As expected and already documented in other stud ies (Kaligari 1997; Kaligari & kornik 2007), we found evident dierences in oristic composition between karst meadows and pastures, with a oristic similarity of approx. 54%. Frequent common species, such as Bromus erectus Carex humilis D orycnium germanicum Briza media Salvia pratensis agg., Buphthalmum salicifolium Asperula cynanchica G alium verum Brachypodium ru pestre H ypochoeris maculata Betonica oncinalis subsp. serotina etc., represent plants characteristic of the Euro pean secondary dry and semi-dry grasslands of the phy tosociological class Festuco-Brometea (Dierschke 1997; Kaligari 1997; kornik 2001; Kaligari & kornik 2002; Illys et al. 2007). As DCA ordination of species showed (Fig. 2), dierences in oristic composition between karst meadows and pastures are probably related to soil conditions in the grasslands: soil pH, moisture, available nutrients, etc. Species composition of meadows includ ed many species characteristic of mesotrophic mead ows from the class M olinio-Arrhenatheretea and which were missing in karst pastures: for example Carex acca T ragopogon pratensis L athyrus pratensis and L eontodon hispidus ese species occur in karst meadows because of the more mesic soil conditions (with more humus and moisture and neutral to slightly acid pH). In contrast, karst pastures are developed on shallower, stonier soils, where bedrock reaches the surface and enables the oc currence of many typical oligotrophic, thermophilous, sclerophyllous and basiphilous species, such as Sesleria juncifolia, Carex humilis, Centaurea rupestris, Satureja subspicata subsp. liburnica, Crepis chondrilloides, G enti ana tergestina, G lobularia cordifolia, G enista sericea and Plantago argentea P LANT FUNCTIONAL TRAIT COMPOSITION In addition to dierences in species composition, we also detected several dierences in the plant functional traits between karst meadows and pastures. Our results are etation types are again clearly divided, which indicates that the karst pastures and meadows contain species with distinct plant functional traits. A t-test showed that karst meadows had signicantly higher mean plant height (Plan_hig), SLA values, hemicryptophytes (LF_He), an nuals (LC_annu) and biennials (LC_bien), proportion of hairy plants (Hair_low, Hair_hig), a tussock growth form (GF_tuss) and a grass-like or palmate leaf form (Lea_for3, Lea_for10) (Tab. 2). In karst meadows, spe cies begin to ower later (Flo_star) and have a longer owering period (Flo_peri) than in pastures. In con trast, karst pastures have a higher proportion of peren nials (LC_pere), early owering species (Flo_star), plants with rosette (GF_Rose), spring green and scleromorphic leaves (LP_1, LA_2). Plants in pastures are also char acterized by signicantly higher LDMC values, simple (Lea_for5), pinnatid (Lea_for8) and pinnate leaf forms (Lea_for11) (Tab. 2). e comparison of relative proportions of C, S and R strategists among relevs showed signicantly higher relative proportions of competitors (C) and ruderals (R) in karst meadows than in pastures. Pastures, in contrast had signicantly higher proportions of stress-tolerators (S) when compared to karst meadows (Tab. 2). FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE ...

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ACTA CARSOLOGICA 40/3 2011 522 consistent with previous assertions that grazing gener ally favoured small-stature species and species with ro sette over high, erect and tussock plants (Harper 1969; Lavorel et al. 1997; Diaz et al. 2001; Peco et al. 2005; Diaz et al 2007 ). In pastures, tall, erect plants are eliminated by large herbivore grazing, while small or prostrate species survive (McIntyre et al 1995). Our study showed that forbs with a perennial life cycle predominated in pastures and meadows. Annu als that reproduce generatively and can quickly colonize gaps were rare in both grassland types. e small num bers of species with short life span was probably due to the low intensity of land use. It has been shown in other studies that annuals (therophytes) are promoted by in tensication of land use (Bullock et al. 1994; McIntyre et al. 1995; kornik et al. 2010), as they are more tolerant of disturbance, owing to their fast growth rates and early, prolic seed set (Grime 1974). LDMC and SLA were also traits that dier signi cantly between karst pastures and meadows. Species in pastures showed lower SLA values and higher LDMC values. Lower values of SLA tend to correspond with relatively high investment in leaf defence and long leaf lifespan; whereas leaves with high LDMC tend to be rela tively tough and are thus assumed to be more resistant to physical hazards (e.g. herbivory, wind, hail...) (Cornelis sen et al. 2003). Species with low LDMC and high SLA tend to be associated with productive, oen highly dis turbed environments. Our results partially concur with those of Diaz et al (2001) and W estobys (1998) suggest ing that a higher growth rate (high SLA) is a disturbance tolerance mechanism, whereas small leaves are an avoid ance mechanism (Briske 1996). A similar interpretation could also be used for traits connected to reproductive strategies. In our study, early owering and fruiting ap peared to be promoted in karst pastures. ese traits have been interpreted as an avoidance strategy in rela tion to grazing (Briske 1996). Furthermore, reproductive strategies can be considered as an example of adaptation to both climate and grazing (de Bello et al. 2005). In our case, earlier owering in karst pastures, when compared to the more mesophilous karst meadows, is probably an example of adaptation to more arid microclimate condi tions. Grimes (1974) CSR triangle provides a system for classifying herbaceous plants as having strategies adapt ed to intense competition (C), abiotic stress (S) and fre quent disturbance (R). e grassland communities of the north Adriatic Karst comprise species with pronounced competitive (C) characteristics. Especially in the case of meadows, they appeared to provide sucient resources to support tall, fast growing competitors. In general, the competitive nature of species is restricted by disturbance and stress conditions. In these grasslands, stress is more severe in karst pastures owing to their extremely shallow and stony dry soils, which experience desiccation dur ing the summer. e low relative proportions of ruderals (R) for both karst pastures and meadows also suggest the reduced impact of disturbance, owing to low intensity land-use (mowing and grazing). W e could conclude that karst meadows and pas tures dier signicantly in both oristic composition and functional trait means, owing to dierent environmental conditions (e.g., topography and soil conditions) and dierent land-use (disturbance). DCA analysis of species composition and some of the traits indicated greater re source availability on karst meadows. In contrast, karst pastures showed more species with combination of traits interpreted as an avoidance strategy in relation to distur bance in low productive habitats. REFERENCES ARSO, 2009: Meteoroloki letopis mesena viina pa davin.[Online] Available from: http://www.arso. gov.si/vreme/podnebje/meteorolo%c5%a1ki%20 letopis/2009pad_vis.pdf [Accesses 24 th October 2011]. ARSO, 2009: Meteoroloki letopis mesena viina pa davin.[Online] Available from: http://www.arso. gov.si/vreme/podnebje/meteorolo%c5%a1ki%20 letopis/2009mes_1.pdf [Accesses 24 th October 2011]. Braun-Blanquet, J., 1964: Panzensoziologie. G rundz ge der V egetationskunde .Springer Verlag, pp. 864, W ien. Briske, D.D., 1996: Strategies of plant survival in grazed systems: A functional interpretation.In: Hodgson, J.G. et al. (eds.) e ecology and managment of graz ing systems CAB International, pp. 37, Oxon. Bullock, J.M., Clear Hill, B., Dale, M.P. & J. Silvertown, 1994: An experimental study of the eects of sheep grazing on vegetation change in a species-poor grassland and the role of seedling recruitment into gaps.Journal of applied ecology, 31, 493. N ATAA PIPENBAHER, MITJA KALIGARI & SONJA KORNIK

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ACTA CARSOLOGICA 40/3 2011 523 Cornelissen, J.H.C., Lavorel, S., Gernier, E., Diaz, S., Bu chmann, N., Gurvich, D.E., Reich, P.B., ter Steege, H., Morgan, H.D., van der Heijden, M.G.A., Pausas, J.G. & H. Poorter, 2003: A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany, 51, 335. arni, A. & M. Kaligari, 1992: Study of the succession line from abandoned "terraced" vineyards in NW Istria (Slovenia).In: Colloquio (ed.) Approcci met odologici per la denizione dell`ambiente sico e biologico mediterraneo: seminario scientico-tecnico di L ecce 17 th th November 1992, Castro Marina, 253, Castro Marina. de Bello, F., Lep, J. & M.T. Sebastia, 2005: Predictive val ue of plant traits to grazing along a climatic gradi ent in the Mediterranean.Journal of applied Ecol ogy, 42, 824. del Pozo, A., Ovalle, C., Casado, M., Acosta, B. & J.M. de Miguel, 2006: Eects of grazing intensity in grass lands of the Espinal of central Chile.Journal of Vegetation Science, 17, 791. Diaz, S. & M. Cabido, 2001: Vive la dirence: plant functional diversity matters to ecosystem process es.Trends in Ecology and Evolution, 16, 646. Diaz, S., McIntyre, S., Lavorel, S. & J.G. Pausas, 2002: Does hairiness matter in Harare? Resolving contro versy in global comparisons of plant trait responses to ecosystem disturbance.New Phytologist, 154, 7. Diaz, S., Lavorel, S., de Bello, F., Quetier, F., Grrigulis, K., & M. Robson, 2007: Incorporating plant functional diversity eects in ecosystem service assessments.Proseedings of the National Academy of Sciences of the United States of America, 104, 20684. Dierschke, H., 1994: Panzensoziologie: G rundlage und M ethode .Ulmer, pp. 683, Stuttgart. Dierschke, H., 1997: Panzensoziologisch-synchorolo gische Stellung des Xerothermgraslandes ( FestucoBrometea ) in Mitteleuropa.Phytocoenologia, 27, 2, 127. Ferlan, L. & V. Giacomini, 1955: Appunti tosocialogici su esempi di pascolo carsico Chrysopogono-Cen taureetum cristatae.Atti I Convegno Friul. Sc. Nat. Udine, 4, 159. Garnier, E., Cortez, J., Billes, G., Navas, M.L., Roumet, C., Debussche, M., Laurnet, G., Blanchard, A., Au bry, D., Bellmann, A., Neill, C. & J.P. Toussaint, 2004: Plant functional markers capture ecosystem properties during secondary succession.Ecology, 85, 2630. Goodall, D.W ., 1954: Objective methods for the classi cation of vegetation. II. An essay in the use of factor analysis.Australian Journal of Botany, 2, 304. Grime, J.P., 1974: Vegetation classication by reference to strategies.Nature, 250, 26. Grime, J.P., Hodgson, J.G. & R. Hunt, 1988: Comparative plant ecology .Unwin Hyman Ltd., pp. 752, Lon don. Grime, J.P., 2001: Plant Strategies, V egetation Processes, and Ecosystem Properties .W iley & Sons, pp. 417, Chichester. Gutser, D. & J. Kuhn, 1998: Schafund Ziegenbeweidung ehemaliger Mhder (Buckelwiesen bei Mittenwald): Auswirkungen auf Vegetation und Flora, Empfeh lungen zum Beweidungsmodus.Z. kol. Natursch, 7, 85. Hansson, M., & H. Fogelfors, 2000: Managment of a semi-natural grassland; results from a 15-year-old experiment in southern Sweden.Journal of vegeta tion science, 11, 31. Harper, J.L., 1969: e role of predation in vegetational diversity. Diversity and stability in ecological sys tems.Brookhaven Symposium in Biology, 22, 48 62. Hegi, G., 1958: I llustrierte Flora von M ittel-Europa .Band IV, 1. Teil. Carl Hanser Verlag, pp. 70, Mnchen. Hegi, G. 1963: I llustrierte Flora von M ittel-Europa Band III, 1. Teil. Carl Hanser Verlag, pp. 452, Mnchen. Hegi, G., 1964: I llustrierte Flora von M ittel-Europa Band V, 4. Teil. Paul Parey Verlag, pp. 2255, Berlin Hamburg. Hegi, G., 1965: I llustrierte Flora von M ittel-Europa Band I. Carl Hanser Verlag, pp. 528, Mnchen. Hegi, G., 1966: I llustrierte Flora von M ittel-Europa Band IV, 2. Teil. Carl Hanser Verlag, pp. 448, Mnchen. Hegi, G., 1974: I llustrierte Flora von M ittel-Europa Band VI/3.Teil. Verlag Paul Parey, pp. 366, BerlinHam burg. Hegi, G., 1987: I llustrierte Flora von M ittel-Europa Band VI, 3. Teil. Paul Parey Verlag, pp. 320, BerlinHam burg. Hill, M.O. & H.G. Gauch, Jr., 1980: Detrended Corre spondence Analysis: an improved ordination tech nique.Vegetatio, 42, 47. Hodgson, J.G., W ilson, P.J., Hunt, R., Grime, J.P. & K. ompson, 1999: Allocating C-S-R plant functional types: a so approach to a hard problem.Oikos, 85, 282. Horvat, I. & S. Horvati, 1934: Chrysopogoneto-Sat ureion subspicatae ein neuer Verband der Bro metalia erecti Braun-Blanquet.Acta Botanica inst. Bot. UnivZagrebensis, 4, 8. FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE ...

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ACTA CARSOLOGICA 40/3 2011 525 Piskernik, M., 1988: Suna travia Slovenskega primorja pred 30 leti pp. 21, Lokev-Divaa. Piskernik, M., 1991: G ozdna, travnika in plevelina veg etacija Primorske .Intitut za gozdno in lesno gosp odarstvo, Vtozd za gospodarstvo, VDO Biotehnika fakulteta Univerze v ljubljani, Strokovna in znanst vena dela, pp. 106, Ljubljana. Poldini, L., 1980: bersicht ber die Vegetation des Kar stes von Triest und Grz (NO Italien).Studia GeoStudia Geo botanica, 1, 79. Poldini, L., 1989: L a vegetazione del Carso isontino e tri estino Edizioni LINT, pp. 209, Trieste. Poldini, L. 1991: Atlante corologico delle piante vascolari nel Friuli-V enezia G iulia: inventario oristico region ale Arti grache friulane, pp 897, Udine. Pyek, P. & A. Pyek, 1991: Succession in urban habitats: an analysis of phytosociological data.Preslia, 63, 125. Quzel, P. & F. Medail, 2003: Ecologie et biogeography des forests du basin M editerranean .Elsevier, pp. 573, Paris. Ricotta, C. & M. Moretti, 2010: Assessing the functional turnover of species assemblageswith tailored dis similarity matrices.Oikos, 119, 1089. Rothmaler, W ., 1995: Exkursionsora von D eutschland .Gustav Fischer Verlag, pp. 753, JenaStuttgart. Schlapfer, M., Zoller, H. & C. Korner, 1998: Inuences of mowing and grazing on plant species composition in calcareous grassland.Botanica Helvetica, 108, 57. SPSS Inc., 2006: SPSS Base 15.00 User`s guite. Chicago, IL: SPSS Inc. Stammel, B., Kiehl, K. & J. Pfadenhauer, 2003: Alterna tive management on fens: Response of vegetation to grazing and mowing.Applied Vegetation science, 6, 245. kornik, S., 2001: A contribution to the knowledge of dry grassland vegetation of the Brometalia erecti Koch 1926 order in Slovenia = Prispevek k pozna vanju vegetacije suhih travi reda Brometalia erecti Koch 1926 v Sloveniji.Acta Biologica Slovenica, 44, 29. kornik, S., Vidrih, M & M. Kaligari, 2010: e eect of grazing pressure on species richness, composition and productivity in North Adriatic Karst pastures.Plant Biosystem, 144, 355. ter Braak, C.J.F. & P. milauer, 2002: CANOCO Refer ence Manual and CanoDraw for W indows User's Guide: Soware for Canonical Community Ordina tion (version 4.5). Microcomputer Power, pp. 500, Ithaca NY USA. W allis DeVries, M.F., Poschlod, P. & J.H. W illems, 2002: Challenges for the conservation of calcareous grass lands in Northwestern Europe: integrating the re quirements of ora and fauna.Biological Conser vation, 104, 265. W atts, D., 2004: Quaternary biotic interactions in Slove nia and adjacent regions: the vegetation. In: Grif ths, H.I.B. et al. (eds) Balkan Biodiversity Pattern and process in the European hotspots Kluwer, pp. 69, Dordrecht. W estho, V. & E. van der Maarel, 1973: e Braun-Blan quet approach.Handbook of Vegetable Science, 5, 156. W estoby, M., 1998: A leaf-height-seed (LHS) plant ecol ogy strategy scheme.Plant soil, 199, 213. W illems, J.H., 1990: Calcareous grasslands in continental Europe. In: Hiller, S.H. et al. (eds.) Calcareous grass lands: ecology and management Bluntisham books, pp. 3, FLORISTIC AND FUNCTIONAL COMPARISION OF KARST PASTURES AND KARST MEADO W S FROM THE ...



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ECOLOGICAL STUDIES OF AN EPIKARST COMMUNIT Y IN SNE NA JAMA NA PLANINI ARTO AN ICE CAVE IN NORTH CENTRAL SLOVENIA EKOLOKE RAZISKAVE EPIKRAKE ZDRUBE V SNENI JAMI NA PLANINI ARTO LEDENI JAMI V SEVERNI OSREDNJI SLOVENIJI Federica PAPI 1 & Tanja PIPAN 2 Izvleek UDK 574.1:551.444.6(497.4) Federica Papi & Tanja Pipan: Ekoloke raziskave epikrake z drube v Sneni jami na planini Arto ledeni jami v severni osrednji Sloveniji V ledeni jami Snena jama na planini Arto na Raduhi smo opravili prvo raziskavo epikrake biodiverzitete v visokogor skih jamah v povezavi z okoljskimi dejavniki. V tej visokogorski jami smo eno leto vzorevali favno na petih mestih. V prenikli vodi smo merili temperaturo, prevodnost, pretok, pH, celokupno trdoto in koncentracije razlinih ionov (kalcij, kalij, nitrat, sulfat in fosfat). V vhodnem delu jame je led prisoten vse leto. Temperatura v jami ne presee 4 C. Te razmere se odraajo v favni, najdeni v prenikli vodi. Prvo vzorno mesto je bilo ves as zaledenelo in brez favne, brez ivali je bilo tudi vzorno mesto z jamskim mlekom na stropu. Na drugih treh vzornih mestih so bili najdeni osebki desetih nevretenarskih taksonov, med katerimi so bili najtevilneji kopepodni rakci. Njihova abundanca je bila v pozitivni korelaciji s pretokom, pH in tem peraturo ter v curku z najtevilnejo favno tudi s prevodnostjo in celokupno trdoto. Velik dele mladih kopepodov v vzorcih nakazuje, da je epikras njihov primarni (to je izvorni) habitat. Raziskave visokogorske epikrake favne lahko pripomo rejo k celovitejemu razumevanju ekologije epikrake favne in njene vloge v razlinih plitvih podzemeljskih habitatih. Kljune besede : epikras, ledena jama, prenikajoa voda, spe leobiologija, favna. 1 University of Nova Gorica, Vipavska cesta 13, SI-5000 Nova Gorica, Slovenia, e-mail: federicapapi@hotmail.com 2 Karst Research Institute ZRC SAZU, Titov trg 2, SI-6230, Postojna, Slovenia, e-mail: pipan@zrc-sazu.si Received/Prejeto: 15.5.2011 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 505, POSTOJNA 2011 Abstract UDC 574.1:551.444.6(497.4) Federica Papi & Tanja Pipan: Ecological studies of an epikarst community in Snena jama na planini Arto an ice cave in north central Slovenia Epikarst biodiversity in relation to environmental conditions was studied for the rst time in an ice cave, Snena jama na planini Arto on Mt. Raduha in north central Slovenia. In this alpine cave ve sampling sites were monitored for fauna in per colation water in the period of one year. Temperature, conduc tivity, discharge, pH, total hardness and concentrations of vari ous ions (calcium, chloride, nitrite, sulphate and phosphate) of water were measured. At the entrance of the cave ice is pres ent all year round and the temperature inside the cave rises to a maximum of 4 C. ese circumstances are reected in the fauna found in percolating water. e rst sampling site in the permanent ice was without fauna, as did the sampling site in an area with moonmilk. In the other three sampling sites, in dividuals of ten invertebrate taxa were found, Copepoda being the most abundant. eir abundance was positively correlated with discharge, pH and temperature of percolation water and additionally in the most abundant drip also with conductivity and total hardness. High proportion of immature copepods in drips shows that epikarst is their primary (i.e. source) habitat. Investigations of the alpine epikarst fauna can help to under stand better the ecology of the epikarst fauna and its roles with in the large range of dierent shallow subterranean habitats. Keywords : epikarst, ice cave, percolating water, speleobiology, fauna.

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ACTA CARSOLOGICA 40/3 2011 506 During the Pleistocene period, the Slovene Alps were covered by permanent snow and ice. In the coldest period of the W rmian, the permanent snow line was between 1300 m above the present sea level and long valley glaciers descended down to 500 m (Kranjc 1984). Systematic investigation of the epikarst fauna (fau na from unsaturated zone including deep soil root zone) in Slovenian Alpine or Pre-Alpine caves has not yet been done. ere are some sporadic reports on the terrestrial fauna from caves in the Slovenian Alps (e.g., Novak & Kutor 1983; Novak et al 2004; Slapnik 2001), but none on the aquatic cave fauna. Epikarst is the uppermost layer of karstic bedrock. It is typically 3 m deep, but its characteristics can vary considerably (Ford & W illiams 2007). From an eco logical point of view, epikarst is both an exceptionally di verse and environmentally heterogeneous habitat (Cul ver & Pipan 2009). Environmental conditions in epikarst are more variable than in caves, and organic carbon dripping from epikarst is an important source of carbon in many caves. e fauna is both diverse and special ized, with Copepoda and, to a lesser extent, Insecta and Amphipoda dominating (Pipan 2005; Pipan et al. 2006). e epikarst aquatic fauna is by far the best known from Dinaric karst in Slovenia (Pipan 2003, 2005) and from patches of isolated karst in Slovenia (Pipan et al 2008). It has also been studied in other countries, including Spain, Romania, and USA (e.g., Camacho et al. 2006; Moldovan et al. 2007; Pipan & Culver 2005). In Slovenia, karst areas comprise 43% of the state (Knez & Kranjc 2009), of which about one third (17%) belongs to the alpine karst (Habi 1969). Precipita tion in the alpine karst varies between 1500 to more than 3000 mm annually and snow cover lasts up to 200 days (Kunaver 1983). Present permanent snow line is at 2700 m a.s.l. (Kunaver 1983). Summer mean tempera ture in the Alps is about 11 C, while winter mean tem perature is about -1.6 C (Gams 1960). e Slovenian calcareous Alps are 100 km long (from west to east) and 40 km wide (from north to south) (Kranjc 1984). Dominant rocks are Triassic limestones and dolomites, up to1,000 m thick, and the highest peak is 2,864 m in the Julian Alps (Audra et al. 2007). e Slo venian Alps consist of two large formations: the Central Alps (Karavanke) and the Southern Limestone Alps (Ju lian and KamnikSavinja Alps). In the Limestone Alps the great majority of the terrain is built of carbonate rocks and there are a few patches of other rocks. In the Karavanke it is just the opposite most of the terrain is impermeable rocks and among them there are some car bonate rocks, nearly all of Mesozoic age (Kranjc 1984). In this contribution, we deal with the drip fauna of the ice cave Snena jama na planini Arto, in the South ern Limestone Julian Alps, where we intensively studied both the copepod and non-copepod fauna from a series of ve epikarst drips. W e hypothesized that an alpine epikarst fauna consists mostly of troglobiotic species, but is less diverse and abundant than the epikarst fauna from the Dinaric karst. W e ask: (1) Is copepod abundance and diversity correlated with dierent physical and chemical parameters measured in percolation water? (2) W hat is a proportion of immature copepod individuals relative to the number of adults as a measure of reproduction? (3) W hat environmental parameters are correlated with the presence of reproducing copepods? INTRODUCTION RESEARCH AREA e investigations were carried out in the tourist cave Snena jama na planini Arto (Cadastre Number 1254, Cave cadastre IZRK ZRC SAZU and Speleological Asso ciation of Slovenia) in the north central part of Slovenia (Fig. 1). e cave is situated on Mt. Raduha in the north eastern part of the KamnikSavinja Alps (Zupan Hajna et al 2008). Snena jama which means Snow cave is the largest horizontal cave (1,327 m) in Mt. Raduha. e cave is developed in the Upper Triassic limestones (Zupan Hajna et al 2008). e ocial entrance is at the one of four large shas, opening at 1,556 m a.s.l. and leading into the rst cave sectiona large gallery. Permanent ice is formed at the entrance part of the cave due to cold air which ows through the shas into the cave (Zupan Hajna et al 2008). Maximum temperature inside the cave is 4 C. In the inner part of the cave there are owstone deposits and moonmilk (Fig. 2). F EDERICA PAPI & T ANJA PIPAN

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ACTA CARSOLOGICA 40/3 2011 507 ECOLOGICAL STUDIES OF AN EPIKARST COMMUNIT Y IN SNE NA JAMA NA PLANINI ARTO AN ICE CAVE IN ... Fig. 2: Calcite moonmilk (white deposit on the ceiling) at the sampling site SJ4 and the sampling device for collecting percola tion water. Fig. 1: G eographical location of the cave Snena jama na planini Arto. MATERIAL AND METHODS Sampling was done using the continuous sampling de vice for collecting epikarst fauna developed by Pipan (2003, 2005). Five such devices were placed under drip ping water in the cave (Fig. 3). e rst of these (SJ1) was 50 m from the entrance, close to the ice-pillar. is sampling site was more or less covered by ice during the whole sampling period (Fig. 4). Another device (SJ2) was set 100 m from the entrance; in January it was covered by ice, too. e third sampling site (SJ3) was on the edge of the ice limit in the cave, 270 m from the entrance. Two collecting devices were positioned in the inner part of the cave: one under the moonmilk, 440 m from the entrance (SJ4), and the other on a stalagmite, close to the moon milk formation, 510 m from the entrance (SJ5) (Fig. 2). Sampling devices were set in September (except SJ3 which was set in December) 2006 and were moni tored for fauna on seven visits from October 2006 till September 2007. e samples of fauna were in situ xed and stored in 70% ethanol. Organisms were sorted in the laboratory of the Karst Research Institute Scientic Re search Centre of the Slovenian Academy of Sciences and Arts (IZRK ZRC SAZU) in Postojna, using a microscope (Nikon Eclipse 600) and identied in the laboratory of the Natural History Museum of Trieste, using micro scopes Leica MZ 16 and Leica DMLB. Temperature (C), conductivity (S/cm) and pH of percolation water were measured in situ using a con ductivity meter (LF 91, W T W ), pH meter (323, W T W ), and a Combo multi-parameter Hanna Instruments. Measurements were performed during each sampling of fauna between October 2006 and September 2007, with additional measurements in 2010/2011. Drip rates were measured by collecting water under drips in a graduated cylinder for timed intervals. Concentrations of cations

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ACTA CARSOLOGICA 40/3 2011 508 (calcium) and anions (chlo ride, nitrite, sulphate and phosphate) as well as total hardness were determined periodically according to Standard Methods for the Examination of W ater and W astewater (1989). Measure ments at the sampling points SJ1 and SJ2 were not always possible because of periodi cal ice covering of the sites without liquid water (Fig. 4). Descriptive statistics was used to describe physical and chemical characteristics of the drips. Pearson cor relation coecients ( r ) were calculated to evaluate copepod abundances with respect to chemical and physical parameters. Statistical analysis and signicance of environmental variables was tested using PAST (Paleontological Statistics Soware Package for Education and Data Analysis, version 2.10) (Ham mer et al. 2001). Fig. 3: M ap of the cave Snena jama na planini Arto, with marked locations of sampling sites (modied from Z upan H ajna et al. 2008, used with permission). Fig 4: Sampling site SJ1 covered by ice. RESULTS Characteristics of the measured physical and chemi cal parameters of dripping water in the cave are shown in Tab. 1. Among these, temperature and pH were the most stable parameters. In both cases, low standard de viation indicates that pH and temperature values tend to be close to the mean and did not vary among drips or during the sampling period. Temperature of the water among sites varied from 0.0 C (frozen water at SJ1 in October) to 4.7 C (at SJ5 in December). More variable was discharge, with data spread over a large range of values (e.g., between 0 ml/min and 880 ml/min, meas ured in December 2006 at two drips). Dry periods with low discharge were in summer, reecting no or low amount of precipitation, and in winter because of the snow cover. A total of 189 copepods, including nauplia, belong ing to two species ( Speocyclops infernus and Bryocamptus sp.) were collected (Tab. 2). Most cyclopoids ( Speocyclops infernus ) were recorded at the beginning of summer (in June), while harpacticoids ( Bryocamptus sp.) were most abundant in June and in September (Fig. 5). Both species showed some modications to subterranean life, includ ing depigmentation, reduction of eyes, and appendages elongation. Copepods were found in three of ve drips, in 17 of 28 samples (Tab. 3). In addition to copepods, 28 other invertebrates, representing at least eight species were found. Copepods were the most abundant animal group, representing 87% of all collected animals, and about 90% of all troglomorphic animals; beside copep ods, these were Amphipoda, Hydracarina and Collem F EDERICA PAPI & T ANJA PIPAN

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ACTA CARSOLOGICA 40/3 2011 509 T ab. 1: Ranges of physical and chemical measurements of ve drips in Snena jama na planini Arto in 2006 and 2010. D rips with copepods are italicized. Parameter / Drip MeanSD (minmax) SJ1 SJ2 SJ3 SJ4 SJ5 Temperature (C) 0.1.2 1.1.8 1.7.5 3.1.3 4.2.3 (-0.3.3) (0.0.0) (0.9.4) (3.1.2) (3.6.7) Conductivity (S/cm) 311.0.6 307.0.4 260.3.9 321.7.8 294.1.1 (306.0.0) (243.0.0) (223.0.0) (309.0.0) (266.0.0) pH 8.41.33 8.35.29 8.23.32 8.36.25 8.19.23 (8.17.79) (7.93.78) (7.73.58) (7.97.82) (7.97.82) Discharge (ml/min) 4.5.4 3.4.0 81.6.7 18.8.6 287.0.7 (0.0.8) (0.0.4) (12.4.0) (5.2.0) (32.0.0) Total hardness (mg/L) 178.5.9 164.5.4 144.0.4 174.0.4 162.5 16.4 (171.5.5) (174) (118.0.5) (187.0.5) (140.0.0) Ca hardness (mg/L) 62.12.36 38.65.33 54.66.89 55.60.14 55.63.27 (56.92.32) (25.69.61) (47.33.38) (53.04.18) (52.43.55) Cl (mg/L) 2.0.0 2.5.7 2.12.63 1.87.25 2.12.25 (2.0.0) (2.0.0) (1.5.0) (1.5.0) (2.0.5) NO 3 (mg/L) 2.54.37 2.54.96 1.91.34 2.88.68 2.43.52 (2.27.80) (1.86.22) (1.51.19) (2.41.88) (1.94.12) SO 4 2 (mg/L) 4.02.19 1.38.15 1.98.11 3.05.19 3.72.50 (2.48.57) (0.57.19) (0.44.04) (0.07.93) (2.04.66) PO 4 3 (mg/L) 0.009.001 0.010.008 0.006.011 0.076.125 0.011.012 (0.009.010) (0.004.016) (0.000.023) (0.000.220) (0.000.028) Carbonate (meq/L) 3.34.0 2.73.0 2.64.26 3.27.22 2.96.15 (3.34.34) (2.73.73) (2.46.83) (3.12.43) (2.86.07) T ab. 2: L ist of taxa, their abundance and level of specialization to subterranean environment, collected from three drips in Snena jama na planini Arto in 2006. Phylum / Subphylum Class / Sublass Order Family Genus Troglomorphic N Sampling site Annelida Oligochaeta No 2 SJ5 Arthropoda / Chelicerata Arachnida / Micrura Acarina: Hydracarina Yes 14 SJ2, SJ3, SJ5 Arthropoda / Crustacea Maxillopoda / Copepoda Cyclopoida Cyclopidae Speocyclops infernus Yes 69 SJ2, SJ3, SJ5 Harpacticoida Canthocamptidae Bryocamptus sp. Yes 112 SJ2, SJ3, SJ5 nauplia 8 SJ2, SJ3, SJ5 Malacostraca / Eumalacostraca Amphipoda Niphargidae Niphargus sp. Yes 3 SJ5 Arthropoda / Hexapoda Entognatha Collembola Isotomidae Yes 3 SJ3, SJ5 Insecta Coleoptera cf. Lathriidae No 1 SJ3 Diptera No 1 SJ5 Psocoptera No 3 SJ2, SJ3 Thysanoptera No 1 SJ3 ECOLOGICAL STUDIES OF AN EPIKARST COMMUNIT Y IN SNE NA JAMA NA PLANINI ARTO AN ICE CAVE IN ...

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ACTA CARSOLOGICA 40/3 2011 510 bola, the last taxon being terrestrial. Amphipods were found at only one site: SJ5, with the highest temperature and discharge measured. In the sites SJ1 and SJ4 no ani mals were found (Tab. 3). Overall, the average rate of copepods was 0.2 per drip per day (Tab. 3). ere were dierences in the phys ico-chemical parameters between copepod and noncopepod drips (Tab. 1). Copepods were found nearly throughout the entire range of pH, discharge and chemi cal qualities of the drips measured, but not in the drips with high conductivity (over 300 S/cm; SJ4) and high values of carbonate (over 3.3 meq/L; SJ4), and, of course, in frozen drips. e highest abundance of Copepoda and the other two troglomorphic taxa (Hydracarina and Col lembola) were found at the sampling site SJ3 with rela tively low temperatures (up to 2.4 C), low conductivity (up to 302 S/cm) and low values of carbonate ions (up to 2.8 meq/L) in comparison with the other sites. Among the ten variables measured, temperature, conductiv ity, and total hardness best explained the variation of copepod abundance at the sampling site SJ3. Statisti cal signicant correlations (p < 0.01) were found be tween these environmental parameters and the copepod abundance. In general, Pearson cor relation coecients show signicant correlation be tween copepod abundance and both discharge and pH (r = 0.90, p < 0.05 in both cases). ere was also sig nicant correlation between number of copepod indi viduals and temperature (r = 0.75, p < 0.05). W e found both mature and immature individuals, rst represented by males with mature spermatophore and females with attached spermatophore, as well as many juveniles (38% of the to tal copepod abundance) and nauplia (4% of the total co pepod abundance). W e did not nd any pre-copulating pairs and only one ovigerous female of Speocyclops infer nus e abundance of mature individuals (males and fe males with mature attached spermatophores) was highly correlated with temperature (r = 0.8, p < 0.05). Fig. 5: Abundance of two copepod species, Speocyclops infernus and Bryocamptus sp. from perco lation water in Snena jama na planini Arto in 2006. T ab. 3: Copepod abundance and their ux per day in ve drips in Snena jama na planini Arto (/ not sampled; sampling impossible due to the ice). Date SJ1 SJ2 SJ3 SJ4 SJ5 27/10/2006 0 0 0 13 1 0 24 7 24 / / 2 18 12 17 25 0 0 0 0 0 0 0 5 1 8 8 11 4 9 08/12/2006 06/01/2007 28/04/2007 14/06/2007 05/08/2007 23/09/2007 Total 0 69 (per day=0.24) 74 (per day=0.26) 0 46 (per day=0.18) DISCUSSION Snena jama na planini Arto is an alpine ice cave, where the alpine climate could limit the distribution of subter ranean fauna. Aquatic species (Copepoda, Amphipoda, and Hydracarina) from epikarst water show troglo morphic characters and are presumably troglobionts sensu stricto (Pipan 2005; Pipan et al 2008; Culver et al F EDERICA PAPI & T ANJA PIPAN

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ACTA CARSOLOGICA 40/3 2011 511 2010). Perhaps individuals of Psocoptera and Diptera are accidentals and ew into the traps from the cave, while the other terrestrial taxa (Oligochaeta, Coleop tera and Collembola) might be specialists of the soil root zone (T. Novak, per. comm.). Terrestrial species that are presumably from the root layer also dominated in percolation water in a cave from isolated karst in north eastern Slovenia (Pipan et al. 2008), but not in caves in the Dinaric karst of Slovenia (Pipan 2005). It is not surprising that aquatic and terrestrial species show dierent patterns of distribution and abundance with respect to karst region (i.e., Dinaric, alpine, and isolat ed). e factors driving the groups to colonize subterra nean habitats are dierent in the two cases. e Messin ian salinity crisis, aecting primarily the Dinaric karst, was a major factor in the colonization of subterranean habitats by aquatic species (Culver & Pipan 2010), while drying conditions during the Pleistocene were a major factor in the colonization of subterranean habitats by terrestrial species, especially in the northern karst re gions (Barr 1968). Nevertheless, the ux of copepods measured as copepod abundance per drip per day was lower in six caves in Dinaric karst (0.1) than in Snena jama na planini Arto (0.2). is dierence is negligible in comparison with data from Organ Cave, W est Vir ginia, USA (Pipan et al 2006), where an average of one copepod per drip per day was found. It is possible that Dinaric epikarst is less conducting, where the residence time of water may be much greater than epikarst in the other two cases, based on residence time estimates of Kogovek (2010). Aquatic as well as terrestrial invertebrates in per colation water are important and sometimes, together with dissolved and ne particulate organic matter, the only source of organic nutrients for organisms in deeper subsurface habitats, especially so in areas of barren sur face landscape such as high mountain karst. Simon et al. (2003, 2010) showed that the dissolved organic carbon in percolating water was an important carbon source and an important source of epilithic biolms in caves. In all ve sampled drips a large variation with re spect to presence/absence and abundance of copepods was detected as a considerable spatial variation in both physical and chemical parameters. At the most abundant drip, SJ3, three environmental parameters: temperature, conductivity and total hardness of percolation water were found to aect the presence of copepods. In gener al, values of these parameters were lower in comparison to other drips. Copepods preferred lower temperatures and lower conductivity values, which correspond to lower concentration of calcium and magnesium ions. As the conductivity might be interpreted as a surrogate for the duration of water retention in underground habitats, these results suggest that SJ3 drip is quickly transmitted through the vadose zone. Such circumstances, in gener al, appear at thinner cave ceilings and/or more fractured bedrock above the ceiling. Y ield of copepods from drips was clearly connect ed with the ow rate: greater the ow is, greater is the abundance. e presence of amphipods in SJ5 can also be understood as the consequence of a greater ow. is is in contrast to previous investigations in Organ Cave, USA (Pipan et al 2006), where the highest density of copepods was found at low ow rates. But drip rates in Snena jama na planini Arto rarely reached 100 ml/min, which was the case in Organ Cave. Upon the data from Snena jama na planini Arto we can conclude that both, the higher and the slower the ow rates are, the less suit able the epikarst habitat may be for copepods and other microinvertebrates. is could be generally true in the alpine epikarst. In the epikarst above Snena jama na planini Arto the optimum conditions appeared in late summer when the highest overall copepod abundance, and the highest abundance of mature individuals were observed in the cave. Accordingly, the abundance of mature individuals was positively correlated with temperature. e absence of fauna in two drips may be explained in two dierent ways. W hile SJ1 drip seems sterile because of low water temperature, or possibility of existence deep ssures but with less storage, it is possible that the absence in SJ5 is the consequence of a thick moonmilk deposition which acts as a lter prevailing fauna from falling into the sam pling device. e other reason might be some substance noxious to fauna, such as those known to inhibit mi crobial activity in the moonmilk deposits (Janices et al 2009). Moonmilk is a secondary cave deposit, common ly associated with biogenic calcite precipitation (Hill & Forti 1997; W alochnik & Mulec 2009). e high proportion (38%) of immature individu als of copepods is in accordance with data obtained from six caves in the Dinaric karst (Pipan et al 2010). In both cases, the ratios of immature individuals show that reproduction in situ is high and that the epikarst harbors this troglobiotic fauna. Animals found in drips represent a biased sample of individuals washed out from the substrates in the epikarst where they lived. Anyway, alpine epikarst, where climatic conditions are much more severe than those in lower karst landscapes, is also relatively abundantly colonized with subterra nean fauna, like the Dinaric epikarst. Investigations of the alpine epikarst fauna, inhabiting shallow subterra nean habitats, might facilitate the understanding of the ecology of the epikarst faunas and their roles within the large range of thermally and otherwise dierent shallow subsurface habitats. ECOLOGICAL STUDIES OF AN EPIKARST COMMUNIT Y IN SNE NA JAMA NA PLANINI ARTO AN ICE CAVE IN ...

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ACTA CARSOLOGICA 40/3 2011 512 W e are indebted to David C. Culver and Tone Novak for the revision of the manuscript, many useful suggestions and discussions. David Culver and Trevor Shaw im proved the language. is study was partly supported by the Slovenian Research Agency, and Ad-futura: Slovene Human Resources Development and Scholarship Fund. Laboratory work was performed at the Karst Research Institute ZRC SAZU and in the entomology laboratory of the Natural History Museum of Trieste. W e thank Mateja Zadel for chemical analysis and dr. Andrea Colla for helping at insect determinations, useful suggestions and technical support. FP also thanks the members of Jamarsko Drutvo rni galeb Prebold (especially Darko Naraglav), cavers of Jamarsko Drutvo Seana (especially Jure Jakovi, Zdenka, Rok and Irena itko, Andrej, Luka and Silvia Peca), researchers of the Karst Research Institute ZRC SAZU (especially Janez Mulec and Trevor Shaw), and other friends (Chiara Focaccetti, Fabio Polese, Andreea Oarga, Brane Koren and Brigitta Slavec). ACKNO WLEDGEMENTS REFERENCES Audra, P., Bini, A., Gabrovek, F., Huselmann, F., Hobla, F., Jeannin, P.Y ., Kunaver, J., Monbaron, M., uteri, F., Tognini, P., Trimmel, H. & A. W ild berger, 2007: Cave and karst evolution in the Alps and their relation to paleoclimate and paleotopog raphy.Acta Carsologica, 36, 1, 53. Barr, T.C., 1968: Cave ecology and the evolution of tro globites.Evolutionary Biology 2, 35. Camacho, A.I., Valdecasas, A.G., Rodrguez, J., Cuezva, S., Lario, J. & S. Snchez-Moral, 2006: Habitat con straints in epikarstic waters of an Iberian Peninsula cave system.Annales de Limnologie-International Journal of Limnology, 42, 2, 127. Culver, D.C. & T. Pipan, 2009: e Biology of Caves and Other Subterranean H abitats .Oxford University Press, pp. 254, Oxford, U.K. Culver, D.C. & T. Pipan, 2010: Climate, abiotic factors, and the evolution of subterranean life.Acta Carso logica, 39, 3, 577. Culver, D.C., Holsinger, J.R., Christman, M.C. & T. Pipan, 2010: Morphological dierences among eye less amphipods in the genus Stygobromus dwelling in dierent subterranean habitats.Journal of Crus tacean Biology, 30, 68. Ford, D. & P. W illiams, 2007: K arst hydrogeology and geomorphology .John W iley & Sons, pp. 562, New Y ork. Gams, I., 1960: On the extreme upper limit of rural set tlements, winter wheat, forest and snow line in Slo venia.Geografski vestnik, 32, 59. Habi, P., 1969: Hidrografska rajonizacija krasa v Slo veniji.Kr Jugoslavije, 6, 79. Hammer, ., Harper, D.A.T. & P.D. Ryan, 2001: PAST: Paleontological Statistics Soware Package for Edu cation and Data Analysis.Paleontologia Electronica, 4, 1, 1. Hill, C. & P. Forti, 1997: Cave minerals of the world .Na tional speleological society, pp. 463, Huntsville. Janices, I., Portillo, M.C., Cuezva, S., Gonzales, J.M., Caaveras, J.C. & S. Sanchez-Moral, 2009: Micro bially induced calcitic moonmilk deposits lead to inhibition of microbial activity in caves.15 th In ternational Congress of Speleology, K errville, T exas, United States of America, July 19, 2009 pp. 378, Kerrville, Texas. Knez, M. & A. Kranjc, 2009: Kras.In: Pleniar, M., Og orelec, B. & M. Novak (eds.), e G eology of Slo venia Geoloki zavod Slovenije, pp. 561, Lju bljana. Kogovek, J., 2010: Characteristics of percolation through the karst vadose zone .Zaloba ZRC/ZRC Publish ing, pp. 168, PostojnaLjubljana. Kranjc, A.A., 1984: Speleological characteristics of alpine karst in Slovenia, northwestern Y ugoslavia.Norsk Geogrask Tidsskri, 38, 3, 177-183. Kunaver, J., 1983: Geomorpholy of the Kanin mountains with special regard to the Glaciokarst.Geografski zbornik, 22, 201. Moldovan, O.T., Pipan, T., Iepure, S., Mihevc, A. & J. Mulec, 2007: Biodiversity and ecology of fauna in percolating water in selected Slovenian and Roma nian caves.Acta Carsologica, 36, 493. Novak, T. & V. Kutor, 1983: e ecology of an Alpine ice cave.Mmoires de Biospologie, 10, 117. F EDERICA PAPI & T ANJA PIPAN

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ACTA CARSOLOGICA 40/3 2011 513 Novak, T., Lipovek Delakorda, S., Seni, L., Pabst, M.A., & F. Janekovi, 2004: Adaptations in pha langiid harvestmen G yas annulatus and G. titanus to their preferred water current adjacent habitats.Acta Oecologica, 26, 45. Pipan, T., 2003: Ekologija cepononih rakov (Crustacea: Copepoda) v prenikajoi vodi izbranih krakih jam .PhD thesis (in Slovene with English abstract and summary), Univerza v Ljubljani, Oddelek za bi ologijo, pp. 130, Ljubljana. Pipan, T., 2005: Epikarst A Promising H abitat. Copepod fauna, its diversity and ecology: a case study from Slovenia (Europe) .Karst Research Institute at ZRC SAZU/ZRC Publishing, pp. 101, Postojna. Pipan, T. & D.C. Culver, 2005: Estimating biodiversity in the epikarstic zone of a W est Virginia cave.Journal of Cave and Karst Studies, 67, 2, 103. Pipan, T., Christman, M.C. & D.C. Culver, 2006: Dy namics of epikarst communities: microgeographic pattern and environmental determinants of epikarst copepods in Organ Cave, W est Virginia.American Midland Naturalist, 156, 75 Pipan, T., Navodnik, V., Janekovi, F. & T. Novak, 2008: Studies of the fauna of percolation water of Huda luknja, a cave in isolated karst in northeast Slove nia.Acta Carsologica, 37, 1, 141. Pipan, T. Holt, N. & D.C. Culver, 2010: How to protect a diverse, poorly known, inaccessible fauna: identi cation and protection of source and sink habitats in the epikarst.Aquatic Conservation: Marine and Freshwater Ecosystems, 20, 748. Simon, K.S., Beneld, E.F. & S.A. Macko, 2003: Food web structure and the role of epilithic biolms in cave streams.Ecology, 84, 2395. Simon, K.S., Pipan, T., Ohno, T. & D.C. Culver, 2010: Spatial and temporal patterns in abundance and character of dissolved organic matter in two karst aquifers.Fundamental and Applied Limnology, 177, 81. Slapnik, R., 2001: Activity and movements of Zospeum isselianum Pollonera 1886 (Gastropoda, Pulmonata, Carychiidae) in a cave in the KamnikeSavinjske Alps (Slovenia).Natura Croatica, 10, 153. Standard M ethods for the Examination of Water and Wastewater 17 th edition, 1989.APHA-A WW AWPCF, pp. 1088. W alochnik, J. & J. Mulec, 2009: Free-living amoebae in carbonate precipitating microhabitats of karst caves and a new vahlkampid amoeba, Allovahlkampa spelaea gen. nov., sp. nov.Acta Protozoologica, 48, 25. Zupan Hajna, N., Mihevc, A., Pruner, P. & P. Bosk, 2008: Paleomagnetism and magnetostratigraphy of karst sediments in Slovenia .Zaloba ZRC/ZRC Publish ing, pp. 266, Ljubljana. ECOLOGICAL STUDIES OF AN EPIKARST COMMUNIT Y IN SNE NA JAMA NA PLANINI ARTO AN ICE CAVE IN ...



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TIN MINING IN THE LIMESTONE CAVES OF PERLIS, MALA Y SIA R UDARJENJE KOSITRA V KRAKIH JAMAH P ERLISA, MALEZIJA Liz PRICE 1 Izvleek UDK 551.435.84:622.345(595) Liz Price: Rudarjenje kositra v krakih jamah Perlisa, Malezija Perlis je majhna drava na severu Malezije. Ima tako obmoja izoliranega stolpastega krasa, kot obmoja kopastega krasa, kjer v tevilnih krakih jamah najdemo aluvijalne sedimente, bogate s kositrom. Rudarjenje v teh jamah je potekalo v 20. sto letju in je po obsegu presegalo primerljiva rudarjenja v krakih jamah. Kljune besede: Perlis, polotoki del Malezije, kositer, ruda v aluviju, rudarjenje, jama. 1 B-12-3, Le Chateau 2, Lorong Syed Putra Kiri, 50460 Kuala Lumpur, Malaysia, e-mail: lizprice@hotmail.com Received/Prejeto: 14.10.2010 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 497, POSTOJNA 2011 Abstract UDC 551.435.84:622.345(595) Liz Price: Tin mining in the limestone caves of Perlis, Malay sia Perlis is a small state in northern Malaysia, and it has both iso lated tower karst hills as well as a long range of limestone hills. Many of the natural caves within this hill range contain alluvial tin deposits that were extensively mined during the 20 th cen tury on a scale that is unique for tin ores from caves. Keywords: Perlis, Peninsular Malaysia, tin, alluvial ore, min ing, cave. I NTRODUCTION Perlis is the smallest state in Malaysia, and also the north ernmost, situated in the northwest corner of the peninsu la, with the border to ailand wrapped round its north ern margin. It has the southernmost limestone range on the mainland of Southeast Asia, the remainder of the limestone within Peninsular Malaysia occurs as isolated outcrops and towers. e Setul Boundary Range is rid dled with caves, many of which have active streams that are rarely found elsewhere in the peninsula. is lime stone range is unique because it is the longest such range in the peninsula, it is formed of the oldest limestone in the country and the natural caves have been mined ex tensively for the tin minerals within their alluvial sedi ments. Nowhere else in the world has tin been mined to such an extent from deep within limestone caves. ere are two groups of limestone hills within Perlis (Fig. 1). e Setul Hills form a continuous belt of mountainous country along the western border of the state (Fig. 1). ey form steep polygonal karst, with fewer clis, more subdued slopes and more forest cov er (Fig. 3). e highest peak of the Setul range is Bukit Pelarit at 553 m. e Setul Hills contain the caves with the tin deposits, mainly in the area around Kaki Bikit. In contrast, in a north-south belt through the centre of the state, the Chuping Hills are formed of eroded frag ments of a strong, white, Permian limestone lying within the core of north-south synclines. ey form a spectacu lar landscape that comprises a series of steep-sided hills and isolated towers rising from alluvial plains, and are clad in dense but stunted vegetation except on tall clis

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ACTA CARSOLOGICA 40/3 2011 498 of bare rock (Fig. 2). ey contain a variety of cave pas sages and undercut foot caves. Between the Setul and Chuping Hills, sandstones form hills rising north to the ailand border, and are overlain by alluvial plains towards the south. Fig. 1: Outline map of northern Perlis. Away from the limestones and the granite, most of the ground is underlain by sandstones and mudstones. Alluvium is extensive on these towards the south, and also oors the wangs within the Sentul H ills. Fig. 2: Steep-sided hills of the Chuping L imestone (Photo: L. Price). Fig. 3: L imestone hills of the Setul Range, with Wang K elian in the foreground (Photo: T Waltham). GEOLOGY OF THE SETUL HILLS is long range of rugged hills is formed in an anticline of the dark, thick-bedded limestones of the Setul Forma tion, of Ordovician to Lower Devonian age. Limestone was not laid down continuously over this period, but was formed during distinct phases separated by intervals of non-deposition; it contains many argillaceous impurities and bands of mudstone, as well as chert and iron oxide. e sequence was folded in Mesozoic times, when it was also intruded by granite (Fig. 1). e Bukit China gran ite forms the highest land in northern Perlis, the tallest peaks being Bukit China at 724 m and Khao China which is across the border in ailand and is 741 m. is is the southern end of a granite body that extends northwards for more than 100 km into southern ailand. e granite has all the features of late-stage miner alisation in its marginal zones, with hydrothermal veins, pegmatites, aplite dykes and greisen. Tourmaline and to paz are common, as well as cassiterite (tin oxide), with relatively minor sulphides. Some of the contacts between the granite and the Setul Limestone are clean, with meta LIZ PRICE

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ACTA CARSOLOGICA 40/3 2011 499 THE MINING INDUSTR Y e centre of the tin mining industry was the small town of Kaki Bukit, on the eastern side of the limestone hills in northern Perlis; its name means at the foot of the hill. Tin mining was an important industry in Perlis from the early 1900s to the 1960s. As it was relatively easy to ex tract the tin from the cave alluvium, many of the caves in the Setul range have been mined. morphic recrystallisation of the calcite to form marble. Other parts exhibit reaction between the limestones car bonate and the granites silica to create complex skarn deposit minerals. Lamprophyre dykes reach far into the limestone and are exposed in a quarry at Kaki Bukit and also in one wall of a cave passage in Gua Kelam 2. Granite weathers rapidly in the tropical climate. Its feldspars and other silicate minerals break down to clays that are rapidly carried away by streams. e quartz and other resistant minerals produce coarse alluvial gravels that ll adjacent valleys and create wide ood plains. W ithin these alluvial gravels, the cassiterite and other heavy minerals are concentrated because of their greater density, forming lenses, strings and horizons of tin in quantities that are suciently rich as to have made mining of them economically attractive. e tin ores are found both in valley alluvium and in the caves (Fig. 4). e alluvial tin on the surface occurs in the oors of deep valleys and depressions within the karst; these are generally surrounded by high clis or steep slopes, and are known locally as wangs, though not all have oors of alluvium. W ang Kelian is the rst big depression south of the granite contact zone (Fig. 1). W ang Tangga is the next alluviated valley to the south. W ang Kelian has sur prisingly little tin in its alluvium and has never been mined, but W ang Tangga has tin-bearing alluvium up to 15 m deep and has been extensively mined. Tin-rich alluvium has been swept into the caves since the formation of the karst, and has proved to in clude some of the richest ore. In most passages the cave alluvium is only a few metres thick, but it reaches much greater thicknesses in large chambers (Jones 1978). At some sites, notably where the current was checked at the foot of waterfalls, the heavy cassiterite is concentrated in lenses and pockets within the alluvium. One pocket in a cave at W ang Mu, south of Kaki Bukit, contained 90% cassiterite and in 1938 yielded tin worth more than a million Malayan dollars (at that time 60 Malayan dol lars = 7 British pounds). But tin values are very variable, with good ore typically carrying between 1% and 5% tin, and much of the alluvium was barren. Alluvial tin occurs in active stream caves, but most now lies in abandoned phreatic passages and chambers. Some of the alluvium is partially cemented by calcite, and in places it is capped by thick calcite owstones that have been dated to ap proximate ages of 280 ka (Smart et al. 1984). e main caves containing the tin lie between W ang Tangga and Kaki Bukit, some 6.5 km south of the granite (Fig. 5), and also in caves further south as far as the Alor Kangar mine, less than 8 km from Kaki Bukit. Fig. 4: Water-rounded alluvial cobbles of cassiterite (Photo: L. Price). Fig. 5: e downstream entrance to G ua K elam 1, with the min ers walkway that reaches through to Wang T angga (Photo: L. Price). TIN MINING IN THE LIMESTONE CAVES OF PERLIS, MALA Y SIA

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ACTA CARSOLOGICA 40/3 2011 500 Little is known of the early history of the tin min ing industry in Perlis. Mining had denitely started by the rst years of the 20 th century, but it is fairly certain that Chinese settlers began to exploit the deposits on a small scale well back in the 19 th century. Prior to the First W orld W ar, primitive methods were used to work the de posits, then European miners arrived in Perlis with more modern systems. is enhanced production gures and brought Perlis to prominence as a tin-producing state. However, mining the alluvium from deep within the caves was dicult and arduous work, and the produc tion of tin ore from Perlis only amounted to 0.7% of the total recovered in Malaya, which at one time was one of the worlds main producers of tin. Records of the mining are known from 1909, but detailed records were only kept aer 1952 when the mines came under the jurisdiction of the Federation De partment of Mines. Aer the Second W orld W ar, Malaya was occupied by the Japanese, during which time the mining industry was virtually dormant, and also many records were lost. e main mines were located within a radius of 4 km of Kaki Bukit. In 1953, there was an average of 25 mines working, of which all were in caves except for two surface workings. Some were small opera tions, with perhaps around ve labourers, while others were huge fully equipped mines. W here possible, the tin miners just followed the natural cave passages, but they also drove access tunnels into some of the caves (Fig. 6). e mining methods were fairly straightforward, as the tin-bearing alluvium is generally free of, or only loosely consolidated by calcite. Some mines, especially those where the alluvium was cemented by calcite, used simple methods where the ore was dug with picks and shovels and taken out by trolley. Out in daylight, the ore was crushed and then separated by dulang washers hand washing using large gently-dished wooden pans to wash the light quartz sand away from the granular cas siterite (Fig. 7). In other mines, high pressure water hoses, known as monitor pumps, were used to wash out the alluvium as a mobile slurry. e pumps were supplied with wa ter pumped in from outside and fed through pipes to the working face. W ater storage tanks were also built in some of the caves. In Gua Baba, the upper part of the cave sys tem was dammed to provide a high level water supply with the necessary pressure to operate the monitors. Separation of the tin ore took place either in the mine or where the washings were pumped out to daylight for processing. Both underground and on the surface, the tin ore was mainly separated in conventional wooden sluices, locally known as palongs; these had rie bars to collect the heavy ore mineral as the alluvium was washed Fig. 6: e articial tunnel driven into a mine beside Wang M engkuang (Photo: L. Price). Fig. 7: D ulang washers improving the grade of ne-grained al luvial cassiterite; these are working at a tin mine in L aos, which is also in a sedimentary basin within a karst terrain (Photo: T Waltham). LIZ PRICE

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ACTA CARSOLOGICA 40/3 2011 501 THE CAVES NEAR KAKI BUKIT Numerous cave entrances and resurgences fringe the valley oors at the base of the limestone range, but en trances are also located high up in the hills. According to the miners, many of these cave systems are connected. Many of the caves still contain mining relics. Several still have the rail tracks for the trolleys, some have wooden walkways suspended by wire cables, and Gua Kelam 2 has an engine room with huge diesel generators. Some of the caves are home to a variety of cave fauna, such as bats, large toads, sh, crabs, and invertebrates such as centi pedes, crickets, millipedes and spiders (Fig. 8). Some of the major caves are listed below. Gua Kelam 1 is a natural cave that gives access through the hill from Kaki Bukit into W ang Tangga (Fig. 9). In 1935, an English miner had a 370 m wood en suspended walkway built above the stream through the entire length of the cave (Fig. 10). is allowed the miners to transport tin ore out from W ang Tangga without wading in the cave stream and to pass cascades of owstone from high-level inlets that almost block the stream passage (Fig. 11). Today the cave is set up as a tourist cave. It is electrically lit with multi-coloured lights, there are information boards. A small entrance fee is charged. Gua Kelam 2 is 3,748 m long and has four entrances (Fig. 9). e hard calcite-cemented alluvium was dug by pick and shovel and transported from the mine in rail trolleys. e material was crushed by mechanical foot stamps and the ore separated by dulang washing. By 1957 the main cave had been worked for a distance of about 610 m, and was cleared of tin-bearing alluvium to a depth of 110 m. Today Gua Kelam 2 is used as a show cave and for adventure caving. Visitors enter on a trolley train, then walk along a concrete walkway for about an hour, passing artefacts that include as old water pipes and mining tools (Fig. 12), and also wooden bridges across the cave lakes. Part of the cave oor has been polished by the stream action to reveal white and black banding in the impure limestone sequence. e exit is from a high level passage in the other side of the hill, below which lies a streamway entrance (Price 1993). down the sluices sloping box channels. Some of the larger mines were tted with modern machinery thou sands of metres inside the hills, and employed gangs of a hundred or more labourers within the caves and mined tunnels. W here the natural cave passages were small, they were enlarged by blasting. In many places, the caves con tained large chambers, and the tin-bearing alluvial de posit could be more than 10 m thick. Some cave passages lled with tin-rich sediment extended to depths of a few hundred metres, and were completely mined out. One of the longest mined caves recorded in 1978 reached a length of about 2,000 m. Some caves were dry. Others contained streams, and ash oods occurred during the rainy season, when some sections of the mines were closed for fear of ooding. e unpredictable ash oods, falling rocks and collapsing wooden walkways were just some of the perils the miners faced in the caves. Surface operations were also concentrated on the Kaki Bukit area, working the alluvium in the open wangs. Gravel pump mines excavate a pond in the al luvium and then pump the gravel slurry from its oor. Hydraulic mines use monitor jets to wash the alluvium from exposed faces. e alluvium in W ang Tangga was protably worked before W orld W ar II, but the ore was exhausted by 1961 and all surface production ceased. Most of the caves were almost mined to exhaustion by the 1960s aer the average labour force had peaked at around 2,500 people. W orld tin prices plummeted in the 1980s, and sealed the fate of the surviving mines. Today a few individual miners still live in the Kaki Bukit area, and some go into the caves to search for any easily avail able tin. Fig. 8: A huntsman spider eating a long-legged centipede, in the small K aki Bukit T emple Cave (Photo: L. Price). TIN MINING IN THE LIMESTONE CAVES OF PERLIS, MALA Y SIA

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ACTA CARSOLOGICA 40/3 2011 502 Fig. 9: Survey of G ua K elam (by the M alaysian Nature Society Cave G roup; redrawn by T Waltham). Gua Baba is a resurgence cave south of Kaki Bukit, and was mined from the 1940s until 1958 (except during the period of the Japanese occupation) as the Khai Fatt Tin Mine (Jones 1978). A narrow stream passage con tinues upstream through ve large chambers. e fourth chamber contained alluvium 10m deep that was famous ly rich in tin, and yielded more than 10 tonnes of tin per month through 1955. A diesel engine pumped water to hydraulic monitors at the work face, and gravel pumps, water pumps and a palong sluice were constructed un derground, with dulang washing sheds built outside the entrance. e miners wooden walkways are still in place, Fig. 10: e miners walkway over the stream through G ua K e lam 1 (Photo: T Waltham). Fig. 11: A large bank of owstone in the G ua K elam 1 stream passage (Photo: L. Price). LIZ PRICE

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ACTA CARSOLOGICA 40/3 2011 503 ACKNO WLEDGEMENTS e author thanks Tony W altham for assistance in preparing this paper and its maps. Fig. 12: An old monitor pipeline with calcite precipitated by seep age water, in G ua K elam 2 (Photo: L. Price). although many are now submerged by water. By 1957 the mining had worked into the cave for a distance of 610 m. e passages are now known to connect to Lo Po Sang which has an entrance high up on the hill, in a system that is around 2,400 m long. Gua W ang Mu is another resurgence cave contain ing some large chambers. e Foh ye Mine has an entrance up on the hill above W ang Mu (Price 2000). It worked gravels from natural cave passages that descend through some large chamber and small streamways, and has been surveyed to a length of 2,100 m. REFERENCES Jones, C.R., 1978: e geology and mineral resources of Perlis, North K edah and the L angkawi I slands .Geo logical Survey of Malaya, District Memoir 17, pp. 257, Kuala Lumpur. Price, L., 1993: Gua Kelam 2.I nternational Caver 7, 11. Price, L., 2000: Perlis 96 and 98.International Caver, 32. Smart, P.L., Andrews, J.N. & B. Batchelor, 1984: Implica tions of Uranium series dates from speleothems for the age of landforms in north-west Perlis, Malaysia: a preliminary study.Malaysian Journal of Tropical Geography, 9, 59. TIN MINING IN THE LIMESTONE CAVES OF PERLIS, MALA Y SIA



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VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMO RELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA AND ABULJA IN BOSNIA AND HERZEGOVINA W BALKAN B IODIVERZITETA IN ENDEMO RELIKTNOST VASKULARNIH RASTLIN KRAKIH GOROVIJ P RENJ VRSNICA IN ABULJA B OSNA IN HERCEGOVINA Sulejman REDIb 1,2 Senka BARUDANOVIb 2 Sabina TRAKIb 2 & Dejan KULIJER 2 1 Academy of Sciences & Art of Bosnia and Herzegovina, Sarajevo, Bosnia and Herzegovina, tel./fax: +387 33 64 91 96, e-mail: sredzic@anubih.ba 2 Center of Ecology & Natural Resources of the Faculty of Science of University of Sarajevo, Bosnia and Herzegovina Received/Prejeto: 7.9.2010 COBISS: 1.01 ACTA CARSOLOGICA 40/3, 527, POSTOJNA 2011 Abstract UDC 581.52:574.1(497.6) Sulejman Redi, Senka Barudanovi, Sabina Traki & De jan Kulijer: Vascular plant biodiversity richness and endemorelictness of the karst mountains Prenj vrsnica abulja in Bosnia and Herz egovina (W. Balkan) e complex of karstic mountains Prenj-vrsnica and abulja in Herzegovina (W Balkan) is characterized by high level of both geomorphology and biodiversity richness. is has been conrmed by a research of plant communities, their structure and dynamics, which took place throughout several seasons from 2005 to 2008. In the investigated area the vegetation cover, as a reliable indicator for specic karstic circumstances, is being dierentiated in a great number of syntaxa (plant com munities) that encompass over 2,500 vascular plants. On the surface of about 100,000 ha identied were up to 236 plant as sociations, 116 alliances and 63 vegetation orders that belong to 34 classes. is amounts 34% of total of vegetation classes at the European level and 100% of so far known vegetation classes in Bosnia and Herzegovina, over 80% of classes at the level of Montenegro and Croatia. ere have been identied nearly 450 endemic and relict species, which is why most of the identied communities are endemic and relict ones, not only at the level of association but also at the level of higher syntaxonomic cat egories, such as alliance and order. e highest diversity level characterizes those communities that make a direct contact with the calcareous geological foundation either in rock crev ices belonging to the class Asplenietea trichomanis Br.-Bl. 1934 corr. Oberd. 1977, or screes on limestone that are comprised by the classes laspietea rotundifolii Br.-Bl.1948 and D rypetea spinosae Quezel 1967, then sub-alpine and alpine pastures Ely no-Seslerietea Br.-Bl. 1948, and rocky grasslands ero-Brachy podietea Br.-Bl. 1947. at high level of oristic and vegetation richness places this area among the most diverse areas both in Europe and whole Mediterranean. Such pattern of vegetation Izvleek UDK 581.52:574.1(497.6) Sulejman Redi, Senka Barudanovi, Sabina Traki & De jan Kulijer: Biodiverziteta in endemo-reliktnost vaskularnih rastlin krakih gorovij Prenj, vrsnica in abulja, Bosna in Hercegovina Obmoje gorovij Prenj, vrsnica in abulja odlikuje ve lika geomorfoloka pestrost in biodiverziteta. To potrjuje tudi raziskava strukture in dinamike rastlinskih zdrub, ki je potekala med leti 2005 in 2008. Vegetacijski pokrov je pomem ben indikator okolja. V obmoju raziskav smo doloili veliko tevilo rastlinskih zdrub, ki zajemajo ve kot 2500 vaskularnih rastlin. Na povrini 100.000 hektarov smo doloili 236 asocia cij, 116 rastlinskih zvez in 63 vegetacijskih redov, ki pripadajo 34 razredom. Zastopanih je kar 34 % razredov, ki jih poznamo v Evropi, vsi razredi Bosne in Hercegovine ter 80 % razredov, poznanih na Hrvakem in v rni gori. Prepoznanih je bilo 450 endeminih in reliktnih vrst, kar pomeni, da je veina zdrub endeminih oziroma reliktnih tako na ravni asociacij, kot tudi na ravneh zvez in redov. Najveja diverziteta je znailna ob neposrednem stika z apnenasto podlago. Tako v razpokah najdemo pripadnike razreda Asplenietea trichomanis (Br.-Bl. 1934 corr. Oberd. 1977), za melia so znailni razredi laspi etea rotundifolii Br.-Bl.1948 in Drypetea spinosae Quezel 1967, alpske panike Elyno-Seslerietea Br.-Bl. 1948 in za kamnite travnike ero-Brachypodietea Br.-Bl. 1947. Veliko oristino in vegetacijsko bogastvo uvra obmoje med tista z najvejo diverziteto v Sredozemlju in Evropi, kar potrjuje izjemen ekoloki pomen Dinarsko-hercegovskega krasa. Kljune besede: Dinarski kras, vegetacija, sintaksonomija, kra ki biotope, kanjon Neretve, endemine in reliktne rastlinske vrste, zatoie, interpretacija z GIS.

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ACTA CARSOLOGICA 40/3 2011 528 S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER (syntaxonomy) and oristic diversity conrms undoubtedly the unique role of dinaric-herzegovina's karst as a complex of unrepeatable ecological factors on global scale. Keywords: Dinaric karst, vegetation, syntaxonomy, karst bi otopes, Neretva canyon, endemic and relict plants, refuge, GIS interpretation. INTRODUCTION In the entire world, there have been noted specic pat terns of diversity in terms of geomorphology, hydrology and biology which are closely related with karst (Katzer 1909; Riter-Studnika 1959; Fairbridge 1968; Herak & Stringeld 1972; Bonaci 1987; W hite 1988; Boievi 1992; Redi 1997; Kogovek et al 1999; Kranjc 2002; Verovnik et al. 2004; Klimchouk et al. 2006; Francisko vic-Bilinski et al. 2004; Florea & Vacher 2007). However, there are some specic features regarding its geogenesis, hydro genesis and syngenesis that characterize the karst of Dinaric Alps (Cviji 1893; Bognar 1987; Kranjc 1994; Culver & Sket 2002; Brancelj 2006; Redi 2008a). All that had led to the creation of diverse karstic phenom ena, such as: karren, sinkholes, poljes, caves, which made the dinaric karst recognizable around the world (Katzer 1909; Milanovi 1979; Spahi 2001; Sket et al. 2001; Trinajsti 2008). Apart from karst poljes, on the territory of mid Dinaric Alps (W Balkan) entire mountain com plexes with numerous karstic attributes occur. Among them special place is taken by the mountains that stretch along the Adriatic coastline from NW to SE (Sliskovic 1995; Prohi et al. 1997; Lui 2003; Redi 2007a). e most unique patterns of karstic-genesis have caused the creation of not only rich over ground and underground orography, but also extremely high biodiversity richness at all levels (Kuan 1969; Horvati 1963; Lakui 1970; Horvat et al. 1974; Lakui & Redi 1989, 1991; Topi & Ilijani 2005, Redi 2007b;). One of the mountain complexes which contain rarely repeatable forms of karstic diversity is the complex of mountains around the Neretva river: Prenj and Vele in east and southeast, vrsnica, Vran and abulja in west and southwest, on the territory of Herzegovina (Katzer 1926; ili 1967, 1979, 1984; Lepirica 2008). In spite of that, this area is still relatively poorly investigated; espe cially its biodiversity for it is the best indicator of karstic diversity and karst as the most unique complex of eco logical factors. e investigation of plant cover (Murbeck 1891; Beck-Mannagetta 1901; Adamovi 1907; Horvat 1933, 1941; Riter-Studnika 1954; Ilijani & Hrak 1990) so far has shown high level of endemism and relictness of vascular ora, as well as of fauna of some animal groups (Sket 1997; Sijari 2000). Apart from the oristic rich ness, the investigation of syntaxonomic diversity so far has conrmed the existence of several both endemic and steno endemic plant communities, of which some are exclusively associated with this area, such as M inuartio handelii-Caricetum Bjeli & ili 1979 at the top of Veli ki Vilinac, H eliospermo retzdornani-Oreoherzogietum il lyricae ili 1970. In the canyon of Neretva river, of about 2,000 m height, and canyons of its tributaries: Rakit nica, Ljuta, Treanica Idbar, Bijela, Neretvica, Doljanka, Glogonica and Dreanka, special refugia of both ora and vegetation from Tertiary, which are characterized by extreme richness (ili 2000; Redi 2004, 2008b), are identied. ere are many leads that this area represents special a kind of the biodiversity hotspot, which should be more closely claried through further investigation. e investigation that has been conducted so far indi cates the existence of development endemic-relict centre situated both at the mountain foots and their peaks as well. Here have evolved many tertiary and glacial relicts. As a special indicator for this richness is consid ered to be the syntaxonomic diversity which integrates the diversity of both ora and habitats (Ilijani 1989; Barudanovi & Redi 2006; Redi 2007c, 2008c, 2008d; arni et al. 2009). Besides, the syntaxonomic diversity is the excellent indicator for the entire ecological diversity. In addition to its fundamental signicance, the syntaxo nomic diversity can be applied like a tool for the sustain able management in this, every day more threatened, karstic area. is implies that the syntaxonomic diversity can be a basis for successful biological and ecological in situ conservation. rough this investigation formulated was the hypothesis that extremely high level of oris tic diversity correlates closely with the syntaxonomic (vegetation) diversity. Main target of this paper was to assess the syntaxonomic diversity in relation to process of karst-genesis, especially in such habitats where the in uence of karstic foundation is the most direct in terms of speciation, ended-genesis, such is the case with rock crevices and screes. is kind of approach will undoubt edly illustrate the crucial role of this dinaric karst area in terms of biodiversity patterns.

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ACTA CARSOLOGICA 40/3 2011 529 GEOGRAPHIC POSITION e investigated area is situated in the complex of high mountains of Herzegovina and determined by coordi nates 17 and 19 E; 43 and 44 N. e considered area represents high upland framework constituted of Prenj, vrsnica and abulja mountains. It is positioned in northern Herzegovina, in the area of middle and upper ow of Neretva river (Fig. 1). As far as geographic regionalization is concerned, this territory is part of Bosnia and Herzegovina that be longs to high karst macro-region or its subunit, meso region of northern Herzegovinas upland. e territory takes approximately 1,000 km, stretching from the ac cumulation lake of Jablanica in the North to Bjelopoljs ka valley in the South, and from the surrounding of Borako lake in the East to distant north-western slopes of abulja mountain in the W est. In the physical and geographical sense, the con cerned part of Neretva river basin is distinguished by its relief, climate, soils, as well as by its geological, hydro logical and biological diversity. Afore named attributes and specic geographical position have aected this rela tively small area in a way that it became rich in natural values and distinguished by extremely high level of geoand biodiversity (Miladinovi 1964). GEOLOGY e investigated area, aer geological classication on regional scale, is a part of the Outer or karstic Dinaric Alps, which is the subunit of Dinaric Alps (Cviji 1989). "Dinaric Alps involve structural complex of carbonate platform with relatively unied carbonate sedimentation (calcite and dolomites) with several intrusions of baux ite, breccias etc. e structural relationships inside the Dinaric Alps are rather com plex. Dierences in altitude are induced by elevation, dierences in erosion rate, lling of local and regional valleys with new depositions, but also by previous second ary (tectonic) accumulation of structural complexes in form of scaling and drawing." (Herak 1983). Movements in vertical sense, which took place at the end of Neogen and dur ing Quaternary, have resulted in rising of faulting of Prenj, vrsnica and abulja mas sifs to their current height and creation of Neretva river canyon with deeply truncated valleys of its tributaries. e most frequently occurring is Mesozoic alluvium of Tri assic, Jurassic or Cretaceous age (Katzer 1926; ii & Pami 1979; ii et al. 1984; Lepirica 2005). e most dominant lithological units are limestone, dolomites and limestone and dolomites combined. e geologically most recent alluvial deposition covers the valley bottom just beneath the mountain slopes. Moun tain lakes are oen associated with glacial processes, which has led to development of special kind of plant communities. O ROGRAPHY In the geomorphologic sense, the investigated area of Neretva river basin is situated in the mega-geomorpho logic region of Dinaric Alps of Bosnia and Herzegovina. More precisely, the area of Prenj, vrsnica and abulja mountains, as well as of valleys of Neretva river and its tributaries, belong to the geomorphologic meso-region of the middle Dinaric Alps. By its morphology, this en PHY SIO GEOGRAPHIC FEATURES OF ENDEMIC DEVELOPMENT CENTER OF PRENJ, VRSNICA AND ABULJA MOUNTAINS VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ... Fig. 1: G eographic position of endemic development centre of Prenj, vrsnica and abulja mounmoun tains on the territory of W. Balkan.

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ACTA CARSOLOGICA 40/3 2011 530 tire area has got polygenetic features due to inuences of several geomorphologic factors which took eects at the moment of its creation and development. It is important to stress that the relief development in northern Herze govina was primarily aected by active exogene geomor phic processes. e morphological structures of Prenj mountain (2,155 m), vrsnica mountain (2,228 m) and abulja mountain (1,798 m), as they are today, were predis posed by massive neo-tectonic upli during Quaternary (Buatlija 1974; Lepirica 2005, 2008). In the past, specic forms of both geoand geomor phogenesis that were followed by karstogenesis have re sulted in extremely high relief diversity and uniqueness of geomorphological shapes going from the lowest to the highest positions of this karstic complex. e diversity in terms of geomorphology represents one of the cru cial determinants of most unique biodiversity in which many endemo-relict both species and communities are contained. HYDROLOGY Main hydrological features to the entire complex of Prenj, vrsnica and abulja mountains are undoubtedly given by Neretva river and its tributaries. From the Neretvas spring toward Mostar city, its right auents are as fol lows: Ljuta, Treanica, Neretvica, Doljanka, Grabovica and Drenica. e most important auents along the le river bank are: Laanica, Konjika Bijela, Idbar, Glogonica, Mostarska bijela. Today, the most impressive part of the Neretvas canyon, its mid part, between Prenj, vrsnica and abulja mountains, is turned into articial hydro-accumulation lakes "Grabovica" and "Salakovac". e upper ow of Neretva river, from its spring un der Gredelj and across Bjelopoljska valley, drains vast area of northern and northeastern Herzegovina. Based on the identied boundary, the surface basin of upper and middle ow of Neretva river covers the area of about 2,500 km. However, due to profoundly karsti ed structure of this surrounding, which is built of tick carbonate and dolomite layers from Mesozoic, the sur face is holed and cut by a system of crevices, gaps and chennels, making the hydrological network become disarranged and transmitted into karstic underground. Hence, the extent of catchment area cannot be dened precisely. As an example, in the Kalinovako Zagorje a sinking creek that ows on the right side from the vil lage Jelace, corresponds with Gornji and Donji Vrutak, which are actually springs that belong to the catchments area of Bistrica creek. Furthermore, the adjacent Tatinac creek and sinking creek at Kalinovik carry large quan tity of water into the Neretva river. e latter represents aquatic linkage with the biggest spring in upper Neretva area Krupac. ese underground currents appear on the surface as powerful karstic springs, diering in their abundance, functionality and stability. Mostly, they dont ow on the surface for a long time. e uctuation of water level in Neretva river is uneven over a year. Aer it reaches maximum in rst half of May, water level declines slowly and outow falls down to reach its lowest values in August and September. From October, with increased precipitation, the outow rises and reaches its maximum in the middle of Novem ber and over entire December. Due to large ice quanti ties and thick snow cover, including low temperatures in mountainous area, water level stagnates in January and February, which is followed by its slow rising in March. From the year 1926 to 1980 the following absolute values of Neretvas outow were obtained: absolute maximum in outow was measured to be 926 m 3 /s, while the abso lute minimum was 4 m 3 /s. e average annual outow of Neretva river at Konjic was 52.3 m 3 /s (Lepirica 2007). In the investigated area occur only several perma nent natural lakes. ese lakes are: Borako and Blidin jsko lake, and on vrsnica mountain: Crvenjak, Gavrani and Lake on Mejdan, including Jezerce and lake under Vrine on Prenj mountain, which are small and of no hy drological importance. eir sole function is to provide cattle with water. Many of these lakes arose by block ing the bottom of sinking rivers that was carried out by cattle breeder, likewise Crepulja on Plaso. e Borako lake is considered to be the biggest and most important natural hydro-accumulation in the upper ow of Nere tva river (Spahi 2001). e hydrological regime in highmountainous area is strongly inuenced by ice and snow retained for a long time. Large quantities of snow are be ing retained at the bottom of sinkholes over entire sum mer. is kind of habitats became shelters for numerous relict both species and communities during glaciation. Beneath mountain glaciers are to be found many unique springs with cold and fresh water. One of such springs is the spring of Jasle beneath Plono peak in the natural reserve with well preserved woods of endemic Bosnian pine ( Pinus heldreichii H.Christ ) ECOCLIMATE e exceptional vertical dierentiation of relief that in cludes pronounced de-levelling in altitude and protru sion of inuences from mild Mediterranean climate (coming through the valley of Neretva river in reduced extent) have had a major impact on development of sev eral climate types and subtypes occurring at this relative ly small area. e area around Neretva river is characterized by modied Mediterranean climate. In the surrounding, at higher altitude, climate changes into sub-montane. Over S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 531 1,000 m above sea level, montane climate stretches over major part of this watershed. At altitude over 1,700 m, on clis and crests of Prenj, vrsnica and abulja moun tains climate is alpine, harsh and cold. At lower altitudes, between 200 and 450 m above sea level, valleys of Nere tvas tributaries are deeply cut in mountains; hence, the area is aected by transitional climate inuences with cantonal features (Lepirica 2007). Based on records of mean air temperatures both monthly and annually, as well as on precipitation rate, measured at the nearest weather stations in this region (Mostar, Jablanica, Konjic and Nevesinje) (Tabs. 1) supervenes that dierent climate types occur: (i) mod ied-Adriatic climate (at the lowest positions) which protrudes all the way up to Jablanica in North, (ii) Adri atic climate (southern slopes of Prenj mountain toward Mostar city), (iii) transitional lowland climate (mid and upper ow of the Neretva river), (iv) pre-montane and montane climate, taking places above 700 m above sea level, and (v) alpine climate above 1,700 m. HUMIDIT Y OF ECOCLIMATE Climate humidity is an extremely important parameter for determination of not only the type and character of climate, but also for denition of vegetation character in cluding its tendencies in relation to future syngenesis. e climate humidity is dened by precipitation quantity un der given temperature conditions. It is expressed in sever al ways. In ecology as a very practical and widely accepted interpretation of this parameter has been conrmed to be the method aer Graanin (1950). Aer this methodol ogy, ratio between precipitation quantity (monthly or an nually) and mean temperature of air (monthly or annu ally) represents rain factor (RF) (Tabs. 1). Due to high precipitation rate, the ecoclimate at each meteorological station is highly humid. However, the ecoclimate in veg etation season becomes either semiarid or arid resulting in the development of xero-thermophyllous vegetation encompassed by orders Ostryo-Carpinetalia orientalis Lakui, Pavlovi & Redi 1982 and Quercetalia pubes centis Br.-Bl. (1931) 1932, which occur at lower mountain parts (Tabs. 1). WARM CHARACTER OF ECOCLIMATE Regarding its warmth character the ecoclimate is very di verse. It is nivale over winter (the meteorological station Nevesinje) and warm, even hot, in summer (Tabs. 1). is warm climate correlates positively with the develop ment of thermophyllous vegetation units from the class Querco-Fagetea Br.-Bl. & Vlieg. 1937, and thermophyl T ab. 1: Characteristics of eco-climate noted at meteorological station Nevesinje. L egend: H umidity* (for tab. 1): ph per humid; h humid; sh semi humid; sa semi arid. Warm character of eco-climate** (For tab. 1): n nivale; hl cold; uhl temperate cold; t warm; v hot; uttemperate warm Meteorological Station: NEVESINJE (900 m) Month I II III IV V VI VII VIII IX X XI XII Average annual value Precipitation (mm) 161 175 161 118 148 101 74 62 118 160 213 280 1.771 Temperature (C) -1.2 0,2 2.9 7.7 12.5 15.6 18 18.4 14.6 10.6 5.9 1.1 8.9 RF m (Graanin) 159.8 875 55.51 15.32 11.84 6.47 4.11 3.36 8.08 15.09 36.1 254.5 198.98 Humidity* ph ph ph ph h sh sa sa h ph ph ph ph Warm character of ecoclimate** n n n uhl t t t t t ut uhl hl ut T ab. 2: Characteristics of eco-climate noted at meteorological station M ostar. Meteorological Station: MOSTAR (65 m) Month I II III IV V VI VII VIII IX X XI XII Average annual value Precipitation (mm) 163 150 134 123 92 84 46 62 99 136 220 230 1.539 Temperature (C) 4.7 6.2 9.7 13.8 18 22 25 24.8 20.8 15.6 10.7 6.8 14.8 RF m (Graanin) 34.68 24 13.81 8.91 5.1 3.8 1.8 2.5 4.75 8.71 20.6 33.82 103.99 Humidity ph ph h h sh sa a a sa h ph ph ph Warm character of ecoclimate uhl uhl ut t t v v v v t ut uhl T VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 532 lous meadows from the class Festuco-Brometea Br.-Bl. & Tx. 1943, as well as rock debris communities on lime stone from the class ero-Brachypodietea Br.-Bl. 1947. Summarized characteristics of humidity and tem perature are given in Tabs. 1, aer records from me teorological stations which are to be found in the inves tigated area. Although the precipitation rate noted at majority of meteorological stations is very high (mari time type), the ecoclimate is considered to be arid (dry) in some periods of year, which results in the occurrence of typical karst ora and vegetation. Plant cover in karst regions is strongly determined by wind. In karstic habitats, especially of southern and western aspects, formative power of wind comes to its full expression. In habitats that are more protected from wind a luxuriant vegetation of karst woods with whitebarked pine Amphoricarpi-Pinetum leucodermis devel ops Fukarek 1966 and even cold woods with spruce and r Abieti-Piceetum illyricum Stef. 1963, which is a true rarity in the Dinaric karst region. W hile southern and south-western slopes of vrsnica mountain are covered by low woods and thickets, its northern slopes are cov ered by luxuriant woods that are considered to be a true natural reserve, such as Masna Luka. SOIL In this area dierent types of soil occur, which were in duced by specic climate types, geological base, wind and other ecological factors. On carbonate bedrock diverse development series of soil occur. Beginning with the un developed ones, these soils are: litosol (rocky type of soil), regosol (on broken rocks), black earth on limestone or calcomelanosol, rendzine, whereas on less steep slopes oc curs brown earth on limestone or calcocambisol. In some places, over at ground, also ilimerised soil type or luvi sol occurs. e hydromorphous soils occur only on river banks of small watercourses, such is the case with plano histosol (alkaline bogs), and gley or eugley which occur along large water bodies, such as alluvium or uvisol. On bare carbonate bedrock lithosol prevails. is is a shallow type of soil, oen of less than 1 mm deep. Simi lar type of soil is regosol, but it occurs only on broken rocks, oen in rock ssures of large carbonate blocks. is soil type has got undeveloped humus-accumulative horizont, of (A)-C prole. Despite being shallow and poor in nutritient, this soil is overgrown mostly by endemic and relict plants which constitute numerous endemo-relict communities in rock crevices belonging to vegetation orders Amphori carpetalia Lakui 1968, M oltkeetalia petraeae Lakui 1968 and Potentilletalia caulescentis Br.-Bl. 1926, as well as communities on screes from orders Arabidetalia a vescentis Lakui 1968 and D rypeetalia spinosae Quezel 1967 On rendzine and calcomelanosol occur broad leaved-deciduous woods and thickets belonging to or T ab. 3: Characteristics of eco-climate noted at meteorological station Jablanica. Meteorological Station: JABLANICA (202 m) Month I II III IV V VI VII VIII IX X XI XII Average annual value Precipitation (mm) 177 171 167 161 107 104 64 101 101 184 342 366 2.045 Temperature (C) 1.5 3.4 7.4 12.6 16.5 19.7 21.8 21.5 18.1 13.2 9.5 3 12.4 RF m (Graanin) 118 50 22.56 12.8 6.48 5.28 2.93 4.69 5.58 13.93 36 122 164.92 Humidity ph ph ph h sh sh a sa sh ph ph ph ph Warm character of ecoclimate hl hl uhl t t t v v t t ut hl t T ab. 4: Characteristics of ecoclimate noted at meteorological station K onjic. Meteorological Station: KONJIC (277 m) Month I II III IV V VI VII VIII IX X XI XII Average annual value Precipitation (mm) 141 177 96 102 102 76 58 54 104 134 198 222 1.464 Temperature (C) 0.5 2.4 6.1 10.9 15.4 18.5 19.8 20.1 16.7 12 8 3 11.1 RF m (Graanin) 282 74 15.73 9.35 6.62 4.11 2.93 2.68 6.22 11.16 24.8 74 131.89 Humidity ph ph ph h sh sa a a sh h ph ph ph Warm character of ecoclimate hl hl uhl ut t t t v t ut uhl hl ut S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 533 ders Ostryo-Carpinetalia orientalis Lakui, Pavlovi & Redi 1982 Quercetalia pubescentis Br.-Bl. (1931) 1932 and Fagetalia Pawl. 1928, and rocky grasslands from or ders Scorzonero-Chrysopogonetalia Horvati & Horvat (1956) 1958 K oelerietalia splendentis Horvati 1975 and thermophyllous meadows from order Brometalia erecti (W Koch 1926) Br.-Bl. 1936. On these soils (if more organic by nature) in sub-alpine and alpine belt prevail grasslands on carbonate foundation encompassed by or ders Seslerietalia juncifoliae Horvat 1930 Crepidetalia di narice Lakui 1966 and Edraiantho-Seslerietalia Redi 2003, while around snowbeds vegetation order Saliceta lia retusae-serpyllifoliae Lakui et al 1979 occurs Brown earth on limestone, so called calcocambisol, is associated with mild slopes and woods of white-barked pine combined with black pine belonging to order Pin etalia heldrechii-nigrae Lakui 1972, as well as beech woods of Fagetalia Pawl. 1928 order, and temperatehumid meadows belonging to order Arrhenatheretalia Pawl. 1928 (Redi et al. 2007). Inspite of karstic terrain, in some places occur swampy and boggy types of soil (planohistosol), such is the case in Masna Luka, Blidinje (on vrsnica mountain) and Borako lake, and on Prenj mountain. In the valley of the Neretva river and its auents colluvium and u visol occur frequently, with relict plant communities of grey willow and sage Petterio-Salicetum incanae Redi et al. 1992). MATERIAL AND METHODS APPROACH AND MISSION e main parameter in this assessment was the biodiver sity (diversity), as understood by the most recent Inter national Community beliefs OUN AGENDA 21, Rio de Janeiro 1992. (Sitarz 1993), and in the scope of new achievements of modern science, especially those of con servation biology and ecology. Besides, we assessed all the elements which are required to determine both forms and levels of biodiversity, like orography, geological and pedological framework, ecoclimate and anthropogenous impact reected through diverse human activities. FIELD METHODS e eld research on spatial and temporal biodiversity organisation, as well as on both geomorphological and hydrological features of endemic centre Prenj, vrsnica, abulja mountains, was conducted in 2007 and 2008 throughout several seasons (early spring, summer, and early autumn). Apart from data collected during this re search, there have we also used data obtained in previ ously conducted eld investigations or expert excursions accomplished by some team members. Many localities on both longitudinal and transversal prole of each moun tain belonging to endemic centre of Prenj, vrsnica and abulja mountains were been analised (Figs. 2 & 3). Each locality was precisely positioned by GPS, as it is shown on the map. A SSESSMENT OF FLORISTIC AND VEGETATION DIVERSIT Y e assessment of oristic diversity of macrophytes (vascular plants) was carried out on previously cho sen transects, one longitudinal and several transversal proles, of Prenj, vrsnica and abulja mountains, as well as on a great number of spots along eld margins, respectively. It was determined the representative area size, which is the space in which all plants were identi ed. Apart from that, herbal material was gathered in the eld and properly preserved and determined aerwards by usage of relevant botanical keys, from both local and international oristic literature (Hayek 1927; Tutin et al. 1964; Beck et al. 1983; Oberdorfer 1983). e analysis of plant communities and vegetation diversity were carried out on exceptionally large num ber of points (Figs. 2 & 3), in all ecosystem types of en demic centre of Prenj, vrsnica and abulja mountains. e size of analised plots was 100 m 2 (for non-forest community types), respectively, 200 and 500 m 2 if for est ecosystem was concerned. Each plot was analysed in terms of abundance and coverage, as well as of sociabilty and vitality assessment for each of detected species. e Braun-Blanquet methodology was applied, or method ology of Zurich-Montpelier's School, which is generally accepted for all kinds of vegetation biodiversity studies (Braun-Blanquet 1964). Formulating of denition and concept of basic veg etational units syntaxa were carried out in accordance with the I nternational Code of Phytosociological Nomen clature (W ebber et al. 2000). Aer this Code, vegetation is organised in associations (basic unit), alliances (com prising more kindred associations), orders (comprising more kindred alliances) and classes (comprising more both oristically and ecologically kindred orders). VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 534 Beside the standard methodol ogy for the assessment of diversity in terms of species and ecology (Magur ran 1988; Scheiner 1992), the meth odology for the assessment of syn taxonomic diversity as an indicator for vegetation and oristic diversity was applied (Redi 2007c, 2007d, 2008b). Each of the organisation units has got its own sux. e nomencla ture of vegetation units is given aer Prodromus biljnih zajednica BiH (Lakui et al. 1978), Prodomus phy tocoenosum Jugoslaviae ad mappam vegetation M 1: 200 000 (Jovanovi et al. 1986) and e D iversity of Eu ropean V egetation (Rodwell et al. 2002). W hen it comes to interpre tation of vegetation diversity, apart from previously recognized commu nities the newly discovered commu nities, which were provisionaly de scribed aer regulation of the Code of phytocoenological nomenclature, were also taken into account (W ebber et al. 2000). e intention of the paper is not to describe such communities in de tail, but to use them as a bioindicator for karstic biodiversity. ose com munities will be fully phytocoeno logical interpreted in our next syn taxonomical communication. In the following text they are marked with an asterisk (*). ANAL Y SIS AND DIGITISING OF DATA B Y GIS METHODOLOGY e largest share of vegetation units was interpreted on maps in scale 1: 25 000. For the spatial analysis and cartographic interpretation of records obtained in the eld, ArcView pro gramme package version 3.2. was ap plied, whereby as a cartographic back ground 1: 25 000 maps were used. e Fig. 2: e area of Prenj mountain with main localities where biological and geo morphological diversity were analised. S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 535 analysis was carried out on the base of vast material collected during ei ther recent or previous eld research (including co-ordinates and records from the eld maps). During the eld work well documented and described records were included into the GIS database, which encompasses information on distribution of abiotic and biotic components in the investigated area (Tab. 5). Hence, the GIS database contains a series of spatial records on both abiotic and biotic component in the investigated area. Information on quantity, distribution and status of plant communities were also import ed into the database and linked with the appropriate GIS layer (Ashdown & Schaller 1990). e spatial analysis of GIS records enabled the identica tion of areas with high ecological and biological value. Fig. 3: e area of vrsnica and abulja mountains with main localities where biwith main localities where bi ological and geomorphological diversity were analised. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 536 RESULTS AND DISCUSSION D IVERSIT Y OF VASCULAR PLANTS e biodiversity of vascular ora in the mountain com plex that surrounds Neretva river (Prenj, vrsnica and abulja mountains.) is extremely rich (Riter-Studnika 1956; Lakui 1973; Lakui et al. 1982; Beck et al. 1983; ili 1984, Redi et al. 2007). Out of almost 5,000 taxa, as it is estimated to occur on the territory of Bosnia and Herzegovina, there are around 2,500 taxa found in this area (Redi 2009). In spite of some previously undertaken oristic in vestigations, the biodiversity level is relatively unknown. Besides, the area of endemic centre Prenj, vrsnica and abulja mountains is unique and hardly repeatable on bigger scale as far as the number of endemic taxa is concerned. Out of 450 endemic taxa, as it is estimated to occur in Bosnia and Herzegovina, nearly 50% dwells in this area (Bjeli 1987; Redi et al. 2008). erefore, the area of Prenj, vrsnica and abulja mountains is des ignated as a specic "hot spot" in respect to its oristic richness, as well as the level of endemism. It is one of the most important endemic development centres in Balkan Peninsula, and one of the largest in the Mediterranean and SE Europe, which is the reason why it is considered as the area of global value (Bjeli & ili 1971; Redi et al 2008). Beside wider distributed species, there is a sig nicant number of species belonging to alpine oral element that link this area with the Alps (Redi 1999, 2003). On the foot of the mountains and in the canyon of Neretva river a signicant number of plants belong ing to Mediterranean oral element occur, which in o ristic respect connects this area with the Mediterranean region. A great number of Illyrian, dinaric and Balkan oral elements found their refuge within this area. Be sides, some species, that are characteristic exclusively for this endemic development centre, have also been found (ili 1970; Bjeli & ili 1971). ere are more than 170 both species and subspe cies which were recognized as signicant geo-fund en compassed by the Park of nature "Blidinje" (ili 2002). From the standpoint of systematic belonging, species of following families prevail: Compositae, Caryophyllaceae, Fabaceae, Scrophulariaceae, L amiaceae, Rosaceae and others. e proportion of plant families in the vegetation of this area are correlated with the proportion of plant families in the ora of Bosnia and Herzegovina (Redi et al. 2007, 2008). e most recent research of ora in the karstic part of Bosnia and Herzegovinas Dinaric Alps indicates that within this mountain complex live more than 500 species of vascular plants that are consid ered to be endemic-relict (Redi 2009). is high level of endemism of vascular ora correlates with signicant extent of the high level of syntaxonomic diversity, which reects the uniqueness of karstic habitats in the most ap propriate way. P HY TOGEOGRAPHY From biogeographical point of view the area of endemic development centre which is to be found within moun tain complex of Prenj, vrsnica and abulja has a very unique position (Fukarek 1949, 1965; Lakui 1969, 1981; Bjeli & ili 1971; Redi & Dizdarevi 1998; ili 2000; Redi 2004). is is the area where two large phy togeographic regions collide: (i) Euro Siberian North American and (ii) Alpine high Nordic region. Besides, there is strong inuence coming from the Mediterranean region in south. EUROSIBERIAN NORTH AMERICAN RE GION extends across the largest portion of the endemic development centre. It includes a broad belt from the lowest sub-montane area to upper timberline. In the investigated area, the region is divided into three lower phytogeographic units, so called, provinces: (i) Illyrian, (ii) Moesian and (iii) Province of relict black pine for ests. Illyrian province on both horizontal and vertical prole of endemic centre of Prenj, vrsnica and abulja T ab. 5: GIS records sets. GIS data set Description Type of feature Scale Position Position of the investigated area given on the map in scale 1:25 000 topographic maps 1:25 000 Geology Geological foundation layers taken from the geological map in scale 1:100 000 polygon 1:100 000 Vegetation Location for each investigated vegetation class given on the base of literature records polygon 1:25 000 Biodiversity Analysis of zones with high biodiversitys level carried out on the base of records contained in the database polygon 1:25 000 S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 537 mountains is dierentiated into the following vegeta tion belts or sub-belts: (i) Xero-thermophyllous decidu ous forests and thickets of pubescent oak, Turkey oak and Italian oak belonging to order Quercetalia pubes centis Br.-Bl. (1931) 1932, in supra-Mediterranean belt; (ii) Xero-thermophyllous deciduous lower forests and thickets of oriental hornbeam belonging to order Ostryo Carpinetalia orientalis Lakui, Pavlovi & Redi 1982 (alliance Carpinion orientalis Blei & Lakui 1966); (iii) Xero-thermophyllous deciduous forests and thickets of hope hornbeam belonging to order Ostryo Carpine talia orientalis Lakui, Pavlovi & Redi 1982 (alliance Seslerio-Ostryon Lakui, Pavlovi & Redi 1982); (iv) Temperate humid sessile oak hornbeam forests (alli ance Carpinion betuli Oberd. 1953); (v) Beech forests be longing to order Fagetalia sylvaticae Pawl. 1928 (alliance Fagion moesiacae Blei & Lakui 1970 and AremonioFagion Trk et al 1989) (vi) Dark coniferous forests of spruce and r (alliance V accinio Piceion Br.-Bl. 1938) and (vii) Low thickets of mountain pine (alliance Pinion mugi Pawl. 1928 "I llyricum" ). Beech forests belonging to order Fagetalia sylvaticae Pawl. 1928 (alliance Fagion moesiacae Blei & Lakui 1970 and Aremonio-Fagion Trk et al 1989) are very di verse and include: (i) Montane temperate humid beech forests (association Fagetum silvaticae montanum Hor vat 1938); (ii) ermophyllous beech forests (alliance Seslerio-Fagion sylvaticae Redi & Barudanovi 2010); (iii) ermophyllous forests of beech and Acer obtusa tum W aldst. & Kit. ex W illd. (association Aceri obtusa ti-Fagetum Fabijani, Fukarek & Stefanovi ex Fukarek, Stefanovi & Fabijani 1967; (iv) Upland mixed woods with beech and r (association Abieti Fagetum (Fukar ek & Stefanovi) Fukarek 1969); (v) Mediterranean mon tane beech forests (association Fagetum "mediteraneomontanum" Redi et al. 1984); (vi) Sub-alpine beech forests (association Aceri Fagetum moesiacae "subalpi num" Fukarek & Stefanovi 1958). Moesian province extends across the warmer share of the area, on horizontal and vertical prole. ese are mainly canyons and steep slopes with southern aspect. e territory of Moesian province is dierentiated into vegetation belts and sub-belts as follows: (i) Xeric forests of Italian oak (alliance Quercion confertae Horvat 1958 ) (ii) Meso-thermophyllous forests of sessile oak (alliance Quercion robori-petraeae Br.-Bl. 1931) (iii) Beech forests belonging to order Fagetalia Pawl. 1928 (alliance Fagion moesiacae Blei & Lakui 1970). Beech forests include: (i) montane beech forests (association Fagetum moesiacae "montanum" Blei & Lakui 1970); (ii) ermophyllous forests of Moesian beech and autumn moor grass (association SeslerioFagetum moesiacae Blei & Lakui 1970); (iii) ermo phyllous forests of Moesian beech and Acer obtusatum W aldst. & Kit. ex W illd. ( association Aceri obtusati Fag etum moesiacae Fabijani, Fukarek & Stefanovi 1963); (iv) Upland forests of Moesian beech and r (associa tion Abieti Fagetum moesiacae Blei & Lakui 1970) and (v) Sub-alpine forests of Moesian beech (associa tion Aceri Fagetum moesiacae subalpinum Fukarek & Stefanovi 1958). W oods of Moesian beech oen alter nate with woods of white-barked pine, or mountain pine thickets Pinetum mugi that makes here the upper tim berline. Province of relict black pine forests extends across the area with endemic white-barked pine Pinus heldre ichii H. Christ and Illyrian black pine pine ( Pinus nigra J.F.Arnold susp. a ustriaca (Hoss) Bid.). In terms of its ecology and territory, the province is dierentiated into several clearly divided phytogeographic sectors: (i) For ests of Illyrian black pine on dolomites belonging to alli ance Pinion austriacae Horvat 1959 (ii) Forests of Illyrian black pine of sub-alliance Orno Ericenion "dolomiti cum" Horvat 1959 (iii) Community with Illyrian black pine in crevices of carboniferous rocks ( Onosmo Pin etum "illyricum"* ) and (iv) Forests of white-barked pine belonging to alliance Pinion heldreichii Horvat 1950 Forests of alliance Pinion heldreichii Horvat 1950 are endemic-relic and typical for this part of Dinaric karst. is alliance includes endemic communities: (i) Mediterranean montane forests of white-barked pine of association Pinetum heldreichii "mediterraneo monta num" Blei & Lakui 1969 ; (ii) Upland forests of whitebarked pine and spruce ass. Piceo Pinetum heldreichii Lakui; (iii) Upland forests of beech and white-barked pine ass. Fago Pinetum heldreichii Jankovi (1972) 1975 (iii) Montane forests of white-barked pine ass. Pinetum heldreichii "subalpinum" Fukarek 1967 and (iv) Associa tion with white-barked pine in crevices of carboniferous rocks Amphoricarpo-Pinetum heldreichii Fukarek 1967 ALPINE HIGH NORDIC REGION continues in ecological sense on sub-alpine belt of Euro Siberian North American region. It encompasses the area of high mountain peaks, above upper timberline, mostly with mountain pine communities Pinetum mugi Lakui 1977 s.lat., and in some cases above low forests of white-barked pine Pinetum heldreichii "subalpinum" Fukarek 1967, or above sub-alpine forests of beech Fagetum "subalpinum" "dinaricum" Tregubov 1957 Here, this region is repre sented by province of high Dinaric Alps. On a vertical prole this province is dierentiated into three belts: (i) Discontinued belt of sub-nivale vegetation around snow beds on carboniferous ground (alliance Salicion retusae Horvat 1949 in Horvat, Glava & Ellenberg 1974) and (ii) Alpine grasslands or high alpine tundra VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 538 Alpine and sub-alpine pastures or high mountains tundra include those communities: (i) Alpine grasslands on the highest places exposed to strong winds with nar row leaved moor grass ( alliance Seslerion juncifoliae Hor vat 1930) (ii) Alpine grasslands on the highest places with dinaric alliance Oxytropidion dinaricae Lakui 1966 (iii) Alpine grasslands on the highest places with Prenjs al liance Oxytropidion prenjae* and (iv) Alpine grasslands in sheltered places (alliance Festucion bosniacae Horvat 1930 corr. Redi 2007). Sub-alpine grasslands (oen of secondary character) include: (i) Sub-alpine grasslands in exposed habitats alliance Seslerion robustae (Lakui et al. 1982) Redi 2003 and (ii) Sub-alpine grasslands in sheltered places Stachydi-Festucenion bosniacae*. Due to numerous oristic and vegetation speci cities, the province of high Dinaric Alps in this area is represented by high Prenjs sector. is phytogeographic sector includes mountains of the endemic development centre of Prenj, vrsnica and abulja, and large propor tion of Vele mountain. is sector is characterized by a great number of endemic species: M inuartia handelii Mattf., Campanula hercegovina Degen & Fiala, Asperula hercegovina Degen, Arenaria gracilis W aldst. & Kit., Am phoricarpos autariatus Blei & E.Mayer, D ianthus pren jus Beck, Euphorbia hercegovina Beck, D ianthus freynii Vandas, Saxifraga prenja Beck, Edraianthus hercegovinus K.Mal, Oxytropis prenja (Beck) Beck, and others. All enlisted, as well as many other species in this area, con stitute around 250 associations, out of which signicant number is of endemic or steno-endemic character (see syntaxonomic overview of associations) that provides re markable and unique features to this area. D IVERSIT Y OF HABITATS AND VEGETATION e area of endemic centre Prenj, vrsnica and abulja is highly diversied in terms of vegetation or ecological diversity. On this, in respect of abiogen, heterogeneous space, 236 ecological units of association level occur, en compassed by several ecological oristic systems of alli ance level (116) and 63 orders, and as much as 34 vegeta tion classes. Vegetation of clis and rocks ( Asplenietea tricho manis Br.-Bl. 1934 corr. Oberd. 1977) Even though they are on the lowest level of ecologi cal integration, these eco systems are very present in the investigated area. Com munities of these particular ecosystems give the strongest proof of uniqueness and sin gularity of this area. Unlike many others, ecosystems of cracks in the rock are devel oped at the whole vertical prole (between 250 m and 2,220 m) of endemic devel opment centre. As they ap pear in all vegetation zones, they are undoubtedly intrazonal in character. erefore, besides their own dierential peculiarity, they also have additional characteristics that are caused by zonal or climate regional ecosys tems they have developed in. other ecosystems at vertical prole. Fig. 4: Cartographic overview of main types of vegetation in area of mountain Prenj. S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 539 Basic determination of this vegetation is carbon ate geological surface (limestone, dolomites and dolo mitic limestone) at the whole vertical prole, with very distinguished inclination of terrain (between 50), sometimes even 120 (in recess of rocks), extremely shal low soil kalkolithosol, and on shattering rocks kalko regosol. ese terrains are very shallow (between sev eral millimetres and several centimetres). ey are rich in mechanical composition, while they are extremely poor in nutritive (organic) substances. Eco-climate is very variable, from somewhat altered Mediterranean and sub-Mediterranean in canyon of Neretva river to mountain climate in Alpine belt of Prenj, vrsnica and abulja. Extremely high variability of basic eco-climate parameters temperature, relative air humidity and eco logically available water, as well as nutritive poorness, are one of the main factors that determined specic tem plates of speciation (development of species), endemicgenesis (development of endemic plants) and syngenesis (development of plant communities). e best example of specic development patterns for endemic plants on bare karst represent rock crevices which was the reason why the development endemic centre was created right in this area. W ithin the ecosystem in cracks of calcareous rocks, at this area, determined are 30 communities at level of as sociation, 10 on level of alliances, 5 on level of order, and they all belong to the class Asplenitea trichomanis Br.-Bl. 1934 corr. Oberd. 1977. All communities in canyon of Neretva river and its conuents are endemic and Ter tiary relict character, while some are of glacial relict character in sub-Alpine and Alpine area such as commu nities of alliance Amphoricarpion autariati Lakui 1968, Amphoricarpion neumayeri Lakui 1968 M icromerion croaticae Horvat 1931 and Potenillion caulescentis Br.Bl. 1926). Communities of this vegetation contain more than 200 dierent taxa, out of which most are of endemic and steno-endemic character. Especially rich in endemic species is community of H eliospermo retzdorani-Oreo herzogietum illyricae ili 1970. Some of them are exclu sively connected to endemic centre ( Campanula herce govina Degen, Salix glabra Scop. var. deylii Rhl, Silene retzdornana (K.Mal) H.Neumayer (Syn.: H eliosperma retzdornana K.Mal), Rhamnus illyrica Gris. ap. Pant. Sibiraea croatica Degen (Syn.: Sibiraea altaiensis (Laxm.) C.K.Schneid. var. croatica (Degen) Beck/ Potentilla speciosa W illd. and others). In syntaxonomical sense, this class is dierentiated into fol lowing orders, alliances and associations. Order: Poten tilletalia caulescentis Br.-Bl. 1926 include alliances of Po tentillion caulescentis Br.-Bl. 1926 and community of Po tentilletum caulescentis (Br.Bl. 1926) Aich. 1933, in the coldest places of north facing slopes in the alpine belt, and shaded carbonate overhangs. is order includes commu nities that continue on alpine communities of carbonate rock crevices of the north Fig. 5: Cartographic overview of main types of vegetation in area of mountains vrsnica and a bulja. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 540 western Dinaric Alps belonging to alliance M icromerion croaticae Horvat 1931which is dierentiated into several endemo-relict communites, such as: Potentilletum clusia nae Horvat 1931, Asplenietum ssi Horvat 1931 Asperulo hercegovinae-Potentilletum appeninae*, Arenario gracilisCampanuletum hercegovinae*, Primulo-Edraianthetum serpyllifoliae*, Primulo-Campanuletum hercegovinae*, Asplenio ssi-Potentilletum apenninanae and Edraiantho serpyllifolii-Potentilletum clusianae* Vegetation in crack rocks of shaded places is encompassed by the alpine al liance Cystopteridion Richard 1972 and M oehringion muscosae Horvat & Horvati 1951, whose most signi cant communities in hilly and upland belt is M oehringioCorydaletum leiospermae* while in the sub-alpine belt occur endemic communitiy Cardamino-Campanuletum cochlearifoliae* Due to strong impact of maritime climate which af fects the southern and southeastern slopes of this moun tain complex, in the rock crevices occur also communi ties that characterize the vegetation of carbonate rock ssures of the litoral southeastern Dinaric Alps. In re spect to vegetation, these communities are encompassed by the endemic order M oltkietalia petraeae Lakui 1968 with two alliances: Centaureo-Campanulion Horvati 1934 in the hotest habitats of both supra-Mediterranean and hilly belt, and Edraianthion tenuifolii Lakui 1968 in warm, dolomite habitats of continental area. e al liance Centaureo-Campanulion Horvati 1934 is very diversied in this area. It includes several endemo-relict communities, of which the most important are: I nulo verbascifoliae-M oltkietum petraeae*, Asplenio-Cotyledon etum horizontalis Horvati 1963 Cephalario leucanthaeI nuletum verbascifoliae*, Achnathero-M oltkaeetum pe traeae*, M icromerio thymifoliae-I nuletum verbascifoliae* and Edraiantho-Seslerietum interruptae Redi 1990 e alliance Edraianthion tenuifolii Lakui 1968 includes the highest positions of endemo-relict communities with Moltkia within its distribution range (around 1,700 m above sea level Boranica on Prenj mountain). e most important communities are: Potentilla speciosaM oltkia petraea Horvat 1941 and Centaureo triumfettiM oltkietum petraeae* e vegetation of rock cracks in the sub-alpine and alpine belt in major share of the investigated karst area belongs to south-dinaric order Amphoricarpetalia Lakui 1968, which is being dierentiated in two alli ances: Amphoricarpion autariati Lakui 1968 in the can yon of Neretva river with following communities: Helio spermo retzdorani-Oreoherzogietum illyricae ili 1970 and M icromerio croaticae-Potentilletum persicinae* e alliance Amphoricarpion neumayeri Lakui 1968 en compasses communities of limestone and dolomite rock crevices in the zone of endemic white-barked pine. e most signicant are: Amphoricarpi-Pinetum leucodermis Fukarek 1966, M oltkio-Pinetum heldreichii* Arenario gracili-M oltkietum petraeae and Asperulo hercegovinaePotentilletum persicinae*. High value to the area is provided also by small fragments of communities which are characteristic for shaded mediterranean carbonate rocks belonging to or der Anomodonto-Polypodietalia O. de Bols & J. Vives 1957 and alliance Polypodion serrati Br.-Bl. (1931) 1947. Vegetation of crevices of dolomitized limestone rocks in the zone of white-barked pine belongs to order Potentil letalia speciosae Quzel 1964 and separated endemic al liance M oltkio-Potentillion speciosae with two alliances: Edraiantho-Potentilletum speciosae* and M olkaeo-Poten tilletum speciosae*. ese ecosystems are very ane to ecosystems of shaded and wet rocks of Mediterranean and sub-Med iterranean belt of class Adiantetea Br.-Bl. 1947, order Adiantetalia Br.-Bl. 1931 and alliance Adiantion capilliveneris Br.-Bl. 1931, which is developed in fragments on recesses in canyon of river Neretva that is under in uence of Mediterranean climate. ey spread on very small surfaces of this area, but they have huge ecological and biological meaning. Vegetation in ecosystems of on carbonate screes ( laspietea rotundifolii Br.-Bl. 1948) Same as ecosystems of cracks in the rocks, car bonate screes or grinders are azonal in character. ey are developed in bases of mountain massive, and all the way to peaks of Prenj, vrsnica and abulja.ey are especially impressive in sub-mountain belt of the whole endemic centre where they form magnicent communities of unique oristic content, appearance, origin and role in system of creation of new types of vegetation, or in system of syngenesis. Main features of these ecosystems are given by vegetation that is adjusted to extreme ecological conditions, constant movement of geological surface, very shallow soils (sirozem), then colluviums in bases, very variable ecoclimate characteristics (specially temperature, relative air humidity and water available to plants). Plants that are part of construction of vegetation of rock creeps have adjusted to these conditions during evolution, so they have developed very deep and oen very branched system of roots, pillow-like bush forms and relatively small surface biomass. Due to extreme and unique of climate forms, geological surface, lack of organic sub stances and other factors of biotope, specic forms of speciation have developed that are characterized by unique endemic genesis. at resulted in development of endemics, relicts and plant species with narrow geo graphical spreading. S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 541 In this vegetation type 30 associations, 7 alliances, included in 4 orders that belong to wider spread class are determined. is class includes south-eastern-dinaric order: Arabidetalia avescentis Lakui 1968 that makes syngenetic connection with the alpine order laspieta lia rotundifolii Br.-Bl. 1926, alliance Corydalion ochro leuceae Lakui 1975 and community M icromerio thy mifoliae-Corydaletum ochroleuceae on screes of upland and hilly belt in the canyon of Neretva river. e main share of this vegetation order is to be found within al pine and sub-alpine screes of north facing slopes of Prenj and vrsnica mountain All vegetation units are of endemic-relict character. us, endemic alliance Saxi fragion prenjae Lakui 1968 is dierentiated in several communities which inhabit screes that occur at sinkhole slopes, of which the most important are: Saxifrago-Pa paveretum kerneri Lakui 1968, V iolo bioraeSaxifrag etum prenjae*, Aubrietum croatiace Horvat 1931, BunioSaxifragetum prenjae*, Seslerio robustae-Scutellarietum alpinae*, Cystopteri-Aquilegietum dinaricae*, Saxifrago prenjae-Adenostyletum alliariacae*, D oronico-Adenostyl etum kerneri* and Papavero kerneri-D oronicetum which is continuation of communities around snow beds. On screes made of ne comminute carbonate rocks com munities Bunio-I beretum carnosae Horvat 1931 and Eu phorbio-V alerianetum bertisceae Lakui 1968 from the alliance Bunion alpini Lakui 1968 occur On the entire vertical prole (from Neretva river to mountain peaks, especially those of abulja mountain) occur screes that are aected by Mediterranean climate and hence belong to order D rypeetalia spinosae Quzel 1967 with alliance Peltarion alliaceae Horvati (1956) 1958, in which high richness of both species and com munities was identied. e greatest signicance in terms of phytogeography have communities as follows: D rypeetum jacquinianae Horvati 1934 G eranio-An thriscetum fumarioidis Horvati 1963 T eucrio arduiniPeucedanetum*, M icromerio thymifoliae-G eranietum macrorrhizi*, Euphorbio-Saturejetum montanae* and Corydalo-Epilobietum rosmarinifoliae. Dry alpine and sub-alpine screes are encompassed by the alliance Silenion marginatae Lakui 1968 that includes following endemo-relict communities: D rypidiSilenetum marginatae Lakui 1968 G eranio-H eracleet um balcanicum Lakui 1968 D rypeetum linneanae Hor vat 1931 D rypidi-H eracleetum orsinii*, D ryopteridetum villarsi (Jenny-Lips 1930) Horvat 1931 Seslerio robustaePetasitetum kablikiani* and Senecio visiniani-H eracleet um orsinii on limestone blocks of southern Prenj moun tain slopes. Screes along with rock crevices provide this area with unique forms of overall biodiversity. Following en demic species found refuge in this area: Saxifraga prenja Beck Papaver kerneri Hayek H eracleum orsinii Guss. T eucrium arduini L. and others. Ecosystems of screes that occur in dry bottoms of sinking creeks are signicantly dierent both in their structure and genesis from mountain screes. Especially important biotopes are basins of many ooding riv ers, sinking creeks, and conuents of Neretva that sink during vegetation period. At these places, where main biotope attribute is represented by pebbles, endemic communities of order Epilobietalia Fleischeri Moor 1958 occur. is order includes the alliance Salicion in canae Aichinger 1933 with endemic-relict communi ties: Petterio-Salicetum incanae Redi et al. 1992/94 (Muratspahi et al. 1991; Redi et al. 1992), Ostryo-Salicetum incanae*, M olinio litoralis-Petasitetum kablikiani* and Salvio oncinali-Salicetum incanae* It should be stressed that such habitats are under inu ence of surface water ow during wet and cold period of a year. Scree communities on stone blocks in the su pra-Mediterranean and hilly belt are encompassed by the Stipetum calamagrostis Br.-Bl. 1918 from the alliance Stipion calamagrostis Jenny-Lips 1930 ( Achnatherion cal amagrostis Jenny-Lips 1930), belonging to separate order G alio-Parietarietalia Bocaiu et al. 1966. Many of them are of endemic character. Representative communities of that kind are to be found in valleys of Grabovica and Drenica river. Vegetation around snow beds on carbonate sub strates ( Salicetea herbaceae Br.-Bl.1947) At many places in Alpine and sub-Alpine belt of these mountains, such as sinkholes, karst valleys, mild elevations of terrain, areas that are lee warded from strong mountain winds, snow remains for long time, while at some places it remains during a whole year. In such manner, special biotopes were formed in proximity of glaciers, inhabited by mostly glacial relicts species of plants and animals, that, at time, developed a num ber of unique, mostly endemic-relict communities. At this area, vegetation in proximity of glaciers is developed on shallow organic mountain humus, which although laying on carbon geological surface, is acidied due to slow decomposition of humus substance. Besides plants, that are exclusively connected with glaciers, some spe cies that belong to neighbouring vegetation types, such as alpine and sub-alpine pastures, are also found. Due to extreme ecological circumstances, biomass is lowest in these ecosystems. Despite that, they are extraordinary important in total image of biodiversity, not only in this area, but also in biodiversity of north hemisphere as a whole. In addition, ecosystems in proximity of glaciers are the best indicators of intensity and nature of climate alterations. Vegetation around snow beds on Prenj and VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 542 vrsnica mountains. is encompassed by orders Saliceta lia retusae-serpyllifoliae Lakui et al 1979 and alliance Salicion retusae Horvat 1949 with following glacial relict communities: Soldanelo-Salicetum retusae Horvat 1933 in Horvat, Glavac & Ellenberg 1974 Polygono vivipariSalicetum retusae*, G entiano tergestinae-Salicetum retu sae*, D ryadeto-Salicetum retusae* and T rifolio-Salicetum retusae *. Most of communities of this vegetation are of gla cial relict origin, and they have important role in as sessment of age of mountain peaks and mountain as a whole. In current conditions they have an irreplaceable role in assessment of trends in area of global changes of living conditions, especially climate. ese are biotopes of signicant number of glacial species, such as: Soldanella alpine L. T rifolium noricum W ulfen Polygonum viviparum L. G entiana tergestina Beck Salix retusa L. S. serpyllifolia Scop. V iola zoysii W ulfen and others. Vegetation in ecosystems of alpine and sub alpine carbonate pastures and high mountain tundra ( ElynoSeslerietea Br.-Bl. 1948) Above highest level of forest vegetation on Prenj, vrsnica and abulja mountains there is area of high mountain pastures. at is specic form of high moun tain tundra that is in a wider geographical area character ized by relative poorness of ora and small production of biomass. However, at the area of those mountains, this vegetation is very rich in species, as well as in commu nities. It is developed above the belt of juniper of pine woods on carbonate surface of alliance Pinion mugi Pawl. 1928, and sometimes above sub-alpine forest beech ( Fag etum "subalpinum" Horvat 1938) and of W hite bark pine Pinetum heldreichii "subalpinum" In alpine belt, moun tain pastures are of primary character. Besides, they are developed in sub-alpine belt in an area where the prima ry vegetation is altered, and they are of secondary char acter here. Considering a signicant area of mountains at this region, pastures are very present here. ey are specic centres of their spreading and richness compared to the whole Dinarides. Since the Dinarids are mountains built of carbonate rocks, specic communities have developed here and in syntaxonomical sense they are included in separate class of Elyno-Seslerietea Br.-Bl. 1948. is class reaches extremely high diversity level on Prenj, vrsnica and abulja mountains. It is dierentiated into 30 most ly endemic and glacial relict communities, which are en compassed by six alliances and three orders. e coldest mountain habitats of alpine pastures are included into dinaric-apennine order Seslerietalia tenuifoliae Horvat 1930 with two alliances: Seslerion tenuifoliae Horvat 1930 that encompasses communities of the highest peaks ex posed to the most devastating mountain winds, such as: L aeveto-H elianthemetum alpestris Horvat 1930, Festuca paniiana-D ianthus brevicalyx Horvat 1934, EdraianthoD ryadetum octopetalae Lakui 1968, Carici laevi-Fes tucetum bosniacae* and Seslerio tenuifoliae-Arctostaphyl letum uvae-ursi Redi et al 1984. Plant communities that inhabit leeward places and deeper calcomelanosol are included in the alliance Festucion bosniacae Horvat 1930 which is constituted of several in oristic respect quite diverse communities: Festucetum bosniacae (Syn.: Festucetum pungentis Horvat 1930 ). Stachydi-Festucetum bosniacae*, L ilio bosniacae-Festucetum bosniacae*, Av enulo blavii-Festucetum bosniacae* and G entiano sym phyandrae-Festucetum bosniacae Lakui 1975 In the southeast, peaks of these mountains are cov ered by vegetation belonging to the southeast-dinaric, high-mountainous order Crepidetalia dinaricae Lakui 1966 e endemic alliance Oxytropidion urumovii Lakui 1964 (Syn.: Oxytropidion dinaricae Lakui 1966) connects this area in phytogeographic sense with the southeastern Dinaric Alps. It includes several endemic communities, such as: Seslerietum tenuifoliae "hercegovi num" Lakui 1969 Asperulo-Festucetum pancicianae Lakui et al 1973 Edraiantho-V eronicetum satureoidis Lakui et al 1973 Elyno-Edraianthetum serpyllifolii Lakui 1968 and Alchemillo-Scabiosetum silenifoliae on vrsnica and Prenj mountains. e highest peaks of vrsnica and Prenj mountains, places that are exposed to the wind, are inhabited by steno-endemic communities of alpine grasslands from the alliance Oxytropidion pren jae which encompasses communities that cover smaller areas, such as: M inuartio handelii-Caricetum pollicensis, Oxytropidi-D ryadetum octopetalae*, Potentillo clusianaeSaxifragetum marginatae*, Potentillo apenninae-Sesleri etum tenuifoliae* and Potentilletum clusiano-apenninae*. Most exposed habitats from littoral side of these mountains are inhabited by a form of eolian vegetation of sub-alpine grasslands of order Edraiantho-Seslerietalia robustae Redi 2003 is alti-montane vegetation or der here includes the alliance Seslerion robustae (Lakui et al. 1982) Redi 2003 (Syn.: Seslerion nitidae Horvat 1930) with the following communities: Seslerietum ro bustae (Syn.: Seslerietum nitidae Lakui et al 1975), Fes tuco bosniacae-Seslerietum robustae*, Carici-Seslerietum robustae, Avenastro blavi-Seslerietum robustae*, Astran tio carniolicae-Seslerietum robustae* Saturejo subspi catae-Seslerietum robustae and Antherico-Seslerietum robustae* In the driest habitats live communities from the alliance Seslerio-Edraianthion pumilii : Carici laeviH elianthemetum balcanici Horvat 1930 ArenarioSes lerietum tenuifoliae* and G enisto dalmaticae-Seslerietum S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 543 tenuifoliae which makes ecological continuity with the vegetation of supra-Mediterranean rocky grasslands to ward lower positions. About 800 taxa of vascular plants participate in the community of this vegetation, out of which many are of endemic relict character. Some of them are exclusively connected to this area, and therefore they are specic steno-endemics. Mountain and sub-mountain meadows are developed on all expositions and elevations of terrain up to 50 degrees, where they are changed into commu nities of either screes or rock crevices. Communities on the peaks are highly exposed to winds, and so the winds are dominant climate factor in their shaping, structur ing, dynamics and syngenesis. ose communities be long to orders of Seslerietalia tenuifoliae Horvat 1930 and Crepidetalia dinaricae Lakui 1966. At shaded biotopes are sub-mountain meadows with Bosnian fescue grass Festucion bosniacae Horvat 1930 corr. Redi 2007, and at some areas with communities with robust sedge Sesle rion robustae (Lakui et al. 1982) Redi 2003. e highest positions and in sense of eco-climate most rigid areas are inhabited by high mountain vegeta tion, that is by its physiognomy is very similar to moun tain meadows, in syntaxonomical sense belong to the class Carici rupestris-K obresietea bellardi Ohba 1974 ese communities structure ecosystems of alpine and sub-alpine pastures and dwarf shrubs. In syntaxonomi cal sense, on researched area this class is represented by one association, of alliance Oxytropido-Elynion Br.-Bl. 1949 and order Oxytropido-Elynetalia Oberdorfer ex Al brecht 1969 Vegetation of thermophyllous grasslands ( Festu co-Brometea Br.-Bl. & R.Tx. in Br.-Bl. 1943) In hill and upland belt, at biotopes of former forests and bushes, currently dierent thermophyllous commu nities of meadows and rocky grasslands are developed. Soils are shallow, mostly kalkomelanosol and rendzine, rarely kalkokambisol. Meadows are very rich in species, among which some are endemic, even of relict character. In syntaxonomical sense, they belong to the order Bro metalia erecti (W Koch 1926) Br.-Bl. 1936, alliance Bro mion erecti Koch 1926, which on the researched area is represented by two communities: Bromo-Plantaginetum mediae Horvat (1931) 1949 and Bromo-D anthonietum alpinae ugar 1972 Vegetation of Mediterranean and supra Medi terranean rocky grasslands and meadows ( eroBrachypodietea ramosi Br.-Bl. 1947; Syn.: CymbopogoBrachypodietea p.p.) Going toward shallower soils and warmer biotopes, in a zone of thermophyllous forests and thicket of orders Quercetalia pubescentis Br.-Bl. (1931) 1932 and OstryoCarpinetalia orientalis Lakui, Pavlovi & Redi 1982 thermophyllous communities of Mediterranean and subMediterranean bullheads and meadows are developed. ese communities are very rich in species (more than 500 taxa of higher plants are in their content), among which many are of endemic character. Syntaxonomically, they are very complex and dierentiated in one order, 6 alliances and 11 communities, while they are unied into one class. ey are encompassed by the supra-Med iterranean order Scorzoneretalia villosae Horvati 1975 that dierentiates into four alliances. At mild slopes, fre quently in larger sinkholes, on deeper carbonate soils live communities of supra-Mediterranean meadows from the alliance Scorzonerion villosae Horvati 1949, especially on abulja and Prenj mountains, such as: D anthonioScorzoneretum vilosae Horvat & Horvati (1956) 1958 and L ilio cathaniae-Agrostetum capillaris*. On more shallow and rocky carbonate soil, at more sloped and to the wind exposed places, the communi ties of rocky grasslands occur. e highest positions are taken by the alliance Chrysopogoni-Satureion Horvat & Horvati 1934 with communities: Scabioso-G lobulari etum Saturejo-Edraianthetum Horvat 1942 and StipoSalvietum oncinalis Horvati (1956) 1958. Going upwards on a vertical prole toward subalpine belt, this rocky grassland are continued by com munities from the alliance Satureion subspicatae Horvat 1959 ( Carici-Centauretum rupestris Horvat 1931 and Sal vio bertoloni-Saturejetum subspicatae *) that are ecologi cally connected with communities of sub-alpine grass lands. In warm dolomite habitats communities from the alliance Festucion pseudovinae Soo 1933 ( K oelerio-Fes tucetum pseudovinae and G entiano cruciataeFestucetum pseudovinae *) occur, as well as the communities Alys setum moelendorani Riter-Studnika (1956) em. 1967 and Reichardio macrophyllae-ymetum aureopunctati* that are inevitably associated with the endemic alliance on dolomite substratum Peucedanion neumayeri RiterStudnika 1967. At habitats that are under high inuence of wind, on very shallow and rocky terrains, ecosystems of pioneer vegetation of rocky grasslands are occasionally present, with class K elerio-Corynephoretea Klika in Klika & No vak 1941, order Alysso-Sedetalia Tx. 1951 and alliance Alysso alyssoidis-Sedion albi Oberdorfer & Mller 1961. Herbal tall thermophyllous vegetation ( TrifolioGeranietea Mller 1962) In the system of progradation, communities of bull heads and thermophyllous meadows have a tendency of traversing to communities of low forest and thermo phyllous thickets. In certain development stages, they VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 544 transfer into very dynamic type of vegetation that con tain species of both meadows and forests habitats. at is how ecotones are formed currently very important habitats where intensive processes of competition and ecological co-existence are in progress. ese habitats and their vegetation are of great importance in evalu ation of conditions and potential possibilities, as well as trends of given type of vegetation to original climax community. Vegetation belongs to the class of T rifolioG eranietea Mller 1962, order Origanetalia Mller 1962 with two alliances G eranion sanguniei R. Tx. in Mller 1962 and T rifolion medii Mller 1962. is vegetation is still poorly researched in those conditions. Vegetation of hygrophyllous and mesophyllous grasslands ( Molinio-Arrhenatheretea R. Tx. 1937) Ecosystems of hygrophyllous and temperate humid meadows on the investigated area are spread on small areas, since surface water is either not present or it is poorly available in these mountains. is type of vegeta tion is mostly developed on deep soils. ose are brown limestone soils kalkokambisol, meadows luvisol, and eugley and pseudogley, on at terrains where the level of ground water is low. On a vertical prole they are de veloped from bases of mountains to sub-mountain areas. In syntaxonomical sense they are very complex and de pending on hydro-thermic regime and oristic composi tion thery are dierentiated into 16 associations, 13 al liances, 7 orders, while they are all unied in the class M olinio-Arrhenatheretea R. Tx. 1937. e deepest soil of both hilly and upland belt is inhabited by mesophyllous grasslands from the order Arrhenatheretalia Pawlowski 1928, alliance Arrhenatherion elatioris Br.-Bl. 1925, which are communities Arrhenatheretum elatioris R. Tx. 1937 ononidetosum and Festucetum pratensis Lakui et al 1975, then alliance Festuco-Agrostion capillaris Redi 1990 with community Festuco-Agrostetum capil laris (Horvat 1951) 1962 em. Trinajsti 1972 and alliance Cynosurion cristati Tx. 1947 with community CynosuroFestucetum rubrae Redi 1990. e grasslands of upland and sub-alpine belt are included into the endemic dinar ic alliance Pancicion serbicae Lakui 1966, of which on Prenj mountain identied are following communities: L ilio bosniacae-Crepidetum conyzifoliae*, K nautio-Fes tucetum nigraescentis* and Onobrychydi-Festucetum ni graescentis *. On humid type of soil and gently sloped ter rain, which is oen the case at Blidinje and Borako lake, there is an occurance of hygrophyllous grassland com munities from the order H oloschoenetalia Br.-Bl. (1931) 1947 and fragments of the community D eschampsietum mediae Br.-Bl. 1931 belonging to the alliance D eschamp sion mediae Br.-Bl. (1947) in Br.-Bl. et al 1952. Similar kind of habitats are covered by hygro-mesic meadows of poljes (at Blidinje lake) from the order T rifolio-H ordeeta lia Horvati 1960 and alliance M olinio-H ordeion secalini Horvati 1934 with following communities: Plantagine tum altissimae Riter-Studnika 1954 D eschampsio-Plan taginetum altissimae Ilijani 1974 and Scillo litardiereiRanunculetum *. Around upland springs we can nd some fragments of swampy meadows from the order D eschampsietalia Horvati (1956) 1958, alliance D es champsion caespitosae Horvati 1930 and D eschampsi etum caespitosae Horvati 1930. e most humid types of soil, which are on the way to get transformed into bogs, are inhabited by hygrophylous meadows from the order M olinietalia W Koch 1926, alliance M olinion coer uleae W Koch 1926 (communities: G ladiolo-M olinietum coeruleae Horvat 1962, M olinietum coeruleae W Koch 1926 and Seslerio uliginosae-Caricetum paniceae *), then the alliance Calthion palustris Tx. 1937 and community Filipenduletum ulmariae from the alliance Filipendulion (= Filipendulo-Petasition Br.-Bl. 1947). e vegetation along forest roads and on clearings in woods belongs to wider distributed order Plantagini-Prunelletalia Ell mauer & Mucina 1993 and alliance Plantagini-Prunellion Eli 1980, while on trampled places vegetation from the alliance Potentillion anserinae R.Tx. 1947 encompassed by the order Potentillo-Polygonetalia R. Tx. 1947 occur. Vegetation of swamp and water ecosystems Even though the whole area of endemic centre of Prenj, vrsnica and abulja mountains is built of perme able carbonate rocks, occasionally there are certain forms of water-resistant surfaces and soils with developed ped ological prole that in most part is moist throughout a year. is vegetation on researched areas is developed along the coastline of Blidinje lake on Dugo polje below vrsnica, especially along Borako lake in foothills of Prenj, along glacial Crljensko lake on Vilinac (vrsnica), along smaller ows (Masna Luka on vrsnica), and par tially along Neretva river and its conuents. Even though this vegetation is developed on small surfaces, it has a great ecological meaning and makes this area very diver sied and rich. In such ecological circumstances dier ent types of communities exist, that create a productive component of several ecosystems. Ecosystems of marshes with reedbeds occur in a form of smaller fragments which develop along standing water and mountain lakes. It is encompassed by the class Phragmiteto-M ag nocaricetea Klika in Klika & Novak 1941 (Syn.: Phrag mitetea Tx. & Prsg. 1942 p.p.), order Nasturtio-G lyceri etalia Pignatti 1953, and communities from the alliance Sparganio-G lycerion Br.-Bl. & Siss. in Boer. 1942. e reedbed community Scirpo-Phragmitetum australis W S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 545 Koch 1926 from the alliance Phragmition communis W Koch 1926, and order Phragmitetalia W Koch 1926 oc curs on alkaline bogs at Borako lake. Going toward more terrestric soil this community makes ecological continuity with sedge communities Caricetum elatae W Koch 1926 and Caricetum gracilis R.Tx. 1937 from the alliance M agnocaricion elatae W Koch 1926. e inshore belt, in which organic biomass achieves a high rate of turnover, comprises fragments of hygrophylous commu nities from the alliance Oenanthion aquaticae Hejn ex Neuhusl 1959. Especially indicative is the presence of community Scheuzherio-Caricetea fuscae (Norh.1936) Tx. 1937 in Masna Luka below vrsnica mountain, as well as dif ferent communities with reed, sedge and other aquatic plants.is class includes the vegetation of blanket bogs from the order Caricetalia davallianae Br.-Bl. 1949, which characterizes the alliance Caricion davallianae Klika 1934 of boreal distribution. ough covering a smaller area, it is represented here by three communi ties: Eriophoro-Caricetum paniceae Horvat 1962, Ve ronico beccabungae-Eriophoretum latifoliae and M entho longifolii-Eriophoretum latifoliae *. In transitional zone of bogs occur fragments of boreal communities Carice tum fuscae Br.-Bl. 1915 and Eleocharetum ovatae Hayek 1923 from the alliance Caricion fuscae W Koch 1926 em. Klika 1934 (Syn.: Caricion canescentis-fuscae ) and order Caricetalia fuscae W Koch 1926 em. Br.-Bl. 1949. In this area, the inshore belt of mountain lakes, as well as the estuaries of their auents, are covered by aquatic vegetation that belongs to the widely spread class Potamogetonetea Klika in Klika & Novk 1941 (Syn.: Potametea R.Tx. & Prsg. 1942). e order CalitrichoPotametalia W Koch 1926 with the alliance Ranunculion uitantis Neuhusl 1959 comprises oligodominant com munities of clear brooks, which occur mainly around Borako lake. e aquatic communites of mountain lakes belong to the order Potamogetonetalia W Koch 1926 (Syn.: Potametalia W Koch 1926) and are charac terized by three ecologically dierent community types: (i) submerged vegetation from the alliance Magnop otamion (Syn.: Potamion eurosibiricum W Koch 1926) with Nasturtio-Beruletum angustifoliae submersae RiterStudnika 1972 and M yriophyllo-Nupharetum W Koch 1926, while in warmer litoral waters communities from the alliance Ceratophyllion demersi So 1927 occur. e oating vegetation, which is composed of plants that are not rooted at the bottom, is comprised by nenuphar communities from the alliance Nymphaeion albae Ober dorfer 1957 e freshwater vegetation in the litoral belt is of patchy character. It is encompassed by the class I setoNanojuncetea Br.-Bl. & Tx. 1943 and its order I soetetalia Br.-Bl. 1931, which is alliance Nanocyperion avescentis W Koch 1926 and Fimbrystilion dichotomae Horvati 1954. In so-called "dead", standing water, of euents that surround the Borako lake, developed communities of oating unrooted vascular plants from the cosmopolite class L emnetea W Koch & Tx.1954, order L emnetalia W Koch & Tx.1954 and alliance L emnion W Koch & Tx.1954. Similar kind of habitats are inhabited by com munities of cryptogammes with green algae from the class Charetea fragilis Fukarek 1961 ex Krausch 1964, or der Charetalia hispidae Sauer ex Krausch 1964 and alli ances Charion fragilis Krausch 1964 em. W Krause 1969 and Charion vulgaris ( Krause ex Krause & Lang 1977) Krause 1981 e vegetation that develops around moun tain springs and smaller streams is of glacial character. It belongs to the class M ontio-Cardaminetea Br.-Bl. & Tx. ex Klika 1948 order M otio-Cardaminetalia Pawlowski in Pawlowski, Sokolowski & W allisch 1928 and alliances Cardamino-M ontion Br.-Bl. 1926 em. Zechmeister 1993, while on tu the alliance Cratoneurion commutati W Koch 1928 occurs Vegetation of mesophyllous tall herb commu nities ( Mulgedio-Aconitetea Hada & Klika in Klika 1948) Same as communities of class T rifolio-G eranietea Mller 1962 communities of tall herbs are developed in the ecotones areas. However, this vegetation develops in the zone of mesophyllous forests and bushes. Soils are mildly nitried kalkokambisol or deeper kalkomelanosol in the zone of mesophyllous forests of beech, spruce and r, as well as spruce and white-barked pine. is vegeta tion was previously developed at the bottoms of sink holes where snow remains longer. is vegetation type belongs to the order Adenostyletalia Br.-Bl. 1931 and in cludes several plant communites from the Alpine-dinar ic alliances: Adenostylion alliariae Br.-Bl. 1925, Petasition dreri Lakui 1968 and K nautio-V eratrion albae *. Vegetation of tall herb communities on nitried soil that develops next to mountain cottages and sheepfolds forms a special dinaric community Senecio rupestrisChenopodietum boni-henrici* from the alliance Rumi cion alpini Klika & Hada 1944, order Rumicetalia alpini Mucina in Karner et Mucina 1993. In a development sense this vegetation is ecological connection between mesophyllous woods and mesophyl lous grasslands. A special value in this type of vegetation are communities of endemic alliances Petasition dreri Lakui 1968 and V eratrion that connect tall herb com munities with sub-alpine meadows of the endemic alli ance Pancicion Lakui 1966. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 546 Vegetation of ecosystems of mesophyllous broa dleaveddeciduous forests and thickets ( Querco-Fag etea Br.-Bl. & Vlieger in Vlieger 1937) Deciduous forests and thickets, on the investigated area are spread on wide belt from lowest parts (around 250 m above see) all the way up to sub-alpine belt (1,600 m and 1,800 m) where they end at sub-alpine low for ests of beech ( Fagetum "subalpinum" Horvat 1949). De pending on hydro-thermic regime they are dierentiated into thermic and semi-humid ones. Terrains where they develop are rendzine and kalkomelanosol, as well as ka lkokambisol, on attened terrain, rarely luvisol. W ithin these ecosystems, a special place is given to dierent beech forests that are developed in hill, hill-Mediterra nean, upland and sub-Alpine belts. Content of this veg etation includes 20 associations, 8 alliances and 3 orders. e deciduous mesophyllous forests in dependence of hydric regime and oristic composition, as well as of soil depth, are dierentiated in three orders: (i) Betulo pendulo-Populetalia tremulae Rivas-Martnez & Costa (progradation stages with trembling poplar and hazel from the alliance Corylo-Populion tremulae Br.-Bl. 1961 and community CapreetoPopuletum tremulae Glii (1950)1975; (ii) Fagetalia sylvaticae Pawlowski in Paw lowski, Sokolowski & W alisch 1928 (beech woods) and (iii) Rhamnetalia fallacis Fukarek 1969 (scrubs of Rham nus L.on limestone blocks in the upland belt with the community Cynancho-Rhamnetum Fukarek 1969 from the alliance L onicero-Rhamnion Fukarek 1969 In the lowest part of the area (upper and mid ow of Neretva river), the order Fagetalia Pawlowski in Pawlowski, Sokolowski & W alisch 1928 is represented by communities of hornbeam and sessile oak QuercoCarpinetum betuli (Horvat 1938) em. Blei 1958 from the alliance Carpinion betuli Oberdorfer 1953, while in warmer habitats it alternates with the illyrian oak-horn beam woods from the alliance Erythronio-Carpinion (Horvat 1958) Marinek in W allnfer et al. 1993. Above this vegetation belt develops the belt of beech woods which is dierentiated into four alliances: (i) illyrian beech forest Aremonio-Fagion Trk et al. 1989 (Syn.: Fagion illyricum p.p.) in cold habitats, oen on north fac ing sloes and deep karstic soil; (ii) moesian beech forests Fagion moesiacae Blei & Lakui 1970, on extremely karstic terrain and shallow soil of AC prole, and on south facing slopes; (iii) xeric beech forests on dolomite Cephalanthero-Fagion Txen 1955 and (iv) forests of beech and hope hornbeam Ostryo-Fagenion Borhidi ex So 1964, on limestone blocks and steep slopes with very shallow soil types. e illyrian beech woods are highly diverse. In the investigated area, they are dierentiated into several com munities: Fagetum "montanum illyricum" Fukarek & Stefanovi 1958, Abieti-Fagetum "illyricum" Horvat 1958 Aceri-Fagetum "subalpinum" Fukarek & Stefanovi 1958 Seslerio -Fagetum Moor 1952 Aceri obtusati-Fagetum Fabijani, Fukarek & Stefanovi ex Fukarek, Stefanovi & Fabijani 1967 and Aceri-T ilietum "mixtum" Stefanovi 1979 (polydominant community in the canyon of Neretva river and its larger auents), and typical karst community from vrsnica mountain Fagetum "mediterraneo-monta num" Redi et al 1984 (Redi & Barudanovi 2010). e moesian beech forests Fagion moesiacae Blei & Lakui 1970 comprise the following communities: Seslerio autumnalis-Fagetum moesiacae, Fagetum moesi acae "montanum" Blei & Lakui 1970 Aceri obtusatiFagetum moesiacae Fabijani, Fukarek & Stefanovi 1963 Abieti-Fagetum moesiacae Horvat, Glava & Ellenberg 1974 Aceri-Fagetum moesiacae Jovanovi 1957 Pino hel dreichi-Fagetum moesiacae "mediterraneo-montanum", Pino heldreichii-Fagetum moesiacae "subalpinum" and Rhamno fallacis-Fagetum moesiacae In the upland belt the very representative commu nities are of beech and r Abieti-Fagetum and commu nities of Rhamnus L. from the order Rhamnetalia fallacis Fukarek 1969, developed on limestone blocks and shal low humus layer. A special value is given to thermophyl lous communities of beech and hope hornbeam OstryoFagenion Borhidi ex So 1964, and forests of beech and white-barked pine Fago-Pinetum heldreichii s.lat. Fragments of acidophilus communities of class Quercetea robori-petraeae Br.-Bl. & R.Tx. 1943, order Quercetalia robori-petraeae Tx. 1931 and group Cas taneo-Quercion petraeae So 1962 em. 1971 are found here, within which fragments of chestnut and sessile oak on acid substrates in Jablanica surrounding were deter mined. Vegetation of ecosystems of broadleaved-decidu ous thermophyllous forests and scrubs (Quercetea pu bescentis (Oberdorfer 1948) Doing Kra 1955; Syn.: Querco-Fagetea p.p.) In warmer habitats, on shallow soils rendzine and organic-mineral humus, colluviums soils in bases of mountains, as well as on kalkokambisol on atter ter rains, form lowest areas to upland belt. ere, forests and scrubs of hornbeam, pubescent oak, Turkey oak, Italian oak and hope hornbeam are developed. In the content of class Quercetea pubescentis (Oberdorfer 1948) Doing Kra 1955 in this area 17 communities of association levels, 7 of alliance level and three of order level are de termined. Even though they form a wide belt, the optimum is in the valley of Neretva river and its conuents, where it gives a specic phyto-geographic mark to this area. In ecological typological sense, we dierentiate low forests S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 547 and scrubs of white hornbeam Carpinion orientalis Blei & Lakui 1966, black hornbeam and autumn blue moor grass Seslerio-Ostryetum, within which are endemic and endemic relict communities. Other types are forests of Italian oak Quercion confertae Horvat 1954 and pubes cent oak Ostryo-Quercion pubescentis as well as forests of Turkey oak Quercion cerris Lakui 1976. Deciduous thermophyllous forests and thickets are much degraded today. Despite this, they contain a whole oristic wealth and signicant number of endemic relict species. ese are special kind of habitats for diverse animal species. In respect to their ecology and physiognomy ther mophyllous woods and thickets are dierentiated in three orders: (i) Fraxino orni-Cotinetalia Jakucs 1961 (thicket communities with Christs-orn Paliuretum aculeate Auct) and Pistacio lentisci-Carpinetum orienta lis* (transitional thicket communities between mediter ranean and supra-Mediterranean belt) from the alliance Paliuro-Carpinion orientalis *. ere are well-preserved woods with pubescent, Turkey and Italian oak around Jablanica, upper ow of Neretva river, and slopes of abulja mountain. ey are comprised by the order Quercetalia pubescentis Br.-Bl. (1931) 1932 and several alliances. e communities of Italian oak with hope hornbeam are encompasssed by the alliance Quercion pubescentis-petraeae Br.-Bl. 1931 and communities: Carpino orientalis-Quercetum pubes centis, Ostryo carpinifoliae-Quercetum pubescentis and Querco-Ostryetum carpinifoliae Horvat 1938 W oods of Italian oak in Herzegovina are of relict character. ey belong to the alliance Quercion confer tae Horvat 1954 and most unique association Quercetum farnetto "hercegovinum" Fukarek (1963) 1966 On acid soil, which is hard to be found here (the valley of Doljanka river), occur fragments of acidophyl lous oak woods Quercetum petraeae "montanum" Stef. (1961) 1964 and the alliance Quercion petraeae Zalyomi & Jakucs ex Jakucs 1960. In similar kind of habitats de velop also fragmented woods with Turkey oak, from the alliance Quercion cerris Lakui 1976. e most frequently occuring are low forests, thickets and shrubs with oriental hornbeam and hope hornbeam from the order Ostryo-Carpinetalia orientalis Lakui, Pavlovi & Redi 1982. Shrubs with oriental hornbeam from the alliance Carpinion orientalis Blei & Lakui 1966 achieve the extremely high diversity lev el. ey are dierentiated into following communities: Seslerio autumnalis-Carpinetum orientalis*, Cruciato glabrae-Carpinetum orientalis, Aceri-Carpinetum orien talis Blei & Lakui 1967 M elampyro trichocalycinaeCarpinetum orientalis, Petterio-Carpinetum orientalis, Rusco aculeati-Carpinetum orientalis Blei & Lakui 1967 and Punico-Carpinetum orientalis e relict wood with hope hornbeam and autumn moor grass from the alliance Seslerio-Ostryon Lakui, Pavlovi & Redi 1982 comprises two communities, which are: Seslerio autumnalis-Ostryetum carpinifoliae Horvat & Horvati 1950 and Fraxino orni-Ostryetum carpinifoliae Vegetation in ecosystems of hygrophyllous forests and shrubs Along the water ows of researched area and along the coastline of Blidinje and Borako lakes, as well as on rare water resistant biotopes, in a form of smaller frag ments, isolated area, and dierent forms of hygrophyl lous communities are developed. e vegetation of hy grophyllous shrubs with purple willow, on swampy kind of soil along watercourses, of the class Salicetea purpu reae Moor 1958, order Salicetalia purpureae Moor 1958, comprises the shrub formations with purple willow Sali cion purpureae Lakui 1975 and Salicion triandrae Mller & Grs 1958. In some places fragments of white willow Salicion albae R. Tx. 1955 occur e inshore belt of rivers and streams, which frequently dries out, com prises fragments of communities from the alliance AlnoQuercion roboris Horvat 1938, as well as of Populion al bae Br.-Bl. ex Tchou 1948 and Alnion glutinosae (Malc. 1929) Meier Dr. 1936 alliances. Even though this vegetation is spread on small a territory at the area of endemic development centre, it has great ecological meaning in total image of rich bio logical diversity. Ecosystems of mesophyllous and thermophyllous shrubs ( Rhamno-Prunetea Rivas Goday & Borja Car bonell 1961) At many places, forest and thicket are degraded and today converted into lower forms of organization of eco systems. Most dominant type of this vegetation are com munities of shrubs that are very oen a long lasting stage in development of more complex forms of vegetation. ese are also specic ecotones that are characterized with extraordinary oristic richness. Shrub communi ties of mesophyllous and mesophyllic-thermophyllous character belongs to the order Prunetalia spinosae R. Tx. 1952 and alliances: (i) Berberdion vulgaris Br.-Bl. 1950 with the community Evonymo-Rhamnetum catharti cae *; (ii) Crataego-Corylion Fukarek 1969, shrubs of hazel with following associations: H elleboro-Coryletum avellanae Redi 1990, Crataego-Coryletum avellanae Fukarek 1969 and (iii) Prunion spinosae So 1951 with the community Crataego-Prunetum spinosae Beus 1971. e greatest physiognomic meaning have the hazel communities, alliance Crataego-Corylion Fukarek 1969, barberry Berberidion and blackthorn Prunion spinosae So 1951. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 548 Vegetation in ecosystems of heaths ( LoiseleurioVaccinietea Eggler ex Schubert 1960) On the investigated area heaths are developed on carbonate shallow soil of type of rendzine and humus in the sub-alpine and upland belt. ese are communities of low evergreen shrubs, in which prevail species from genus Juniperus L. Heaths are here encompassed by the order Rhododendro-V accinietalia Br.-Bl. in Br.-Bl. & Jen ny 1926 and the alliance Juniperion nanae Br.-Bl. et al. 1939 in sub-alpine belt, while in upland and hilly belt the alliance Juniperion communis occurs. Distinguished ecological meaning is given to com munities of mountain peaks from the alliance Juniperion nanae Br.-Bl. et al 1939, in upland belt, in the zone of white-barked pine Pinetum heldreichii "mediterraneomontanum" Blei & Lakui 1969 and the community Juniperetum communis-intermediae Stefanovi 1974. Vegetation in ecosystems of relict pine forests ( Erico-Pinetea Horvat 1959) On the investigated area relict pine forests have a great meaning. ey are so dierent in both ecologi cal and in oristic sense compared to other congenital types of vegetation that they belong to a separate phytogeographical province relict pine forests. ey are very common on dolomites and dolomitic limestones. ere fore, they have a role of refuge of Tertiary ora and veg etation. Syntaxonomical, this vegetation is dierentiated into three vegetation orders: (i) Erico-Pinetalia (Ober dorfer 1949) em. Horvat 1959 (relict black pine forests on dolomite) with two alliances: Fraxino orni-Ericion Horvat 1958 and Fraxino orni-Pinion nigrae Zupani 2007 (ii) Pinetalia heldeichii-nigrae Lakui 1972 (for ests of white-barked pine and illyrian black pine) in the upland and sub-alpine of Prenj, vrsnica and abulja mountains and (iii) order Rhododendro hirsuti-Ericetalia carneae Grabherr, Greimler & Mucina 1993 (shrubs of mountain pine with heath on dolomite in both alpine and upland belt) with the relict alliance Pino mugi-Eri cion and endemo-relict community G entiano dinaricaeEricetum carneae identied on Prenj mountain. Vegetation order Pinetalia heldreichii-nigrae Lakui 1972 is dierentiated into the alliance Pinion nigrae Lakui 1972 (forests of illyrian black pine in the can yon of Neretva river and its auents) with relict black pine forests combined with hope hornbeam OstryoPinetum nigrae Trinajsti 1999 and forests of black pine with "omorikoides" spruce Pino nigrae-Piceetum abietis "omorikoides" Lakui & Redi 1989 in the sub-alpine belt of vrsnica mountain. Forests of tertiary relict white-barked pine from the alliance Pinion heldreichii Horvat 1950 are dierentiated into three associations: Pinetum heldreichii "mediteran neo-montanum" Blei & Lakui 1969 Junipero-Pinetum heldreichii (Blei 1960) Fukarek 1970 and Fago-Pinetum heldrechii Jankovi (1972) 1975. Beside the community of Illyrian black pine Pinion nigrae Lakui 1972, special feature to this area is given by forests of endemic Balkan white-barked pine Pinion heldreichii Horvat 1950 that is further dierentiated to several endemic relict associations. Along with these associations, a special rareness are fragments of com munity of spring heath in the sub-alpine belt Pino mugiEricion carneae, that is on Prenj mountain represented by endemic association of G entiano dinaricae-Ericetum carneae*. Vegetation in ecosystems of mountain pine ( Roso pendulinae-Pinetea mugo eurillat 1995 in eurillat et al. 1995) Upper timberline in the investigated area is rep resented by mountain pine Pinetum mugi Hada 1956. Due to strong mountain winds and deep and long-last ing snow, mountain pine has attened physiognomy. Soils are organic kalkomelanosol and rendzine. Syntaxo nomical, vegetation of mountain pine is dierentiated into order Junipero-Pinetalia mugi e alliance Pinion mugo Pawlowski in Pawlowski, Sokolowski & W allisch 1928 comprises communities of mountain pine from the sub-alliance V iolo biorae-Pinenion mugo Redi 2000 with association Pinetum mugi "dolomiticum" G eranio silvaticae-Pinetum mugi and Pinetum mugi "calcicolum" Lakui et al. 1973. e alliance Erico carneae-Pinion mugi Leibundgut 1948 em. Redi 2000 includes the community Erico carneae-Pinetum mugo Redi 2000 on warmer dolomite type of soil in the sub-alpine belt, which on southern and western slopes of vrsnica mountain. descends to 1,300 m above sea level. Vegetation in ecosystems of dark coniferous for ests ( Vaccinio-Piceetea Br.-Bl. in Br.-Bl. et al 1939) At north and northwestern slopes of vrsnica, com munities of dark conifer forests are developed in frag ments. ey are especially representative below the peaks Plono and Vilinac, toward Masna Luka. ey are devel oped on carbonate surface, on soils of A-C prole hu mus and rendzine. It is important to emphasize that on these biotopes there is a mix of thermole populations of spruce Picea abies (L.) H.Karst and the coldest popu lations of white barked pine Pinus heldreichii H. Christ. Syntaxonomically, this vegetation is dierentiated on two orders: (i) V accinio-Piceetalia Br.-Bl. 1939, alliance V ac cinio-Piceion Br.-Bl. 1938 with communities: Pino heldre ichii-Piceetum abietis* and Abieti-Piceetum abietis pineto sum heldreichii* and (ii) Calamagrosti-Abietetalia Fukarek 1969 (forests of dinaric r on limestone blocks), with the S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 549 alliance Calamagrosti-Abietion Horvat 1963 and the com munity Calamgrosti-Abietetum albae Horvat 1950. At coldest biotopes there are communities of r and spruce Abieti-Piceetum abietis bosniacum Lakui et al 1979, and at limestone blocks there are communities of Calamgrosti-Abietetum albae Horvat 1950, especially presented on northern slopes of Prenj. Vegetation of arable, rural and abandoned eco systems Anthropogenic impact in some areas is so distin guished that it lead to conversion not only primarily to secondary, but also secondary to less organized commu nities. By their origin, they are tertiary. ose are com munities on cultivated surfaces, abandoned biotopes, along houses, roads, trampled places. Soils, without consideration of their original nature and type, are nitri ed and troubled. Depending on intensity and nature of action of anthropogenic factor in the past, tertiary veg etation is dierentiated to following communities that participate in building of several ecosystems: (i) ecosys tems of arable and abandoned places from the class Stel larietea mediae R. Tx., Lohmeyer & Preising in R. Tx. ex von Rochow 1951, order Atriplici-Chenopodietalia albi R. Tx. (1937) Nordhagen 1940 and alliances Panico-Setarion Siss. in W estho et al 1946 and Polygono-Chenopodion polyspermi W Koch & R. Tx. 1937; order Chenopodietalia Br.-Bl.1931 em.1936 with communities of Chenopodion muralis Br.-Bl. (1931) 1936 and H ordeion Br.-Bl. (1931) 1947 alliances, on warmer nitried soil: (ii) ecosystems of humid and nitrophyllous vegetation from the class Biden tetea tripartiti Tx. Lohmeyer & Preising 1950, order Bi dentetalia Br.-Bl. & Tx. 1943 and M enthion pulegii Lakui 1973 community; (iii) ecosystems of trampled and poorly aerated soils of the class Plantaginetea majoris Txen & Preising in Txen 1950, order Plantaginetalia majoris R. Tx. 1950 and alliances Polygonion avicularis Br.-Bl. 1931 ex Aich. 1933 and Agropyro-Rumicion crispi Nordhagen 1940 (iv) ecosystems of abandoned nitried places from the class Chenopodietea Br.-Bl. 1951, order Onopordetalia Br.-Bl. & R. Tx. 1943 ex Klika & Hada 1944 and the alli ance Arction lappae Tx. 1937 em. Siss. 1946 and (v) eco systems of dried nitried soil from the class Artemisietea vulgaris Lohmeyer et al. ex von Rochow 1951 encom passed by the order Artemisietalia Lohmeyer ap. Tx. 1947 and alliances Artemision absinthii Lakui et al 1975 and the endemic alliance Cirsion candelabri *. Communities of tertiary vegetation unlike previous types of primarily and secondary vegetation mostly do not contain endemic species. Exceptions are communi ties form the alliance Cirsion candelabri* that are present along roads on Prenj and vrsnica mountains. Besides, in optimal measure, these communities and their oris tic structure are benecial to richness of biological diver sity and diversity of biotope. GIS INTERPRETATION OF MAIN T YPES OF KARST VEGETATION Distribution of basic types of communities is given in GIS interpretation in original proportion 1: 25 000 (Figs. 4 & 5) for the whole area of endemic development centre in complex of mountains of Prenj, vrsnica and abulja. Due to variable mosaic distribution caused by dis tinguishing relied as well as other factors of non-biotic nature, cartography and in this proportion, it was impos sible to present communities at level of association. In order to do it this overview was conducted on level of groups, order and sometimes class, or type. However, the oered GIS cartographic interpretation could denitely be used for the following purpose: to consider and assess real conditions of biological and geological morpho logical and hydrological diversity of this complex area. Total area of endemic centre of Prenj, Cvrsnica and abulja mountains covered by this research was about 105,000 ha. In the area of Prenj mountain coverage is about 51,754.38 ha, while in the complex of vrsnica and abulja mountains it is 51,752.72 ha. In both areas at higher altitudes (that are at the same time very valu able from aspect of biological and geo-morphological di versity) mountain pastures are dominating, rock creeps, glaciers and crak in the rocks and sub-mountain forests of white barked pine Pinetum heldreichii s.lat, sub-moun tain forests of beech and maple Aceri-Fagetum "subalpi num" are present. In lower parts, on south expositions, on shallow soils deciduous woods and thicket of Moe sian beech, black hornbeam are present, while in the val leys, especially in canyon of river Neretva, shrubbery and coppice with white hornbeam Carpinetum orientalis s.lat are present. Besides what has already been mentioned, the cartographic presentation provides an overview over distribution of similar types of vegetation that represent the real condition of bio-cenological, as well as oristic biodiversity at the territory of the endemic centre. e situation on the mountain of Prenj is mostly the same as at the territory of mountains of vrsnica and abulja research area. On this mountain complex dier ent forms of deciduous forests and their pro-gradation degradation stages dominate (Figs. 4 & 5). In the spectre of vegetation, the biggest proportion is of deciduous forests of beech and its dierent develop ment stages (especially at the territory of abulja), then thermole woods of oak and their degradation stages coppice and shrubbery of white hornbeam and black hornbeam, as well as vegetation of thermole meadows and bullhead (Figs. 4 & 5). VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 550 HIGH LEVEL OF VEGETATION RICHNESS Considering this extremely important parameter, as one of the best indicators for specic ecological diversity, the area of endemic centre of Prenj, vrsnica and abulja is considered to be one of the richest and most unique ones not only in Bosnia and Herzegovina and Dinaric Alps, but also in the entire Mediterranean area and Europe (Tab. 6). us, this area represents the biodiversity "hot spot" in SE Europe, and perhaps even wider (Redi 2008a, 2008c). is claim is illustrated in 314 associations, 113 alli ances, 60 orders and 33 classes which were described so far on the territory of Bosnia and Herzegovina (Lakui et al. 1978) (Fig. 6). Compared with the vegetation diver sity of Montenegro Blei & Lakui (1976) found high proportion of syntaxa in endemic development centre Prenjvrsnicaabulja. As much as 92% of classes and 88% of association of Monte negro were numerically con tained in the vegetation of this endemic centre. is in cludes especially the moun tains Magli, Volujak, Dur mitor, Bjelasica, Sinjajevina, Prokletije, Komovi, Haila, Loven and Rumija (Lakui 1970; Blei & Lakui 1976; Lakui & Redi 1989). Comparison of vegetation diversity is made with Croatian vegetation (Trinajsti 2008). e most recent survey of vegetation in neighbouring Croatia showed a high level of syntaxonomical diversity (Tab. 6). Croa tian vegetation belongs to 42 classes, 61 orders, 120 al liances and 408 associations. ere is a high proportion of associations of endemic centre in the total number of plant communities of Croatia (58%). However, endemic development centre Prenjvrsnicaabulja is richer in alpine and sub-alpine communities of the vegetation class Elyno-Seslerietea Salicetea herbaceae and Aspleni etea trichomanis (Tab. 6 & Fig. 6). Fig. 6: Comparative overview of syntaxonomical diversity of Bo snia and H erzegovina with same parameters used in region and Europe. Fig. 7: Comparative overview of syntaxosonomical diversity of endemic centre of Prenj, vrsnica and abulja mountains at Bo snia and H erzegovinas level. T ab. 6: Comparative overview of vegetation diversity of the endemic centre of Prenj, vrsnica and abulja mountains with same pa rameters used for diferent geographical areas. Geographic region Class Order Alliance Association Source Europe 80 233 928 ? Rodwell et al. 2002 Bosnia andHerzegovina 33 60 113 314 Lakui et al. 1978 Montenegro 37 53 97 267 Blei & Lakui 1976 Croatia 42 61 120 408 Trinajsti 2008 Vranica mountain (Bosnia and Herzegovina) 27 44 73 165 Redi 2007 Endemic centre Prenjvrsnicaabulja 34 63 116 236 this paper (Redi et al. ) T ab. 7: Proportions of some syntaxonomical categories of the endemic centre of Prenj, vrsnica and abulja mountains in the vegetation of selected geographical area. Region Class Order Alliance Association Endemic centre Prenjvrsnicaabulja 34 63 116 236 Proportion in Europe (%) 42.5 27 12.5 Proportion in Bosnia-Herzegovina (%) >100 >100 >100 75 Proportion in Montenegro (%) 92 >100 >100 88 Proportion in Croatia (%) 81 >100 97 58 S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 551 CONCLUSIONS e mountain complex Prenj, vrsnica and abulja in Herzegovina is considered to be one of the largest karst regions in Dinaric Alps and even wider. Besides, this mountain complex encompasses some of the highest peaks of central and litoral Dinaric Alps. Specic forms of geogenesis, orogenesis and gen esis of climate that took place in the long history of the Earth, have led to extremely high diversity of karst phe nomena, such as sinking rivers, karren, sinkholes, and caves, which are today most unique habitat types with rich wilderness. In the sub-alpine and alpine belt cirques and snow beds occur; as a remnant of glaciation that happened in past and aer going through karstic hydrogenesis re sulted in development of mountain (glacial) lakes. ese lakes are today most unique ecosystems with a spectre of both plant and animal species of glacial-relict character. is unique forms of geogenesis, orogenesis, hy drogenesis and genesis of climate have caused specic directions of pedogenesis, which led to occurence of diverse developmental series of carbonate soils vari ous forms of lithosol on bare rocks, regosol in crevices of carbonate rocks, calcomelanosol and rendzine, while on at ground deeper soil types such as calcocambisol, modied terra rossa and calcoluvisol occur in the lowest areas. By means of powerfull currents at mountain foot, recent alluvial depositions and colluvium evolve daily. e unique pattern of evolution of abiotic ecosys tem components was a fundamental determinant for specic forms of karstogenesis in this area, which cor responds directly with unique patterns of genesis of ora and syngenesis, respectively development of ora and vegetation. ese unique and diverse forms of karsto genesis provide the answer to high oristic richness, as well as to extremely high level of endemism and relict ness. Because of that, this area has got every right to be designated as the endemic development centre, and aer this research and modern evaluation process even as a biodiversity hotspot of great importance not only for the Mediterranean, but on global scale. If compared with the European ecological diversity (Ellenberg 1986; Rodwell et al. 2002), the territory of this endemic centre shows extremely high level of diversity (Tab. 6). Out of 928 vegetation units, as it was recorded in Europe by now, 13% were recognized at this territory. Vegetation units at level of order, which are to be found here, participate with 27% in the vegetation diversity of Europe, while share of classes amounts 43% in the Euro pean vegetation (Tab. 7). In addition, the ecological diversity of this area ex ceeds extremely high diversity like the one that was not ed on silicate carbonate massif of Vranica mountain in central Bosnia (Lakui et al 1979; Redi 2007c, 2007d). It is important to stress that diversity at some levels, for example at level of higher syntaxa (Fig. 7), is higher than in Bosnia and Herzegovina. Similar relation is in view of proportions and num ber of vegetation orders, as well as classes as highest forms of vegetation integrity. ese data unquestionably indicate that this endemic centre is truly unique, biologi cally rich and that it makes specic "hot vegetation spot" in both Mediterranean and whole Europe. On the other side, these data indicate still rather weak research of bio logical diversity, and specially syntaxonomy of plant cov er in karst at the territory of Bosnia and Herzegovina. AKNO WLEDGEMENTS is paper was co-nanced by means from the scientif ic-investigation project: Procjena ugroenosti i obrasci odrive zatite genofonda endema u razvojnom centru Prenj vrsnica abulja ("Assessment of threats and pat terns for the sustainable protection of endemic gen pool in the development centre Prenjvrsnicaabulja), Project Number: 03-39-5980-67-2/08, co-nanced by the Federal Ministry of Education and Science of Federation of Bosnia and Herzegovina. Major contribution to the projects realization was provided by consultants: academician Ljubomir Berberovi, academician Taib ari and academician Mithat Usupli, Center for Carstology at the Academy of Science and Arts of Bosnia and Herzegovina. Field work in technical respect was realized by the support of technical assistant Sedik Veli. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...

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ACTA CARSOLOGICA 40/3 2011 554 Muratspahi, D., Redi, S., & R. Lakui, 1991: Asocijacija Rusco-Carpinetum orientalis Ble. et Lki 1966 u dolini rijeke Neretve.Glasnik Republ. zavoda zat. prirode Prirodnjakog muzeja, 24, 7. Murbeck, S., 1891: Beitrge zur Kenntnis der Flora von Sdbosnien und der Hercegovina.pp. 192, Lund. Oberdorfer, E., 1983: Panzensoziologische excursions ora .Verlag Eugen Ulmer, pp.1051, Stuttgart. Prohic, E., Hausberger, G. & J.C. Davis, 1997: Geochemi cal patterns in soils of the karst region, Croatia.Journal of Geochemical Exploration, 60, 139. Redi, S., 1997: e Protection of the Diversity of Vascular Plants in the Karst Poljes of the Dinaric Mountains.In: Sasowsky, I.D., Fong, D.W & E.L. W hite (eds) Conservation and Protection of the Bi ota of K arst Presentations Extended abstracts and eld trip guide for the conference, 13 th th Febru ary 1997, Nashville, Tennessee. State College, PA, 82,. Nashville. Redi, S., 1999: e syntaxonomical dierentiation of the Festuco-Brometea Br.-Bl. & R. Tx. 1943 ex Klika & Hada 1944 in the Balkans.Annali di Botanica, 57, 167. Redi, S., 2003: e syntaxonomy and syngenesis of the Elyno-Seslerietea Br.-Bl. 1948 in the Balkan penin sula.Annali di Botanica (nuova serie), 3, 53. Redi, S., 2004: Biogeographic diferentiation of biodi versity of the Balkan peninsula with reection at the Bosnia and Herzegovina.In: Spahi, M. (ed.) Abstract Book of plenary and session lecuters of First congress of geographes of Bosnia and H erzegovina with international participation 22 th th Septem ber 2004. Geografsko drutvo Bosne i Hercegovine, 23, Sarajevo. Redi, S., 2007a: Karst elds (poljes) of Dinaric Alps at biodiversity centres with global values.In: 50 th In ternational Association for V egetation Science Sym posium, Abstracts Book 23 th th July 2007, Swan sea, UK. International Assciation for vegetation science, 47, Swansea. Redi, S., 2007b: Role of glaciation centres in the bio geography richness of Balkan peninsula (SE Eu ropa).In: ird I nternational Biogeography Society Conference, Symposium 2. Quaternary impacts on H olarctic Biogeography, Book of Abstracts, 9 th th January 2007, Tenerife, Canary Islands. e inter national biogeographical society, 45, Tenerife. Redi, S., 2007c: Syntaxonomic diversity as an indica tor of ecological diversity case study Vranica Mts. at the Central Bosnia.Biologia, Section Botany, 62, 173. Redi, S., 2007d: e syntaxonomy of the vegetation of the continental Dinaric Alps (W Balkan).Col lecion of Papers Devoted to Academician Kiril Micevski, Macedonian Academy of Sciences and Arts, 249, Skopje. Redi, S., 2008a: Ecohydrological approach to sustain able management of oodable karst elds in Dinaric Alps (W estern Balkan).In: Zalewski, M. (ed.) Ab stract of I nternational Conference: "Ecohydrological Processes and Sustainable Floodplain M anagement, I AP.6 19 th th May 2008, odz, Poland. Interna tional institute of the Polish academy of sciences European regional centre for ecohydrology under auspice of UNESCO, p. 44, odz. Redi, S., 2008b: Jasnin encijan G entiana jasnae (Lakui) Redi ukras bosanskih planina.Fond eko svijet, 26, 22. Redi, S., 2008c: Sytaxonomic diversity in the evalua tion of biodiversity hotspots in Mediterranean re gion.In: Chitry, M. (ed.) Abstracts of 17 th I nterna tional Workshop European V egetation Survey, 1 th th May 2008, Brno. Masaryk University, 96, Brno. Redi, S., 2008d: e syntaxonomic diversity as a tool in the evaluation of biodiversity hot spots in Medi terranean region.In: Pei, V. (ed.) e Book of abstracts and programme of plenary and session lectures of III I nternational Symposium of Ecologist of the Republic M ontenegro 8 th th October 2008, Herceg Novi. e Montenegrinian Ecological Soci ety 33, Herceg Novi. Redi, S., 2009: Patterns of richness and endemism of vascular plants in biodiversity hotspots in Mediter ranean Central Dinaric Alp.Biologia, Section Botany (submitted). Redi S., Muratspahi, D. & R. Lakui, 1992: Some Forests and shrubs phytocoenoses from valley of the Neretva river.Poljoprivreda & umarstvo, 38, 95. Redi, S. & M. Dizdarevi, 1998: Biogeografska karta Bosne i Hercegovine. In: Zupevi, O. (ed.) Atlas svijeta za osnovnu i srednje kole Sejtarija, pp. 35 37, Sarajevo. Redi, S. (ed.), 2007: Biodiverzitet endemnih razvojnih centara na podruju H ercegovine kao doprinos strem ljenjima T argets 2010 .Monograja, EKO BiH, pp. 306, Sarajevo. Redi, S., Barudanovi, S. & M. Radevi (eds.), 2008: Bosna i H ercegovina zemlja raznolikosti. Pregled i stanje bioloke i pejzane raznolikosti Bosne i H er cegovine .Federalno ministarstvo okolia i turizma BiH, pp.164, Sarajevo. S ULEJMAN REDIb, S ENKA BARUDANOVIb, SABINA TRAKIb & D EJAN KULIJER

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ACTA CARSOLOGICA 40/3 2011 555 Redi, S. & S. Barudanovi, 2010: e petterns of di versity of forest vegetation of the Crvanj Moun tain in the Herzegovina (W est Balkan Peninsula).umarski list, 134 (5), 261. Riter-Studnika, H., 1954: Flora i vegetacija livada krakih polja Bosne i Hercegovine.Godinjak Biolokog instituta, 7, 25. Riter-Studnika, H., 1956: Flora i vegetacija na dolo mitima Bosne i Hercegovine.Godinjak Biolokog instituta, 9, 73. Riter-Studnika, H., 1959: Kraka polja BiH kao reliktna stanita biljaka.Zatita prirode, 16, 28. Rodwell, J.S., Schamine, J.H.J., Mucina, L., Pignatti, S., Dring, J. & D. Moss, 2002: e diversity of European vegetation: An overview of phytosociological alliances and their relationships to EUNIS habitats .National Reference Centre for Agriculture, Nature and Fish eries, pp. 168, W ageningen. Scheiner, S.M., 1992: Measuring pattern diversity.Ecol ogy, 73, 1860. Sijari, R., 2000: Fauna Rhopalocera (Lepidoptera) of Mt. Vran (Herzegovina).W issensghaliche Mit teilungen des Bosnisch-Herzegowinischen Landes museums, Naturwissenscha, He C, 7, 325. Sitarz, D. & P. Simon (eds.), 1993: Agenda 21: e Earth Summit Strategy to Save Our Planet .Earth Press, Rowley, MA, USA. Sket, B., 1997: Distribution of Proteus (Amphibia: Urodela; Proteide) and its possible explanation.Journal of Biogeography, 24, 263. Sket, B., Dov, P., Jali, B., Kerovec, M., Kuini, M. & P. Trontelj, 2001: A cave leech (Hirudinea, Eepob dellidae) from Croatia with unique morphological features.Zoologica Scripta, 30, 223. Sliskovic, I., 1995: On the hydrogeological conditions of western Herzegovina (Bosnia and Herzegovina) and possibilities for new groundwater extractions.International Journal of Rock Mechanics and Min ing Sciences and Geomechanics Abstracts, 32, 7, 310. Spahi, M., 2001: Prirodna jezera Bosne i H ercegovine: L imnoloka monograja .Harfo-graf, pp.170, Tuzla. ili, ., 1967: Oreoherzogia pumila (Turra) Vent. subsp. illyrica ili, subsp. nova novi endemini takson zapadnog dijela Balkanskog poluostrva.Glasnik Zemaljskog muzeja Bosne i Hercegovine, 6, 61. ili, ., 1970 : Heliospermo retzdorani-Oreoherzo gietum illyricae, nova zajednica na hercegovakim planinama.Radovi Akademije nauka i umjetnosti BiH, Posebna izdanja 15, Odjeljenje prirodnih i matematikih nauka, 4, 303. ili, ., 1979: M onograja rodova Satureja L., Calamin tha M iller, M icromeria Bentham, Acinos M iller i Clinopodium L. u ori Jugoslavije .Zemaljski muzej BiH, pp. 440, Sarajevo. ili, ., 1984: Endemine biljke .Svjetlost, Zavod za udbenike i nastavna sredstva, pp. 227, Sarajevo. ili, ., 2000: ili, ., 2000: List of species (Pterido phyta and Spermatophyta) for the Red Book of Bosnia and Herzegovina.W issensghaliche Mit teilungen des Bosnisch-Herzegowinischen Landes museums, Naturwissenscha, He C, 7, 287. ili, ., 2002: Endemine i rijetke biljke Parka prirode Blidinje .Matica hrvatska, Ogranak itluk, pp. 279, itluk. Topi, J. & Lj. Ilijani, 2005: Campanula kapelae the new species? within Campanula pyramidalis agg.Natu ra Croatica, 14, 3, 249. Trinajsti, I., 2008: Biljne zajednice Republike H rvatske .Akademija umarskih znanosti, pp. 179, Zagreb. Tutin, T.G., Heywood, V.H., Burges, N.A., Valentine, D.H., W alters, S.M. & D.A. W ebb, 1964: Flo ra Europea Vol. I-V. Cambridge University Press, Cambridge. Verovnik, R., Sket, B. & P. Trontelj, 2004: Phylogeogra phy of subterranean and surface population of wa ter lice Asellus aquaicus (Crustacea: Isopoda).Mo lecular Ecology, 13, 1519. W eber, H.E., Moravec, J. & J.P. eurillat, 2000: Inter national Code of Phytosociological Nomenclature. 3rd edition.Journal of Vegetation Science, 11, 739. W hite, W .B., 1988: G eomorphology and H ydrology of K arst T errains .Oxford University Press, pp. 464, New Y ork. VASCULAR PLANT BIODIVERSIT Y RICHNESS AND ENDEMORELICTNESS OF THE KARST MOUNTAINS PRENJ, VRSNICA ...


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