66 ACTA BOT. CROAT. 78 (1), 2019 Acta Bot. Croat. 78 (1), 66–81, 2019 CODEN: ABCRA 25 DOI: 10.2478/botcro-2019-0004 ISSN 0365-0588 eISSN 1847-8476 Algal assemblages in springs of different lithologies (ophiolites vs. limestone) of the Konjuh Mountain (Bosnia and Herzegovina) Jasmina Kamberović1*, Anđelka Plenković-Moraj2, Koraljka Kralj Borojević3, Marija Gligora Udovič2, Petar Žutinić2, Dubravka Hafner4, Marco Cantonati5 1 University of Tuzla, Faculty of Natural Sciences and Mathematics, Univerzitetska 4, 75000, Tuzla, Bosnia and Herze- govina, jasmina.kamberovic@untz.ba 2 University of Zagreb, Faculty of Science, Department of Biology, Rooseveltov trg 6, 10000 Zagreb, Croatia 3 Minnesota Drive, Great Sankey, Warrington, UK, WA5 3SY 4 Bartulovići 4, 20357 Blace, Croatia 5 MUSE – Museo delle Scienze, Limnology and Phycology Section, Corso del Lavoro e della Scienza 3, I-38123 Trento, Italy Abstract – The biodiversity of algal communities and environmental conditions were investigated in the springs of Mt. Konjuh. The assemblages of 20 springs emerging from different lithologies (limestones and ophiolites, re- spectively) comprised 234 algal taxa. Diatoms and cyanobacteria were the most species-rich groups. The most common alkaliphilic, circumneutral, and eutraphentic diatoms were represented by the genera Gomphonema, Nitzschia, Navicula, Cymbella, and Achnanthidium, and by the cyanobacterial genus Phormidium. Hierarchical clustering and SIMPROF analysis based on relative algal abundance clustered springs into six groups, separating them mainly according to spring type and lithology. Indicator species for groups and springs on different litholog- ical substrata were singled out, revealing 33 taxa with preferences for ophiolites, and 20 taxa with preferences for carbonates. The values of the Shannon-Wiener diversity index were moderately high per spring location, and similar for the two groups of springs on different lithologies. A higher similarity in species composition was noted between springs on ophiolites and limestones than between springs on ophiolites and other types of siliceous sub- strata. The present study suggests that algal assemblages in springs emerging from ophiolites, even those made up by a preponderance of silicates, should be analyzed separately from those related to springs on other siliceous substrata. The results obtained showed that most of the springs studied are affected by anthropogenic impacts and morphological alterations leading to the dominance of highly competitive meso- and eutraphentic algal species, thus emphasizing the importance of further investigation and conservation of these habitats. Keywords: biodiversity, diatoms, cyanobacteria, springs, geological substratum, ecological preferences, trophic status * Corresponding author, e-mail: jasmina.kamberovic@untz.ba Introduction The importance of spring habitats as biodiversity hotspots and refugia for biodiversity conservation (Cantonati et al. 2012a, Taxböck et al. 2017) has been emphasized by the de- scription of a large number of rare and endangered algal spe- cies (Sabater and Roca 1990, 1992, Cantonati 1998, Werum 2001, Poulíčková et al. 2005, Nascimbene et al. 2011). Half of the pennate freshwater diatoms described in Europe were recorded from spring ecosystems (Werum and Lange Berta- lot 2004). Studies dealing with the algal biodiversity of Eu- ropean springs have increased since the mid 1990s (Sabat- er and Roca 1990, Cantonati 1998, Falasco and Bona 2011, Werum and Lange-Bertalot 2004, Fránková et al. 2009). Some of them resulted in the discovery of new diatom gen- era and species (Cantonati and Lange-Bertalot 2006, 2009, 2010, 2011, Cantonati et al. 2009, 2010). Studies on algae in Bosnia and Herzegovina date back to the late 19th century and the early 20th century (Protić 1897, 1899, 1901, 1908, Gutwinski 1899, Beck 1928), whereas the first detailed stud- ies on diatoms were carried out in the framework of a wider study of diatom assemblages in several lakes and springs of ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 67 the Balkan Peninsula (Hustedt 1945). However, crenic algal assemblages in Bosnia and Herzegovina have been investi- gated only at a few karst spring locations (Blagojević 1974, 1976, 1979, Hafner 2009, Dedić et al. 2015). Results deriving from the study of algal ecology in springs (Cantonati et al. 2012b, c) have application in the monitoring of freshwater ecosystems. As a commonly stud- ied algal group, diatoms are a useful proxy for water quality, reflecting both physical and chemical characteristics of a wa- ter body (Kelly and Whitton 1995, Rott et al. 1999, Werum and Lange-Bertalot 2004, Wojtal 2013). Diatom assemblag- es can be grouped according to different lithology (Werum and Lange-Bertalot 2004). For instance, Cymbella tridentina Lange-Bertalot, M. Cantonati & A. Scalfi is a typical cren- ophilous species living on carbonate substrate (Cantonati et al. 2010), while Achnanthidium dolomiticum M. Cantonati et Lange-Bertalot is more common in springs fed by drain- age basins dominated by dolomite lithology (Cantonati and Lange-Bertalot 2006). Diatom genera such as Achnanthes s.l. and Cymbella are generally more represented on carbonate substrates, whilst Eunotia and Pinnularia are more frequent on silicate rocks (Cantonati 1998). Algal assemblages in springs on ophiolitic or ultramaf- ic rocks have been rarely studied (Werum 2001). Although widespread on several continents (Dilek and Furnes 2011), these substrata cover less than 1% of the Earth's surface (Coleman and Jove 1992). Dinaric ophiolites constitute one of the largest exposures of mantle rocks on Earth (Lugović et al. 1991). Ophiolites from Bosnia belong to the western belt of ophiolites in the Balkans as fragments of a long chain which passes through western Serbia (Mt. Zlatibor), Kosovo and the western belt of the Mirdita ophiolite in Albania to the Pindos and Othris ophiolites in central Greece. The ophi- olites and associated sediments (mostly carbonates and ophi- olitic melange) cover a surface of tens of thousands square kilometers in Bosnia and Herzegovina, with the Krivaja- Konjuh mountain complex, which is the largest ophiolite zone in this area, positioned in the north-eastern part of the country (Trubelja et al. 1995, Babajić 2009). Still, despite its wide-ranging coverage, freshwater algal assemblages on Di- naric ophiolite substrata are largely unexplored. Unlike oth- er siliceous rocks, ophiolites have less than 45% concentra- tions of silica, generally contain high amounts of Fe, and trace metals such as Ni and Cr (Alexander et al. 2007, Tru- belja et al. 1995). Results of chemical studies on ultramaph- ic rocks of the Krivaja-Konjuh ophiolitic complex demon- strated modal composition with high content of MgO, low content of CaO, and a high MgO: FeO ratio of about 5 and more (Operta 2017). The main objectives of this study were to: (i) explore al- gal biodiversity in crenic habitats of Mt. Konjuh, north-east- ern Bosnia and Herzegovina, including rarely investigated springs on ophiolites, (ii) describe ecomorphological, phys- ical and chemical properties of these spring habitats, (iii) analyze the influence of geological substrata on algal assem- blages, and (iv) describe the ecological preferences of domi- nant algal groups. Materials and methods Study area Konjuh Mountain is a part of the central Dinarides on the Balkan Peninsula, with an altitude of 1326 m a.s.l. In 2009 it was designated a Protected Landscape (IUCN Category V) with ecosystems characterized as Tertiary relicts. It belongs to a zone with a moderately continental climate, with a mean annual temperature varying between 9.2–10 °C, and annu- al precipitation ranging from 900 to 1250 mm (Kudumović Dostović 2012). As part of the ophiolitic complex, Mt. Kon- juh consists mainly of peridotite and serpentinite rocks. In addition, ophiolitic melange, gabbro, shale and limestones are also found (Ristić et al. 1967). Physical and chemical analyses Ecomorphological characteristics of springs (shading, current velocity, spring type) were assessed using methods described in Spitale (2007). Discharge was measured by a graduated bucket. The flow discharge for large rheocrenic springs was estimated by multiplying the cross section area, obtained from the values of average depth and width of the stream channel, and average velocity, which was measured by the float method. Temperature, dissolved oxygen, pH, and conductivity were measured directly on site with a Hanna HI 9828 Multimeter (Hanna Instruments Inc., U.S.A.). Water for analyses of chemical parameters was taken at the same time. Total alkalinity, calcium, and magnesium were analyzed via titration, and sulphates were measured using the turbidimet- ric method (APHA, AWWA and WEF, 1999). Ammonia, ni- trates and nitrites were analyzed via a spectrometric meth- od according to the standard procedures (BAS ISO 7150-1: 2002, BAS ISO 7890-3:2002, BAS EN 26777:2000) in Bosnia and Herzegovina (Institute for Standardization of Bosnia and Herzegovina, 2002a, b, 2000, respectively). Algal sampling and identification A total of 196 algal samples were collected at the spring- head and 15 meters downstream during the spring, summer and autumn of 2013. Altogether 20 springs were seasonally studied: nine springs on ophiolites (2 rheocrenes, 6 rheohe- locrenes, and one hygropetric spring), one (rheohelocrenic) spring on ophiolitic melange, and ten springs on carbonate (limestone) substrate (7 rheocrenes and 3 rheohelocrenes). The springs are situated at altitudes ranging between 402 and 1003 m a.s.l (Fig. 1). Epilithic algae were taken by brushing the surfaces of 5–6 stones (Kelly et al. 1998), samples of epi- bryon were collected from aquatic bryophytes, while epip- elon was collected from the sandy bottom with a cylinder. Abundance of diatom and non-diatom algae was estimat- ed as number of cells per square centimeter (epilithon and epipelon) or number of cells per gram of dried bryophyte mass. Non-diatom species were identified in fresh samples, whilst diatoms were acid cleaned (Hustedt 1930) and mount- ed in Naphrax (Brunel Microscopes Ltd., U.K.). At least 400 diatom valves were identified in each slide using random KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 68 ACTA BOT. CROAT. 78 (1), 2019 transects under an Olympus BX41 light microscope (Olym- pus Corporation, Japan) at a magnification of 1000x. Quan- titative data on diatoms and non-diatoms were integrated according to Stevenson and Bahls (1999). Because of their estimation per different units, and the need for uniformity for statistical analysis, the absolute cell numbers were trans- formed into relative (percentage) abundances. In addition to the abundance, which shows the average percentage of the taxa within samples, data on the taxa frequency are also given to provide information on the occurrence of taxa in the analyzed group of samples. Identification was done following Starmach (1985), Cvijan and Blaženčić (1996), Krammer and Lange-Berta- lot (1986, 1988, 2000, 2004 a,b), Komárek and Anagnosti- dis (1998, 2005), Krammer (2000, 2002), John et al. (2002), Hofmann et al. (2011) and Lange-Bertalot et al. (2017). Data processing and statistical analyses Diversity was calculated by using species richness (S) as the number of identified taxa, the Shannon-Wiener index of species diversity, H'(ln), (Shannon and Weaver 1949) and Pielou’s index of equitability, J' (Krebs 1999). Ecological re- quirements of diatoms for moisture, pH and trophic status were obtained from van Dam et al. (1994). The diatom tro- phic index (TI) by Rott et al. (1999), the Croatian trophic di- atom index (TIDHR) and the Ecological Quality Ratio – EQR (Anonymous 2013, 2016) were used in the evaluation of tro- phic and ecological status of spring ecosystems. All data were transformed by using the square root function prior to statistical analyses. Non-metric multidi- mensional scaling (NMDS) and hierarchical group average clustering based on mean values of the relative algal abun- dance for each spring were used for spring ordering. Simi- larity profile analysis (SIMPROF) with iterative permutation procedure (999 permutations) was used for the detection of boundaries among clusters. The ordination was conducted on the Bray-Curtis similarity matrix of species data (Legen- dre and Legendre 1998). Normal distribution of the main physical and chemical parameters and values of diversity in- dices were tested by the Kolmogorov-Smirnov test in order to select parametric or nonparametric test. The significance of differences in the main physical and chemical factors be- tween springs emerging from two different lithologies (ophi- olites vs. limestone) was analyzed using the nonparametric Mann-Whitney U test (p < 0.05). Parametric t-test was used to test for differences between diversity indices in springs from contrasting lithologies. The IndVal analysis (Dufrene and Legendre 1997) was used to identify characteristic spe- cies of: i) springs on different lithological substrata, and ii) different clusters. The significance of indicator value of each species was estimated by Monte Carlo permutation tests. Statistical analyses were performed in IBM SPSS Statistics for Windows version 22.0 (IBM Corp., U.S.A.), Primer 6.0 (Clarke and Warwick 2001), and PC-ORD 5 software pack- ages (Grandin 2006). Results Environmental variables Ecomorphological characteristics of springs, including spring names with short spring codes, location, shading, ve- locity and discharge are described in On-line Suppl. Tab. 1. Springs on ophiolites were characterized by low but rela- tively steady discharge (≤ 2 L s–1), water temperature below 8 °C, low turbidity, low calcium content, high magnesium, and high dissolved oxygen concentrations (On-line Suppl. Tab. 2). On the other hand, springs on carbonates (lime- stone) had an unstable discharge varying from complete dry- ing up to 150 L s–1 (On-line Suppl. Tab. 1). Carbonate rheo- crenic springs had higher values of oxygen concentration than the rheohelocrenes, neutral to slightly alkaline water, and high calcium content. High nitrate concentrations (up to 18 mg L–1) were noted in several springs independently from the lithology. The values of alkalinity and conductivity ranged from 16 to 70 mg CaCO3 L–1, and 136–584 µS cm–1, respectively (On-line Suppl. Tab. 2). Springs emerging from ophiolites showed higher values of temperature, pH, magne- sium and lower values of conductivity, discharge, sulphates and calcium than springs on carbonates (On-line Suppl. Tab. 3). During field studies many negative anthropogenic influ- ences were noted. The most pronounced effects are altera- tions of morphology of the spring area for the purpose of water abstraction, deforestation with accompanying effects Fig. 1. Location of the study sites presented by spring codes on the Mt. Konjuh in Bosnia and Herzegovina. ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 69 of tree transport, road infrastructure building and trampling by cattle. Species composition, diversity, ecological preferences of diatoms and trophic status A total of 234 algal taxa were identified in 196 samples collected from 20 springs. All identified taxa are listed in Tab. 1. The most abundant algal groups were diatoms and cya- nobacteria, with 187 and 34 taxa, respectively. The other al- gal classes were Xanthophyceae (3), Florideophyceae, Ulvo- phyceae, Chlorophyceae and Conjugatophyceae (2 in each), Chrysophyceae and Klebsormidiophyceae (1 in each). To- tal number of recorded taxa ranged between 46 (rheocrenic spring on ophiolite – 2KE) and 76 (rheohelocrenic spring on ophiolite – SK). The most common algal genera were Gomphonema (17), Nitzschia (15), Navicula (11), Phormi- dium (9), Cymbella (9), and Achnanthidium (8). The species Achnanthidium minutissimum, Planothidium lanceolatum, Meridion circulare, Cocconeis pseudolineata and Amphora pe- diculus were recorded in more than a half of all investigated springs. Besides the above mentioned, also very abundant were Cocconeis lineata, Gomphonema micropus, Cocconeis euglypta, Odontidium mesodon and Tapinothrix varians. Hierarchical group average clustering and the SIMPROF test identified 6 assemblages (Fig. 2) mainly related to spring type (rheocrenes and rheohelocrenes) and geological sub- stratum (ophiolites, carbonates, and ophiolitic melange). Species contribution to the clusters was described by the SIMPER analysis (On-line Suppl. Tab. 4), and statistically significant indicator species (p < 0.05) for each cluster pro- vided by the IndVal analysis are listed in Fig. 2. Species indi- cator values and contribution of the most abundant species could not be technically estimated for springs MV (cluster 1) and ZL (cluster 6), because the first and the sixth cluster included only one spring. However, the most frequent and abundant species in spring MV (cluster 1) were Achnanthid- ium minutissimum, Achnanthidium pyrenaicum, Odontidium mesodon, Nitzschia fonticola and Amphora pediculus, whilst the most abundant and frequent species in spring ZL (clus- ter 6) were Meridion circulare, Gomphonema micropus, Eu- notia soleirolii, Nitzschia dubia and Ammatoidea sp. Cluster 2 (rheocrenes on ophiolites) was characterized by the xero- tolerant diatom Humidophilla perpusilla, distinguished by both IndVal and SIMPER analysis. Taxa distinguished by IndVal were the eutraphentic diatom Gomphonema parvu- lum and the cyanoprokaryote Pseudanabaena sp. SIMPER analysis singled out several species with high contributions: Planothidium lanceolatum, Chlorogloea microcystoides, Me- ridion circulare, Phormidium formosum, and other taxa listed in On-line Suppl. Tab. 4. Rheocrenes on carbonates (clus- ter 3) were characterized by Rhoicosphenia abbreviata and Nitzschia fonticola (given by IndVal), and a high contribu- tion of the rheophilic cyanobacterium Tapinothrix varians, the crenophilous diatoms M. circulare and Odontidium me- sodon, and the epiphytic diatom species P. lanceolatum and Cocconeis lineata. Cluster 4 comprised rheohelocrenes on ophiolites and carbonates, and was characterised by the di- atoms Cocconeis pseudolineata, Encyonopsis cesatii, Diplo- neis fontanella, and Tryblionella angustata. IndVal also sin- gled out Achnanthidium exile, Amphora lange-bertalotii var. tenuis, Cymbella hantzschiana, Gomphonema subclavatum, and Nitzschia capitellata. Representatives of shaded rheo- crenes (cluster 5) were eutraphentic diatoms, such as Am- phora pediculus, Encyonema minutum, Fallacia subhamulata, Gomphonema angustatum, Achnanthidium affine, and Cym- bella diminuta. According to NMDS, springs were separated by litholo- gy, with carbonate springs distributed on one side and ophio- Fig. 2. Hierarchical group average clustering based on algal assemblages in the studied springs (presented by spring codes) on the Mt. Konjuh and characteristic species for each cluster identified by the IndVal analysis. Number of clusters was calculated with the SIMPROF analysis. KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 70 ACTA BOT. CROAT. 78 (1), 2019 Tab. 1. Algae of springs in the Mt. Konjuh, GS – geological substrata (O – ophiolites, C – carbonates, OM – ophiolitic melange); A – mean abundance per sample, %; F – frequency per sample, %; max. – maximum relative abundance per sample, %; N – number of springs where the taxon was found. Taxa GS A F max. N Cyanophyceae Ammatoidea sp. O-C-OM 8.5 7.1 50.0 4 Aphanocapsa minuta (Kylin) Whitton O-C 0.4 1.0 0.7 2 Chamaesiphon confervicola A.Braun C 32.5 1.0 58.5 1 Chamaesiphon fuscus (Rostafinski) Hansgirg O-C 1.9 2.0 5.9 3 Chamaesiphon incrustans Grunow C 1.1 1.5 3.1 1 Chamaesiphon polonicus (Rostafinski) Hansgirg O 9.1 3.6 28.6 5 Chamaesiphon sp. C 3.5 2.0 5.9 3 Chamaesiphon subglobosus (Rostafinski) Lemmermann O 0.2 0.5 0.2 1 Chlorogloea microcystoides Geitler O-C 23.0 9.2 78.6 6 Chroococcaceae O-C-OM 21.2 4.1 72.7 5 Chroococcus sp. O 0.1 0.5 0.1 1 Chroococcus turgidus (Kützing) Nägeli O-C 2.2 4.6 5.0 5 Clastidium setigerum O.Kirchner O 0.3 0.5 0.3 1 Cyanothece aeruginosa (Nägeli) Komárek O 0.03 0.5 0.0 1 Hassallia pulvinata Frémy O-C 8.0 2.0 20.1 2 Hyella sp. C 0.3 0.5 0.3 1 Leptolyngbya perforans (Geitler) Anagnostidis et Komárek C 28.8 1.0 56.2 1 Leptolyngbya sp. O-C-OM 7.3 10.2 39.6 9 Lyngbya martensiana Meneghini ex Gomont O-C 1.9 3.1 6.1 3 Lyngbya natans Hansgirg O 0.9 0.5 0.9 1 Oscillatoria sp. C 18.0 0.5 18.0 1 Phormidium corium Gomont ex Gomont O 4.8 1.0 8.6 1 Phormidium formosum (Bory de Saint-Vincent ex Gomont) Anagnostidis et Komárek O-C 22.2 16.8 99.0 9 Phormidium griseoviolaceum (Skuja) Anagnostidis C 9.9 0.5 9.9 1 Phormidium incrustatum Gomont ex Gomont C 5.0 1.0 8.7 1 Phormidium kuetzingianum (Kirchner ex Hansgirg) Anagnostidis et Komárek C 1.5 1.0 1.6 1 Phormidium retzii Kützing ex Gomont O-C 22.6 9.7 89.3 7 Phormidium rotheanum Itzigsohn O 2.4 1.0 2.8 1 Phormidium subfuscum Kützing ex Gomont O-C 32.2 2.0 81.3 3 Phormidium tinctorium Kützing ex Gomont C 31.0 3.1 66.7 3 Pleurocapsa aurantiaca Geitler O 0.3 0.5 0.3 1 Pleurocapsa minor Hansgirg O-C 8.7 3.6 35.6 4 Pseudanabaena sp. O-C 6.1 6.1 19.6 3 Tapinothrix varians (Geitler) Bohunická et J.R.Johansen O-C 36.9 28.1 98.7 14 Florideophyceae Audouinella sp. C 21.4 5.1 70.0 3 Batrachospermum sp. C 6.8 0.5 6.8 1 Chrysophyceae Phaeodermatium rivulare Hansgirg C 10.8 3.6 58.4 3 Xanthophyceae Ophiocytium majus Nägeli O 0.1 0.5 0.1 1 Tribonema sp. O-C 4.2 7.1 17.8 5 Vaucheria sp. C 4.9 8.7 24.7 3 Bacillariophyceae and Fragillariophyceae Achnanthidium affine (Grunow) Czarnecki O-C-OM 2.8 14.8 28.4 9 Achnanthidium caledonicum (Lange-Bertalot) Lange-Bertalot O-C 8.8 4.6 36.5 5 Achnanthidium dolomiticum M. Cantonati et H. Lange-Bertalot O 2.4 1.5 3.3 1 Achnanthidium exile (Kützing) Heiberg O-C 4.0 9.2 10.8 8 Achnanthidium minutissimum (Kützing) Czarnecki O-C-OM 20.6 87.8 82.1 20 Achnanthidium pyrenaicum (Hustedt) H.Kobayasi O-C-OM 7.4 29.1 53.4 13 Achnanthidium rosenstockii (Lange-Bertalot) Lange-Bertalot C 0.8 0.5 0.8 1 Achnanthidium straubianum (Lange-Bertalot) Lange-Bertalot O-C 1.5 4.6 4.2 4 Adlafia bryophila (J.B.Petersen) Gerd Moser, Lange-Bertalot et D.Metzeltin O-C 0.6 3.1 1.4 5 Adlafia minuscula (Grunow) Lange-Bertalot O-C-OM 0.9 12.2 2.8 11 Amphipleura pellucida (Kützing) Kützing O-C 1.3 11.7 4.4 7 Amphora copulata (Kützing) Schoeman et R.E.M.Archibald C 1.5 0.5 1.5 1 Amphora indistincta Levkov O-C-OM 2.2 21.9 16.6 12 Amphora lange-bertalotii var. tenuis Levkov et Metzeltin O-C 1.4 10.2 7.3 7 Amphora ovalis (Kützing) Kützing O-C-M 3.0 14.8 15.9 8 Amphora pediculus (Kützing) Grunow ex A.Schmidt O-C 3.6 65.8 28.9 19 ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 71 Taxa GS A F max. N Brachysira vitrea (Grunow) R.Ross C 0.7 0.5 0.7 1 Brebissonia lanceolata (C.Agardh) Mahoney et Reimer O-C 3.0 8.2 27.6 7 Caloneis alpestris (Grunow) Cleve C 0.1 0.5 0.1 1 Caloneis fontinalis (Grunow) Lange-Bertalot et Reichardt O-C-M 0.8 28.6 4.1 18 Caloneis silicula (Ehrenberg) Cleve O-C 0.4 3.1 0.5 4 Caloneis sp. C 0.5 0.5 0.5 1 Caloneis tenuis (W.Gregory) Krammer O-C-M 0.5 4.6 0.9 7 Chamaepinnularia parsura (Hustedt) C.E.Wetzel et Ector C 0.2 0.5 0.2 1 Cocconeis disculus (Schumann) Cleve O 3.0 3.1 8.2 1 Cocconeis pediculus Ehrenberg C 0.2 1.0 0.2 1 Cocconeis euglypta Ehrenberg O-C-OM 5.2 39.8 38.0 15 Cocconeis placentula var. klinoraphis Geitler O 0.5 0.5 0.5 1 Cocconeis lineata Ehrenberg O-C-OM 6.9 58.7 85.5 18 Cocconeis pseudolineata (Geitler) Lange-Bertalot O-C-OM 6.7 57.7 74.7 20 Cymatopleura solea var. apiculata (W.Smith) Ralfs C 0.4 1.5 0.8 3 Cymbella affinis Kützing O-C 1.7 8.7 12.5 5 Cymbella cymbiformis C.Agardh O-C 7.4 4.6 29.2 2 Cymbella excisiformis Krammer O 0.8 2.0 1.4 1 Cymbella diminuta (Grunow) Reichardt C 0.9 3.1 2.1 2 Cymbella hantzschiana Krammer O-C 1.7 13.8 4.7 10 Cymbella laevis Nägeli C 1.2 1.0 2.1 1 Cymbella parva (W.Smith) Kirchner O-C 1.3 12.2 5.5 9 Cymbella tridentina Lange-Bertalot, M.Cantonati et A.Scalfi O 5.1 3.1 20.1 2 Cymbella vulgata Krammer C 0.2 1.0 0.3 2 Cymbopleura amphicephala (Nägeli) Krammer O 0.8 2.6 1.6 1 Cymbopleura florentina (Grunow) K.Krammer O 0.4 1.5 0.7 1 Cymbopleura subaequalis (Grunow) Krammer C 0.7 0.5 0.7 1 Cymbopleura subaustriaca Krammer O-C-OM 1.9 5.1 7.0 3 Decussata hexagona (Torka) Lange-Bertalot C 0.4 0.5 0.4 1 Delicata minuta K.Krammer O-C 0.4 1.5 0.6 2 Denticula kuetzingii Grunow O-C 0.9 4.1 2.2 5 Denticula tenuis Kützing O-C 3.2 16.8 21.8 9 Humidophila contenta (Grunow) Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot et Kopalová O-C 0.5 8.7 1.9 9 Humidophila paracontenta var. magisconcava (Lange-Bertalot) Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot et Kopalová O-C 1.3 9.2 6.7 11 Humidophila perpusilla (Grunow) Lowe, Kociolek, Johansen,Van deVijver, Lange-Bertalot et Kopalová O-C 5.7 27.0 76.7 14 Humidophila sp. OM 0.5 0.5 0.5 1 Odontidium hyemale (Roth) Kützing O-C 1.1 3.6 2.2 5 Odontidium mesodon (Ehrenberg) Kützing O-C-OM 7.4 35.2 65.2 17 Diploneis elliptica (Kützing) Cleve O-C 1.0 4.6 4.1 4 Diploneis fontanella Lange-Bertalot O-C-OM 1.0 19.9 3.4 14 Diploneis krammeri Lange-Bertalot et E.Reichardt O-C-OM 1.3 26.0 13.2 16 Diploneis minuta J.B.Petersen O-C 0.5 3.1 0.7 3 Encyonema hebridicum (Gregory) Grunow O-C 0.6 2.6 1.2 3 Encyonema minutum (Hilse) D.G.Mann O-C 0.7 4.6 2.3 5 Encyonema neogracile Krammer O 0.4 0.5 0.4 1 Encyonema silesiacum (Bleisch) D.G.Mann O-C 0.7 5.6 1.6 5 Encyonema triangulum (Ehrenberg) Kützing O 0.6 1.0 0.7 1 Encyonema ventricosum (C.Agardh) Grunow O-C-OM 2.5 16.3 30.5 9 Encyonopsis cesatii (Rabenhorst) Krammer O-C 3.9 35.2 24.3 15 Encyonopsis fonticola (Hustedt) Krammer O-C 2.9 10.2 10.2 9 Encyonopsis krammeri Reichardt O-C-OM 1.0 16.8 5.5 12 Encyonopsis microcephala (Grunow) Krammer O 1.1 3.1 2.5 1 Encyonopsis minuta Krammer et E.Reichardt M 0.7 3.1 1.8 1 Epithemia argus (Ehrenberg) Kützing O-C-OM 1.4 2.0 4.3 1 Epithemia frickei Krammer O-C 1.0 1.5 2.2 3 Epithemia turgida (Ehrenberg) Kützing C 1.1 2.6 3.3 1 Eucocconeis flexella (Kützing) Meister O 0.6 1.5 0.9 2 Eucocconeis laevis (Østrup) Lange-Bertalot O-C-OM 0.7 6.1 2.7 8 Eunotia ambivalens Lange-Bertalot et Tagliaventi O 1.4 1.0 1.6 1 Eunotia arcubus Nörpel et Lange-Bertalot O-C 7.8 11.2 60.7 7 Eunotia soleirolii (Kützing) Rabenhorst O-OM 1.4 9.2 4.1 4 Fallacia lange-bertalotii (E.Reichardt) E.Reichardt C 0.7 2.0 1.2 2 Tab. 1. Continued KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 72 ACTA BOT. CROAT. 78 (1), 2019 Taxa GS A F max. N Fallacia subhamulata (Grunow) D.G.Mann C 1.4 4.1 7.8 4 Fragilaria amphicephaloides Lange-Bertalot O 1.4 2.0 1.9 1 Fragilaria recapitellata H. Lange-Bertalot et D. Metzeltin C 4.1 5.6 26.9 3 Frustulia vulgaris (Thwaites) De Toni C-OM 0.3 3.1 0.5 4 Gomphonema angustatum (Kützing) Rabenhorst C-OM 1.5 12.8 11.6 8 Gomphonema angustivalva E.Reichardt O-C 3.3 1.0 6.2 2 Gomphonema angustum C.Agardh O-C-OM 1.7 27.6 10.5 14 Gomphonema brebissonii Kützing O 0.4 1.0 0.4 1 Gomphonema elegantissimum Reichardt et Lange-Bertalot O-C 4.7 18.4 42.6 8 Gomphonema exilissimum (Grunow) Lange-Bertalot et E.Reichardt O-C 1.4 15.8 7.1 11 Gomphonema micropus Kützing O-C-OM 2.0 44.9 23.3 17 Gomphonema occultum E.Reichardt et Lange-Bertalot C 0.6 2.0 1.2 2 Gomphonema olivaceum (Hornemann) Brébisson C 0.6 4.6 1.8 2 Gomphonema olivaceoides Hustedt C 0.8 0.5 0.8 1 Gomphonema parvulum (Kützing) Kützing O-C-OM 1.8 18.4 11.7 15 Gomphonema procerum Reichardt et Lange-Bertalot O-C 0.2 1.5 0.5 2 Gomphonema pseudoaugur Lange-Bertalot O 0.3 0.5 0.3 1 Gomphonema pseudobohemicum Lange-Bertalot et E.Reichardt C 1.6 2.6 3.7 2 Gomphonema pumilum (Geitler) Bohunická et J.R.Johansen OM 3.1 24.5 46.0 12 Gomphonema subclavatum (Grunow) Grunow O-C-OM 0.6 12.2 1.3 10 Gomphonema tergestinum (Grunow) Fricke O-C 2.0 12.8 7.0 10 Grunowia sinuata (Thwaites) Rabenhorst O-C 1.6 3.1 5.9 2 Gyrosigma acuminatum (Kützing) Rabenhorst C 6.1 4.1 14.5 1 Gyrosigma obtusatum (Sullivant & Wormley) C.S.Boyer C 1.5 2.0 2.4 1 Halamphora normanii (Rabenhorst) Levkov O-C 1.2 3.1 3.6 4 Halamphora veneta (Kützing) Levkov O 0.7 0.5 0.7 1 Hannaea arcus (Ehrenberg) R.M.Patrick C 0.1 0.5 0.1 1 Hantzschia amphioxys (Ehrenberg) Grunow O-C 0.5 4.1 0.9 3 Luticola mutica (Kützing) D.G.Mann C-OM 0.8 3.1 2.0 4 Meridion circulare (Greville) C.Agardh O-C-OM 5.9 64.3 38.7 20 Navicula antonii Lange-Bertalot C 1.7 21.9 7.5 10 Navicula cariocincta Lange-Bertalot C 1.0 3.6 3.1 3 Navicula cataracta-rheni Lange-Bertalot O 2.0 6.1 8.9 3 Navicula cryptotenella Lange-Bertalot O-C-OM 1.7 12.8 22.4 8 Navicula cryptotenelloides Lange-Bertalot O 0.8 0.5 0.8 1 Navicula gregaria Donkin C 1.1 0.5 1.1 1 Navicula leistikowii Lange-Bertalot O-C 0.9 13.8 2.7 9 Navicula oblonga (Kützing) Kützing O 0.6 0.5 0.6 1 Navicula radiosa Kützing O-C 1.2 17.9 4.0 11 Navicula sp. C 1.9 4.1 10.3 3 Navicula tripunctata (O.F.Müller) Bory de Saint-Vincent O-C 1.2 17.3 6.2 9 Naviculadicta geisslerae (Jahn) Jahn O 0.7 0.5 0.7 1 Naviculadicta tridentula (Krasske) Lange-Bertalot O 0.6 2.6 1.6 3 Naviculadicta sp. C 4.4 0.5 4.4 1 Neidiomorpha binodiformis (K.Krammer) M.Cantonati, H.Lange-Bertalot et N.Angeli O-C 0.3 1.5 0.4 2 Neidium longiceps (Gregory) Ross O-C 0.3 2.0 0.5 3 Nitzschia amphibia Grunow O-C 1.6 7.1 7.2 8 Nitzschia capitellata Hustedt O-C 0.8 5.6 1.8 7 Nitzschia dissipata (Kützing) Grunow O-C 1.0 15.3 4.1 11 Nitzschia dubia W.Smith O-C-OM 2.1 14.3 11.2 7 Nitzschia fonticola (Grunow) Grunow C 4.5 10.2 33.5 6 Nitzschia frustulum (Kützing) Grunow O-C-OM 1.2 10.2 3.4 9 Nitzschia heufleriana Grunow C 0.6 1.5 0.8 1 Nitzschia inconspicua Grunow O-C 0.7 3.6 1.3 6 Nitzschia linearis (C.Agardh) W.Smith O-C 2.7 31.6 58.5 16 Nitzschia palea (Kützing) W.Smith O-C 2.4 14.8 12.0 12 Nitzschia palea var. tenuirostris Grunow C-OM 4.8 7.1 31.6 5 Nitzschia oligotraphenta (Lange-Bertalot) Lange-Bertalot C 0.5 0.5 0.5 1 Nitzschia sp.1 C 2.6 0.5 2.6 1 Nitzschia sp.2 O 0.9 0.5 0.9 1 Nitzschia umbonata (Ehrenberg) Lange-Bertalot C 1.1 0.5 1.1 1 Tab. 1. Continued ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 73 Taxa GS A F max. N Nupela lapidosa (Krasske) Lange-Bertalot O 0.7 0.5 0.7 1 Orthoseira roeseana (Rabenhorst) O'Meara O-C 0.7 2.0 1.4 3 Pinnularia acutobrebissonii Kulikovskiy, Lange-Bertalot et Metzeltin O 0.9 2.0 1.1 1 Pinnularia borealis Ehrenberg O 0.5 2.6 0.7 2 Pinnularia gibba Ehrenberg C 0.4 0.5 0.4 1 Pinnularia kuetzingii K.Krammer O 0.7 2.0 0.8 1 Pinnularia sp. OM 0.5 0.5 0.5 1 Pinnularia subcapitata var. elongata Krammer O 1.0 2.0 2.0 1 Pinnularia subrupestris K.Krammer O-C-OM 1.1 6.1 3.3 7 Placoneis paraelginensis Lange-Bertalot O-M 0.3 3.1 0.5 5 Platessa conspicua (A. Mayer) Lange-Bertalot M 2.8 1.5 8.1 1 Platessa lutheri (Hustedt) Potapova C 0.9 1.5 1.6 1 Planothidium dubium (Grunow) Round et Bukhtiyarova M 1.4 22.4 10.3 8 Planothidium frequentissimum (Lange-Bertalot) Round et L.Bukhtiyarova O-C-OM 1.8 21.9 6.6 18 Planothidium joursacense (Héribaud-Joseph) Lange-Bertalot O 3.6 0.5 3.6 1 Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot  O-C-OM 10.7 86.7 64.9 20 Planothidium minutissimum (Krasske) E.A.Morales C-OM 10.1 1.0 15.8 2 Planothidium reichardtii Lange-Bertalot et Werum O-C 1.0 5.1 3.9 8 Platessa holsatica (Hust.) Lange-Bertalot C-OM 4.4 4.1 28.9 5 Psammothidium grischunum (Wuthrich) L.Bukhtiyarova et Round C 3.7 13.3 15.7 6 Psammothidium subatomoides (Hustedt) Bukhtiyarova et Round O-C-OM 1.5 8.7 10.7 7 Pseudostaurosira brevistriata (Grunow) Williams et Round C 0.1 0.5 0.1 1 Reimeria sinuata (Gregory) Kociolek et Stoermer C 0.3 1.0 0.4 1 Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot C 0.7 9.2 2.9 5 Rhopalodia parallela (Grunow) O. Müller O-C 0.5 5.1 0.8 3 Rossithidium anastasiae (Kaczmarska) Potapova O-C-OM 2.8 11.2 15.3 7 Rossithidium petersenii (Hustedt) Round et Bukhtiyarova O-C 1.4 6.6 3.0 6 Rossithidium pusillum (Grunow) Round et L.Bukhtiyarova C 0.2 1.0 0.3 1 Sellaphora seminulum (Grunow) D.G. Mann O-C 0.6 4.6 1.7 6 Sellaphora pseudopupula (Krasske) Lange-Bertalot C 0.4 1.5 0.7 3 Sellaphora pupula (Kützing) Mereschkovsky C 0.7 9.2 2.4 6 Sellaphora stroemii (Hustedt) H.Kobayasi O 0.6 0.5 0.6 1 Sellaphora nigri (De Notaris) C.E. Wetzel et L. Ector O-C-OM 2.9 25.0 22.5 17 Simonsenia delognei (Grunow) Lange-Bertalot C 1.1 1.5 3.1 1 Staurosirella martyi (Héribaud) Morales et Manoylov C 0.1 0.5 0.1 1 Stauroneis anceps Ehrenberg O-OM 0.3 2.6 0.4 3 Stauroneis jarensis Lange-Bertalot O 0.3 0.5 0.3 1 Stauroneis phoenicenteron (Nitzsch) Ehrenberg C 0.3 0.5 0.3 1 Stauroneis smithii Grunow O-C 0.8 6.1 3.6 5 Surirella angusta Kützing O-C-OM 1.1 10.2 5.0 9 Surirella brebissonii var. kuetzingii Krammer et Lange-Bertalot O 0.2 0.5 0.2 1 Surirella linearis W.Smith O-C 2.1 18.4 12.8 12 Surirella minuta Brébisson O-C 1.0 4.6 2.9 5 Surirella spiralis Kützing O 0.7 0.5 0.7 1 Surirella terricola Lange-Bertalot et E.Alles O-C-OM 0.9 8.7 3.3 9 Tryblionella angustata W.Smith O-C 3.0 10.7 13.9 11 Tryblionella angustatula (Lange-Bertalot) Cantonati et Lange-Bertalot O 1.2 3.6 3.1 3 Ulnaria ulna (Nitzsch) P.Compère O-C-OM 0.8 24.5 4.1 16 Chlorophyceae Chaetophora sp. O 12.9 1.0 20.0 1 Microspora sp. O 0.6 0.5 0.6 1 Klebsormidiophyceae Klebsormidium flaccidum (Kützing) P.C. Silva, K.R. Mattox et W.H. Blackwell O 3.2 1.5 5.1 1 Ulvophyceae Ulothrix sp. O 0.3 0.5 0.3 1 Ulotrichales C 0.8 1.0 1.7 1 Conjugatophyceae Cosmarium obtusatum (Schmidle) Schmidle O 0.1 1.5 0.1 1 Spirogyra sp. C 0.5 1.5 1.0 1 Tab. 1. Continued KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 74 ACTA BOT. CROAT. 78 (1), 2019 lite springs on the other side of the plot (Fig. 3). The highest Shannon-Wiener diversity and the highest number of taxa were recorded from the epibryon sample of the spring SK on ophiolitic substratum (2.95 and 47, respectively). The low- est diversity (0.08) and the lowest number of taxa (5) were found in the epilithon summer sample in the spring TU on carbonate substratum. The rheocrenic spring ST, character- ised by the species Tapinothrix varians and Cocconeis linea- ta, had the lowest values of Pielou's Index, indicating several dominant and subdominant species in the sample and huge unevenness in species abundances. Parametric t-test per- formed on diversity indices of springs on different geologi- cal substrata did not show statistical differences (p > 0.05). However, statistical differences for used trophic indices be- tween two groups of springs were found (Rott's Trophic In- dex – TI, t = – 2.4, p = 0.02, Croatian Diatom Trophic Index – TIDHR, t = 2.75, p = 0.007, and the Ecological quality ratio – EQR which is based on TIDHR). Higher mean values of tro- phic indices were noted in samples from ophiolites (Tab. 2). Spring ZL on ophiolitic melange was not considered for sta- tistical analysis as it was the sole site on this geological sub- stratum. Values of TI varied between 1.22 (spring MV) and 2.59 (spring ZL). Meso-eutrophic and eutrophic conditions were detected for eleven and eight springs, respectively. Ac- cording to the TI, an oligotrophic status was assessed only for the spring MV. Assessment of ecological status of springs based on values of EQR and TIDHR indicated good status for all localities, except for the spring MV (excellent status) and spring PO (relatively good status) (Tab. 2). Values of pH preferences were found for 58.3% of identi- fied diatoms: 51.4% alkaliphilous (most frequent taxa: Am- phora pediculus, Cocconeis lineata, C. euglypta, Gomphonema angustum, G. micropus, Meridion circulare, Navicula tri- punctata, Planothidium lanceolatum), 33.9% circumneutral (e.g., A. minutissimum, Odontidium mesodon, Encyonema minutum, Encyonopsis cesatii, Navicula radiosa, etc.), 9.2% acidophilous (e.g. Neidium affine var. longiceps, Pinnularia subcapitata var. elongata, Tryblionella angustata), and 5.5% alkalibiontic (rarely occurring species such as Achnanthid- ium exile and Halamphora veneta). Moisture-condition preference indicator values were found for 50.2% of the diatom taxa: 17% of taxa were clas- sified as ‘never, or only very rarely occurring outside water bodies’ (Meridion circulare, Nitzschia fonticola, Ulnaria ul- na), 21.3% occurred ‘mainly in water bodies, sometimes on wet places’. The highest number of species (46.8%) ‘occurred mainly on wet and moist places’, while a low number of spe- cies (11.7%) ‘occurred mainly on wet and moist or temporar- ily dry places’ (Caloneis tenuis, Diploneis fontanella, Luticola mutica). The lowest number of taxa (3.2%) belongs to the category ‘occurring nearly outside of water bodies’ (Adlafia bryophila, Diploneis minuta, Halamphora veneta). Geological preferences Geological preferences of algae for the two main geo- logical substrata, carbonates and ophiolites were analyzed. As only one site on this geological substratum was available, the spring on ophiolitic melange could not be included in statistical analyses. A total of 49 species (31%) were found exclusively in samples taken from springs on ophiolites, and were mainly identified from one site only, and the most common (relative frequency > 3%) were: Navicula catarac- ta-rheni, Chamaesiphon polonicus, Tryblionella angustatula, Cymbella tridentina, Cocconeis disculus and Encyonopsis mi- crocephala. A total of 73 taxa (40%) were found exclusively in samples from carbonate substratum, and the most com- mon (frequency > 5%) were: Planothidium dubium, Navicula antonii, N. cariocincta, Psammothidium grischunum, Gom- phonema angustatum, Rhoicosphenia abbreviata, Sellaphora pupula, Vaucheria sp., Nitzschia palea var. tenuirostris, N. fon- ticola, Fragilaria recapitellata, Audouinella sp., Gomphonema olivaceum, Gyrosigma acuminatum, Phaeodermatium rivu- lare, Platessa holsatica, Cymbopleura diminuta, Encyonopsis minuta and Phormidium tinctorium. Results of the IndVal analysis of algal assemblages for the factor ‘lithology’ (Tab. 3) comprised statistically signif- icant (p ≤ 0.01) indicator species with algal frequency in the samples > 10%. Although many species were identified in the springs on both investigated substrata, they showed higher abundance or frequency on one substratum. A pref- erence for ophiolitic substrata, with IndVal values > 30, was established for: Achnanthidium minutissimum, Humidophi- la perpusilla, Diploneis krammeri, Encyonopsis cesatii, Gom- phonema elegantissimum, Navicula radiosa, Planothidium lanceolatum and Nitzschia linearis. By contrast, a preference Fig. 3. Non-metric multidimensional scaling (NMDS) of sites (pre- sented by spring codes) based on Bray-Curtis matrix of similari- ties of algal assemblages in relation to geological substratum and clusters. Trophic preferences were found for 101 species (54%), and were as follows: oligotraphentic (13.95%, Encyonopsis cesatii, Gomphonema angustum, Surirella spiralis), oligo-me- sotraphentic (13.9%), mesotraphentic (9.9%), meso-eutrap- hentic (12.9%), eutraphentic (32.75%, Amphora copulata, A. pediculus, Cocconeis spp., Navicula tripunctata), hypereutra- phentic (3%, Nitzschia palea, Nitzschia umbonata) and oligo – to eutraphentic (13.9%). ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 75 Tab. 2. The values of diversity and trophic indices of studied springs on the Mt. Konjuh with ranges of minimum (min), maximum (max), mean and standard deviation (SD), t – test values and p – statistical significance for groups of springs on carbonates (101 samples) and ophiolites (95 samples), N – number of samples per spring, S – number of taxa, J' – Pielou's index, H'(ln) – Shannon – Wiener diversity index, TI Rott – Rott's trophic index, TIDHR – Croatian trophic index, EQR – ecological quality ratio with water quality ranges. Substrata Spring code N S H’ J' TI Rott TIDHR EQR ophiolites 1KE 12 48 1.65 0.63 2.61 2.23 0.71 (II) 2KE 12 46 1.50 0.6 2.14 2.31 0.67 (II) 1BO 9 55 1.90 0.67 2.40 2.41 0.64 (II) 2BO 6 63 1.93 0.57 1.77 2.48 0.61 (II) 1VU 9 63 2.07 0.65 2.32 2.49 0.6 (II) 2VU 6 49 2.01 0.66 2.29 2.44 0.62 (II) MI 5 56 2.09 0.7 2.14 2.28 0.69 (II) SK 10 76 2.22 0.69 2.32 2.41 0.64 (II) KA 9 54 1.76 0.62 2.03 2.30 0.68 (II) min. 6 0.32 0.14 1.39 1.49 0.49 (III) max. 47 2.95 0.91 3.27 2.77 1 (I) mean 19.21 1.79 0.62 2.23 2.36 0.65 (II) SD   9.09 0.59 0.17 0.49 0.19 0.77 carbonates GB 12 66 2.04 0.7 2.32 2.19 0.72 (II) KR 8 62 1.34 0.47 2.32 2.21 0.72 (II) KS 12 64 1.77 0.64 2.31 2.16 0.74 (II) MV 9 54 1.69 0.58 1.22 1.76 0.9 (I) PO 12 70 1.58 0.56 1.89 2.54 0.59 (III) ST 12 46 1.30 0.51 1.90 2.10 0.76 (II) TA 12 55 1.30 0.5 1.98 2.28 0.69 (II) TU 12 65 1.50 0.52 2.11 2.38 0.65 (II) UB 8 67 2.23 0.74 2.56 2.41 0.63 (II) ZP 12 58 2.23 0.75 2.19 2.34 0.66 (II) min. 5 0.08 0.03 0.70 1.05 0.47 (III) max. 43 2.83 0.91 3.18 2.82 0.96 (I) mean 19.19 1.74 0.61 2.06 2.24 0.7 (II) SD   9.23 0.67 0.2 0.51 0.34 0.13 ophiolitic melange (mean) ZL 9 48 1.61 0.61 2.57 2.25 0.7 (II) t (ophiolites vs. carbonates) –0.02 –0.56 –0.56 –2.4 2.75 –2.75 p (ophiolites vs. carbonates)   0.99 0.57 0.57 0.02 0.007 0.007 for carbonate substrata (IndVal > 30) was noted for the fol- lowing species: Amphora pediculus, Odontidium mesodon, Gomphonema micropus, G. pumilum, Navicula antonii, and Planothidium dubium. Discussion Diatoms were the dominant algal group in the investi- gated springs. Our results were mostly compared with those obtained from the Alps, due to similarities in the sampling approach based on different spring types. A comparison of the presented results with previous investigations conducted in the south-eastern Alps (Cantonati 1998) revealed many species in common, such as Achnanthidium minutissimum, Odontidium (Diatoma) mesodon, Achnanthidium pyrenai- cum, Cocconeis euglypta, Gomphonema pumilum s.l. and Meridion circulare. However, several species frequent in al- pine springs (Brachysira brebissonii, Encyonema minutum, and Odontidium hyemalis) were either rarely present or not detected in our study. A higher similarity was found with the results of Angeli et al. (2010), who investigated diatoms in anthropogenically influenced, low altitude springs, where many eutraphentic taxa were recorded (Cocconeis, Planothid- ium, and Nitzschia). The highest number of taxa identified, excluding diatoms, belongs to the genera Phormidium and Chamaesiphon, which is in accordance with the results of Cantonati et al. (1996). Several species frequently recorded in this study (Chlorogloea microcystoides, Chamaesiphon po- lonicus, Tapinothrix varians, Audouinella sp., Vaucheria sp. and Tribonema sp.) were also found in previous investiga- tions (e.g., Cantonati et al. 2012c). On the other hand, spe- cies frequently noted in alpine springs, such as Xenotholos kerneri, Hydrurus foetidus, Chamaesiphon starmachii, Tapi- nothrix janthina and Plectonema tomasinianum were not re- corded in our study, presumably due to their preference for higher elevations with lower water temperature and/or sili- ceous substratum. For instance, a typical rheobiontic species of crenic habitats, Hydrurus foetidus, was not found in this study, possibly due to the lower water velocity of the springs investigated. Our results can be well compared with those KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 76 ACTA BOT. CROAT. 78 (1), 2019 Tab. 3. Indicator values (IndVal) and frequency of indicative species of ophiolitic and carbonate springs on the Mt. Konjuh at p <0.01 (4999 permutations using Monte Carlo test). Taxa IndVal Frequency in group 1 Frequency in group 2 gr ou p 1 op hi ol ite s Achnanthidium minutissimum (Kützing) Czarnecki 58.8 96 81 Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot  58.5 92 82 Encyonopsis cesatii (Rabenhorst) Krammer 50.5 58 22 Humidophila perpusilla (Grunow) Lowe, Kociolek, 46.8 49 14 Nitzschia linearis (C. Agardh) W. Smith 40 47 23 Diploneis krammeri Lange-Bertalot et E. Reichardt 38.9 45 13 Navicula radiosa Kützing 34.6 37 6 Gomphonema elegantissimum E. Reichardt et Lange-Bertalot 30.6 33 9 Cymbella hantzschiana Krammer 28.1 29 4 Gomphonema parvulum (Kützing) Kützing 23.8 27 12 Gomphonema tergestinum (Grunow) Fricke 23.8 27 4 Diploneis fontanella Lange-Bertalot 22.5 32 12 Encyonopsis krammeri Reichardt 21.1 26 11 Gomphonema exilissimum (Grunow) Lange-Bertalot et E. Reichardt 20.9 26 10 Amphora lange-bertalotii var. tenuis Levkov et Metzeltin 20.4 22 3 Navicula leistikowii Lange-Bertalot 20.2 24 7 Encyonopsis fonticola (Hustedt) Krammer 19.3 21 4 Cymbella parva (W.Smith) Kirchner 17.5 21 7 Achnanthidium exile (Kützing) Heiberg 16.8 19 3 Navicula cataracta-rheni Lange-Bertalot 15.4 15 0 Tryblionella angustata W. Smith 15.1 18 6 Chlorogloea microcystoides Geitler 14.6 17 5 Pseudanabaena sp. Lauterborn 13.6 14 1 Nitzschia amphibia Grunow 13.3 14 3 Surirella terricola Lange-Bertalot et E. Alles 12.6 14 5 Eunotia soleirolii (Kützing) Rabenhorst 11.5 12 0 Rhopalodia parallela (Grunow) O. Müller 10.7 12 1 Encyonema silesiacum (Bleisch) D.G.Mann 10.1 12 2 Diploneis elliptica (Kützing) Cleve 9.9 10 1 Eucocconeis laevis (Østrup) Lange-Bertalot 9.6 10 3 Nitzschia capitellata Hustedt 9.2 12 2 Tryblionella angustatula (Lange-Bertalot) Cantonati et Lange-Bertalot 9.0 9 0 Hantzschia amphioxys (Ehrenberg) Grunow 8.3 9 1 gr ou p 2 ca rb on at es Amphora pediculus (Kützing) Grunow ex A. Schmidt 56.1 50 83 Odontidium mesodon (Ehrenberg) Kützing 44.3 18 50 Navicula antonii Lange-Bertalot 39.4 0 39 Planothidium dubium (Grunow) Round et Bukhtiyarova 39.4 0 39 Gomphonema micropus Kützing 34.7 29 52 Gomphonema pumilum (Geitler) Bohunická et J. R. Johansen 31.2 12 35 Achnanthidium pyrenaicum (Hustedt) H. Kobayasi 29.1 21 38 Navicula tripunctata (O. F. Müller) Bory de Saint-Vincent 26.1 3 29 Psammothidium grischunum (Wuthrich) L. Bukhtiyarova et Round 23.9 0 24 Nitzschia oligotraphenta (Lange-Bertalot) Lange-Bertalot 22.5 4 24 Achnanthidium affine (Grunow) Czarnecki 20.0 5 22 Gomphonema angustatum (Kützing) Rabenhorst 18.3 0 18 Nitzschia fonticola (Grunow) Grunow 18.3 0 18 Navicula cryptotenelloides Lange-Bertalot 18.0 4 19 Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot 16.5 0 17 Sellaphora pupula (Kützing) Mereschkovsky 16.5 0 17 Vaucheria sp. A. P. de Candolle 15.6 0 16 Phormidium retzii Kützing ex Gomont 14.9 3 16 Nitzschia palea var. tenuirostris Grunow 11.0 0 11 Fragilaria recapitellata H. Lange-Bertalot et D. Metzeltin 10.1 0 10 ALGAE IN SPRINGS OF DIFFERENT LITHOLOGY ACTA BOT. CROAT. 78 (1), 2019 77 from 16 springs and streams of the Dolomiti Bellunesi Na- tional Park in Northern Italy (Cantonati 2008), where a to- tal of 65 algal taxa apart from diatoms, and 1–12 taxa per spring location were recorded. The exceptional heterogeneity of springs, resulting in richness of rare and endangered spe- cies already highlighted in several studies (Aboal et al. 1998, Werum and Lange-Bertalot 2004, Cantonati et al. 2006) was confirmed for the area investigated. Overall, one third of all taxa identified in the present study (36.3%) was noted in one spring only, which is slightly lower than 40% recorded by Cantonati and Spitale (2009) and clearly less than 58% noted by Bertrand et al. (2004). All of the aforementioned studies refer to springs as unique habitats, each requiring a specific diversity estimation. Hierarchical clustering based on relative algal abun- dance clustered springs into six groups. These results were in partial agreement with Cantonati et al. (2012b, c), whose spring classification included 6 diatom-based and 7 groups based on algal taxa except diatoms. Species typical for rheo- crenes on carbonates, such as Achnanthidium pyrenaicum and Nitzschia fonticola (Cantonati et al. 2012b), have been detected with high abundance in the spring MV. Groups of rheocrenes on carbonates (3rd and 5th cluster) showed great- est similarity with rheocrenes on carbonate substrata with NO3– enrichment or shading described by Cantonati et al. (2012b). Diatom species shared by Alpine and Dinaric car- bonate springs were Odontidium mesodon, Achnanthidium pyrenaicum, Amphora pediculus, Meridion circulare, Cocco- neis lineata, C. pseudolineata, and Sellaphora nigri. This con- gruence was expected, since the measured increased nitrate concentrations in these springs determined a higher number of eutraphentic species (e.g., Amphora pediculus). The cyano- prokaryote Tapinothrix varians and the red alga Audouinella spp. showed affinity for shaded springs, as detected in some earlier studies (Cantonati et al. 1996, Cantonati et al. 2012c). However, certain differences were observed when the springs on ophiolites were compared with springs emerging from si- liceous aquifers. In a comparison of algal assemblages, the highest resemblance was observed between rheocrenes on ophiolites and group 4 (rheocrenes on carbonate substrata with lower conductivity or seasonal desiccation) by Canto- nati et al. (2012b). Species in common were Planothidium frequentissimum, Achnanthidium dolomiticum, Meridion cir- culare and several indicators of desiccation, e.g. Humidophila perpusilla and H. contenta. Moreover, two new records for the diatom flora of Bosnia and Herzegovina were found in the investigated ophiolite springs: Achnanthidium dolomiti- cum, a species previously recorded in springs on dolomite substrata (Cantonati and Lange-Bertalot 2006), and Cym- bella tridentina, a rheophilic species of carbonate headwaters (Cantonati et al. 2010). A common feature of dolomite and ophiolite springs is a relatively high magnesium concentra- tion, presumably a key determinant for the distribution of A. dolomiticum. The largest group, rheohelocrenic springs on ophiolites, was compatible with hygropetric rheocrenes on carbonate substrata in Cantonati et al. (2012b), with sev- eral species in common: Denticula tenuis, Encyonopsis ce- satii, and E. microcephala. Despite different lithology, springs on ophiolites and springs on carbonate substrata had a very similar algal composition. The greatest concurrence in algal communities recorded in this study with springs on siliceous substrata from previous studies was noted for the sole spring on ophiolitic melange (ZL), with several species of Gompho- nema and Eunotia in common. The influence of lithology on algal assemblages has been emphasized by several authors (Cantonati 1998, Werum 2001, Cantonati et al. 2006, Wojtal 2013), typically consid- ering springs on different, carbonate and siliceous, litholo- gies. Moreover, phycological studies focusing on springs on ophiolites (silicate rocks) were very rare. When algal assem- blages of springs on peridotite and serpentinite (ophiolites) from this study were compared with springs on amphibo- lite in Germany (Werum 2001), a high similarity in species composition was noted. However, many species commonly occurring on ophiolites in this study, such as A. minutissi- mum, Humidophila perpusilla, Diploneis fontanella, Nitzschia linearis and Planothidium lanceolatum, were not good indi- cators of aquifer lithology because of their widespread dis- tribution (Werum 2001). On the contrary, species that had lower abundance and higher frequency, as detected by Ind- Val analysis, proved to be more reliable indicators of an aqui- fer’s lithology. The IndVal analysis singled out the following species as having a preference for ophiolites: Amphora lange- bertalotii var. tenuis, Cymbella hantzschiana, C. parva, Encyo- nopsis cesatii, E. krammeri, Gomphonema parvulum, G. terg- estinum and Navicula leistikowii. Some of the species listed are typical for calcareous fens, such as E. cesatii (Fránková et al. 2009), others are widespread such as G. parvulum (Abar- ca et al. 2014). Results of the indicator-species analysis for carbonate substrata were in accordance with previous studies (Can- tonati et al. 2012b, c), in which several rheophilic species (Achnanthidium pyrenaicum, Odontidium mesodon, Gom- phonema micropus, G. pumilum, Planothidium dubium) or species with a preference for nitrogen enrichment (Na- vicula tripunctata and Amphora pediculus) were noted. A low resemblance has been found when our results on ophi- olite-spring algal assemblages were compared with results from previous investigations in springs emerging from oth- er types of siliceous substrata such as granite and rhyolite (Werum 2001, Kapetanović and Hafner 2007). Species typi- cal for rheocrenes and helocrenes on siliceous substrata of the south-eastern Alps (Cantonati et al. 2012c) or detected in the Pyrenees (Sabater and Roca 1992), were not found in the ophiolite springs studied. In the latter, genera typical for siliceous substrata, such as Tabellaria, Eunotia and Pinnular- ia, were either missing or occurring only with a few species. Nevertheless, a large number of shared species (110) suggests fewer differences in the structure of algal communities be- tween carbonates and ophiolites (this study) than between ultramaphites (this study) and other types of siliceous sub- strata (e.g. granite, rhyolite; algal community composition data obtained from the literature). In addition, a species typi- cal for carbonate substrata, Tapinothrix varians (Cantonati KAMBEROVIĆ J., PLENKOVIĆ-MORAJ A., KRALJ BOROJEVIĆ K., GLIGORA UDOVIČ M., ŽUTINIĆ P., HAFNER D., CANTONATI M. 78 ACTA BOT. CROAT. 78 (1), 2019 et al. 1996), was found in all springs regardless of lithologi- cal substratum, and especially in springs with higher current velocity. On the other hand, Tapinothrix janthina, the vicari- ant species typical for springs on silicates, was not found in the present study. Our results suggest that algal assemblages in springs emerging from ophiolites and springs emerging from other types of siliceous substrata should be separately analyzed in future studies. The total number of species in springs can greatly de- pend on the lithology or the heterogeneity of microhabitats. Nascimbene et al. (2011) recorded higher numbers of dia- toms, plants and lichens from springs on siliceous substra- ta, whilst other algal groups (including cyanoprokaryotes) were more numerous on carbonate substrata. On the other side, Frankovà et al. (2009) pointed out that the number of diatom species decreased significantly from mineral-rich to mineral-poor spring fen sites. Although a higher number of taxa was noted in springs on carbonates than in those on ophiolites, differences between averages of diversity indices were not significant. The total number of diatom taxa found in the 20 springs of Mt. Konjuh (187) was higher than in 17 Swiss springs (118 taxa; Taxböck and Preisig 2007), very sim- ilar to Pyrenean springs (194 taxa; Sabater and Roca 1992), and lower than in 30 springs in south-eastern Alps (250 taxa; Cantonati 1998). Although the average value of the Shan- non-Wiener Diversity Index indicated a moderately high diversity of algal species (H` = 1.75), it was lower than the value obtained for Alpine springs (H` = 2, Cantonati 1998). Several authors emphasized a relationship between lower al- titudes and higher nitrate concentrations in springs (Aboal et al. 1998, Cantonati and Spitale 2009), which can result in a decrease of diatom species (Bertrand et al. 2004). A wide range in the total number of species per individual sample (from 5 to 47) and a high number of taxa per spring location (from 46 to 76) indicate the need to sample different micro- habitats to achieve a more accurate biodiversity assessments of spring habitats. The algal assemblages in wetland habitats around springs and streams of the Vranica Mountain, Bosnia and Herze- govina (Kapetanović and Hafner 2007), especially those on siliceous substrata, differ from those of the springs studied on Mt. Konjuh by the preponderance of acidophilous taxa (Brachysira brebissonii, Eunotia incisa, Tabellaria flocculo- sa, and many species from the genera Neidium and Euno- tia). Acidophilous species were rarely found on Mt. Kon- juh, due to ultramafic and calcareous lithologies resulting in higher pH values. On the other hand, many eutraphen- tic species were noted (32.7%). This proportion was higher than that recorded by Cantonati and Spitale (2009) in the Dolomiti Bellunesi National Park in Italy (22%). Altough natural diatom distribution could be altered by antrhopo- genic influences, Wojtal (2013) pointed out that springs with high nutrient concentrations also show heterogene- ity of diatom assemblage composition indicating the great potential of springs for biodiversity assessment. Angeli et al. (2010) pointed out that any morphological alteration of springs leads to great disturbance of their diatom communi- ties, which was also the case in many springs studied here- in. Phormidium retzii, an indicator of nitrogen enrichment (Cantonati et al. 2012c), was found in seven springs on Mt. Konjuh, indicating anthropogenic impact and a deteriorat- ed ecological status. Values of Rott’s TI were high due to the dominance of highly competitive meso- and eutraphentic algal species, which is compatible with the results of some previous in- vestigations of low-altitude springs in Germany (Werum 2001), and anthropogenically impacted springs in Austria (Gesierich and Kofler 2010), thus emphasizing that an in- crease in nitrate leads to decreasing numbers of sensitive and rare species. The mean values of the Rott TI (2.06 and 2.23 in carbonate and ophiolitic springs, respectively) belong in the same trophic class as a half of the investigated springs from the River Adige basin, and are higher than Rott’s TI from springs of Trentino (Angeli et al. 2010). Adversely, the EQR and TIDHR values indicated for one level lower trophic class than Rott’s TI in the present investigation, presumably due to geographical adaptation of the species indicator val- ues. However, the use of indices developed for rivers in the trophic-status assessments of springs could undergo certain limitations, in particular in the case of spring types markedly differing from streams. Conclusion The study provides the first information on the biodi- versity of algal communities in the mountain springs of north-eastern Bosnia and Herzegovina. Although springs had long been neglected in hydrobiological investigations, recent studies emphasize the importance of these habitats in biodiversity conservation. Despite the visible anthropogenic impacts in springs of Mt. Konjuh, a moderately high diver- sity of algal species per spring location was detected. Results of this study revealed that springs on ophiolites should be analyzed separately from springs on other siliceous substrata in future studies dealing with community structure in dif- ferent spring types. Due to the high heterogeneity of spring habitats, in order to preserve spring species diversity of Mt. Konjuh the conservation of springs as a group of habitats is required, and not only the protection of individual spring sites. Special focus should be directed towards preserving small rheohelocrenic springs, which are usually ignored in management and protection plans, even though they have a great potential in species conservation. Acknowledgements We are grateful to Elvir Babajić for help in the identifica- tion of the lithology of substrata, to Selvir Kamberović for help in data processing, and the Federal Ministry of Educa- tion and Science in Bosnia and Herzegovina for providing funding for a research visit to the MUSE – Museo delle Sci- enze, Trento, Italy. 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