Diatom assemblages on Shara and Nidze Mountains, Macedonia

501

Nova Hedwigia 81 3mdash4 501mdash537 Stuttgart November 2005

DOI 1011270029-503520050081-0501 0029-5035050081-0501 $ 925copy 2005 J Cramer in der Gebruumlder Borntraeger

Verlagsbuchhandlung D-14129 Berlin middot D-70176 Stuttgart

Diatom assemblages on Shara and Nidze Mountains Macedonia

by

Zlatko Levkov Svetislav Krstic Teofil Nakov and Ljupco Melovski

Institute of Biology Faculty of Natural SciencesGazi Baba bb PO Box 162 MK-1000 Skopje Republic of Macedonia

With 64 figures and 2 tables

Levkov Z S Krstic T Navkov amp L Melovski (2005) Diatom assemblages on Shara and NidzeMountains Macedonia - Nova Hedwigia 81 501-537

Abstract The diatom microflora collected from glacial lakes peat-bogs springs streams riversand wetlands on 352 localities on Shara Mountain and 124 localities on Nidze Mountain Macedoniawas investigated between 1995 and 2003 A total of 418 taxa from 76 genera were identified Thetaxa are predominantly oligotrophic or dystrophic Total floristic similarity between the neighbouringmountain sytems (100 km apart) was only 2274 One of the sites has been influenced by glaciationwhile the other contained habitats that served as glacial refugia Taxa from the refugia are highlysimilar to the those of the ancient Ohrid and Prespa lakes in Macedonia The morphological featuresand taxomic position of 11 taxa are discussed

Key words Diatoms mountains Nidze Shara refugia glacial lakes

Introduction

Research on biodiversity has become an important focus in the last decade of the 20th

century primarily due to increasing evidence of the loss of biodiversity on a globalscale (Pimm et al 1995) At the same time there has been an increase in the recognitionof the scope of diatom biodiversity This recognition is partly the result of theimplementation of molecular techniques (Medlin et al 1991) more sophisticatedEM microscopy (Haringkansson 2002) and implementation of more detailed morpho-metric analysis of the diatom valve (Lange-Bertalot 2001) In particular the workof Lange-Bertalot and co-workers has trended towards a finer designation at thespecies level Such efforts have established numerous taxa grouped in ldquosippenrdquo orspecies complexes which are very difficult to separate morphologically or ecologically

Corresponding author zlevkoviunonapmfukimedumk

502

as in the case of Navicula Bory sensu stricto (Lange-Bertalot 2001) or PinnulariaEhrenberg (Krammer 2000) Also Round et al (1990) triggered an intensive changein diatom nomenclature by the transfer of numerous taxa into an expanded number ofdiatom genera This activity has had a variety of results In some cases the result wasthat a taxon became assigned to a different more restricted genus (Round amp Bukhtiyarova1996 Kingston 2000) or to the same genus but by different authors (Andresen et al2000) This practice has been criticized (Kociolek 1996 Lange-Bertalot 2001) becauseof higher level transfers that are seen as simplistic and lack a proper investigation oftype specimens Type material revisions enable a much more realistic view ontaxonomy ecology and distribution of specific taxa (Sterrenburg 1994 1995) as wellas finding possibly new previously overlooked species (Cox 2003)

Revision of data on the distribution of certain cosmopolitan taxa such as Amphoracoffeaeformis (CAgardh) Kuumltzing in Argentina (Sala et al 1998) or StephanodiscusEhrenberg species (Haringkansson amp Locker 1981) brings to light possiblemisidentification mainly caused by the application of European flora keys to materialsfrom entirely different geographic areas and therefore fitting the species into knownspecies (Kociolek amp Spaulding 2000) The result has been severe criticism of thecosmopolitan distribution of diatoms (Mann amp Droop 1996 Snoeijs amp Potapova1998) and gradual abandonment of the concept in the latest revisions of specificgenera (Krammer 2000 2002 2003 Lange-Bertalot 2001) Cosmpolitian diatomdistribution can be expected in North Europe where aquatic ecosystems have repeatedlybeen influenced by glaciation events (Mann 1999 Kociolek amp Spaulding 2000)while the only centres of diatom speciation could be the old relict lakes (like LakesOhrid or Prespa in Macedonia) that have conserved their ancient conditions (Jurilj1949 1954 1956 Jerkovic 1971 Levkov et al 2003ab) Investigations of oligo-trophic lakes in Europe usually confirm the existence of new (or possibly new)diatom taxa (Lange-Bertalot amp Metzeltin 1996) and a significant diatom diversity(Hustedt 1950 Levkov et al 2001) However the number of natural ecosystems(with low anthropogenic impact) is constantly decreasing as various human impacts(Muumlller et al 1998) continue to impact environments and which consequently leadsto a decrease in the number and abundance of sensitive and rare diatom taxa (Lange-Bertalot amp Steindorf 1996) These ecosystems are therefore a subject of investigationsince diatoms are regarded as reliable indicators of acidification (Arzet et al 1986Marchetto amp Schmidt 1993 Niederhauser amp Schanz 1994) or changes in trophicstatus (Wunsam amp Schmidt 1995 Lotter et al 1998 Hall amp Smol 1999)

Besides the increasing applicability of diatoms to different types of environmentalstudies careful taxonomy becomes crucial in obtaining reliable data on diatom diversityand distribution As pointed out by Sterrenburg (2002) the basic shortcoming in thepublic concern about diversity loss is that ldquowe should first know what there is in anarea before we can determine what we are losingrdquo With particular attention to thechanges in taxonomic concepts of numerous genera and species and also the lack ofdetailed data on composition and distribution of diatom taxa in mountain aquaticecosystems in Macedonia a long-term survey (1995-2003) of these habitats on differentmountains in Macedonia has been conducted Hitherto published data on SharaMountain glacial lakes (Levkov et al 1998 2001) are scarce while there are no data

503

for Nidze Mountain aquatic habitats Consequently the results presented here amountto data revision for Shara Mountain and presentation of an initial database for diatomcomposition and distribution on Nidze Mountain The main objective of this study wasto determine the diatom species composition and the presence of possibly new taxa andto establish and promote the diatom collection at Institute of Biology Faculty of NaturalSciences in Skopje Here we give an overall list of determined and observed taxa onboth mountains as part of the Diatom Check List of Macedonia which will laterinclude biogeography and the intensity of threat to diatom microflora and habitats

Finally regarding the implementation of different taxonomy systems [reflected inStoermerrsquos (2001) intriguing questions who should I follow and what shall Ido] we have decided to merge the two existing systems (Lange-Bertalotrsquos schoolof taxonomy on one hand and Round et alrsquos revisions on the other) Thus likeAboal et al (2003) we intended to so position ourselves as not to lose the sense ofa modern approach to diatom taxonomy and hopefully to add new insight into thecomplex area of diatom research

Material and methods

The mountain Shar Planina is a long horst situated in the northwestern part of the Republic ofMacedonia and in the southwestern part of Serbia and Crna Gora (Kosovo) About 55 of its areais in the Republic of Macedonia (Fig 1) and it is elevated between the Metohija valley to thenorthwest and Polog to the southeast Its crest is more than 90 km long of which more than 80 kmare above 2000 m elevation The highest peak is Turchin (Titov Vrv) - 2748 m asl The mainhypsometrical characteristic of Shar Planina mountain is its large territory above 2000 meters elevation- 3105 km2 or 199 of the total area According to Kolchakovski (2002) geotectonically it belongsto the West-Macedonian zone It is characterized by complex geology and geomorphology Siliciousbedrocks dominate while limestone appears sporadically generally on high peaks

Glacial lakes are a characteristic feature of Shar Planina massif According to Krivokapic (1968)there are 39 lakes of which 25 are permanent On the territory of the Republic of Macedonia 19permanent and eight intermittent glacial lakes were identified The largest lakes on Macedonianterritory are Bogovisnko Ezero (66880 m2) on 1960 m asl and Crno Ezero (33520 m2) on 2170m asl The deepest point of Bogovinsko Ezero is 22 m The depth of Crno Ezero is not known butit is much deeper than Bogovinsko Ezero

Kajmakchalan lies in the southern part of the Republic of Macedonia and in the northern part of Greeceand is a part of the Nidze massif (Fig 1) which is oriented southwest to northeast and in some parts westto east direction Its crest is about 27 km long from Pelagonia valley on the west to the KozjakMountain to the east It is one of five mountains in Macedonia with an elevation above 2500 meters buthas only 83 km2 of its territory above 2000 m asl The highest peak is Kajmakchalan - 2520 m asl

Geotectonically it belongs to the neo-tectonic Selechka-Kajmachalanian block It is characterized byless complex geology (dominated by siliceous bedrocks) and geomorphology than other mountainsin Macedonia

Sampling was made at 352 localities on Shara Mountain and 124 localities on Nidze Mountain Thematerial was collected from most of the water bodies in the investigated area as glacial lakes peat-bogs springs streams rivers and wetlands at elevations between 810 and 2450 m asl Plankton(only from the largest glacial lakes on Shara Mountain) epilithon epiphyton epipelon and epipsammonwere collected by planktonic net by bottom sampler (at greater depths than 05 m) or manually andfixed in 4 formalin Sampling was conducted during summer months June to September from1995 to 2003 Microscope slides were prepared by acid digestion method (Krammer amp Lange-Bertalot 1986) and mounted in Canada-balsam (1995-1999) or Naphrax (2000-2003) mounting

504

media The slides were deposited in the Diatom Collection at the Institute of Biology Faculty ofNatural Sciences in Skopje R Macedonia Species identification and photomicroscopy were madeon a Nikon E-800 microscope at magnification x 1000 following Hustedt (1930) Schmidt et al(1874-1959) Krammer amp Lange-Bertalot (1986-1991) Lange-Bertalot amp Metzeltin (1996) Krammer(1997ab 2000 2002 2003) and Lange-Bertalot (2001) We also followed the work of AchnanthesBory sensu lato (Round amp Bukhtiyarova 1996) Hippodonta Lange-Bertalot Metzeltin amp Witkowski(Lange-Bertalot et al 1996 Lange-Bertalot 2001) Placoneis Mereschkowsky (Cox 2003) GyrosigmaHassall (Sterrenburg 1994 1995) For taxa of the family Fragilariaceae the nomenclature of Roundet al (1990) has been followed Figures were prepared according to Bayer et al (2001) Theabundance of the most important species for this study was estimated by counting of 400 valves perslide Calculation of similarity index was made using Jaccardrsquos index formula given in Washington(1984) Statistical analyses (discriminant analyses - DA) were performed using STATISTICA forWindows (StatSoft 1995) A total of 75 valves were analysed or 25 valves per group Numericalfeatures used for DA were length breadth lengthbreadth ratio and striae density in 10 microm

Results

Taxonomic observations

A total of 418 taxa belonging to 76 genera were determined in the material 324 taxafrom Shara Mountain and 217 from Nidze Mountain Of these identified taxa 123were common to both regions (Table 1) Genera with the highest number of specieswere Pinnularia Ehrenberg (68) Navicula Bory sensu stricto (33) Gomphonema

Fig 1 Study area

505

Table 1 List of identified diatoms on Shara (I) and Nidze (II) Mountains

List of taxa I II

Achnanthes boyei Oslashstrup +Achnanthes conspicua Mayer +Achnanthes exigua Grunow + +Achnanthes montana Krasske +Achnanthes oblongella Oslashstrup +Achnanthes rupestris Krasske +Achnanthidium clevei (Grunow) Czarnecki +Achnanthidium coarctatum Breacutebisson + +Achnanthidium minutissimum (Kuumltzing) Czarnecki + +A minutissimum var gracillima (Meister) Bukhtiyarova + +Actinocyclus normanii (Gregory ex Greville) Hustedt + +Adlafia bryophila (JBPetersen) Moser Lange-Bertalot amp Metzeltin +Adlafia minuscula (Grunow) Lange-Bertalot +Amphipleura pellucida (Kuumltzing) Kuumltzing + +Amphora aequalis Krammer +Amphora libyca Ehrenberg + +Amphora montana Krasske +Amphora ovalis (Kuumltzing) Kuumltzing + +Amphora pediculus (Kuumltzing) Grunow + +Anomoeoneis sphaerophora (Kuumltzing) Pfitzer +Asterionella formosa Hassall +Aulacoseira alpigena (Grunow) Krammer + +Aulacoseira granulata (Ehreberg) Simonsen +Brachysira brebissonii Ross in Hartley +Brachysira serians (Breacutebisson) Round amp DGMann +Brachysira zellensis (Grunow) Round amp DGMann +Caloneis alpestris (Grunow) Cleve +Caloneis bacillum (Grunow) Cleve + +Caloneis leptosoma (Grunow) Krammer +Caloneis schumanniana (Grunow) Cleve +Caloneis silicula (Ehrenberg) Cleve + +Caloneis tenuis (Gregory) Krammer + +Campylodiscus hibernicus Ehrenberg +Campylodiscus noricus Ehrenberg +Cavinula cocconeiformis (Gregory) DGMann amp Stickle +Cavinula pseudoscutiformis (Hustedt) DGMann amp Stickle + +Cavinula scutelloides (WSmith) Lange-Bertalot +Cavinula variostriata (Krasske) DGMann amp Stickle +Cocconeis disculus (Schumann) Cleve + +Cocconeis pediculus Ehrenberg + +Cocconeis placentula Ehrenberg + +Cocconeis placentula var euglypta (Ehrenberg) Cleve + +C placentula var intermedia (Heacuteribaud amp Peragallo) Cleve +Cocconeis placentula var lineata (Ehrenberg) Van Heurck + +Cocconeis placentula var pseudolineata Geitler +Cocconeis pseudothumensis Reichardt +Craticula ambigua (Ehrenberg) DGMann +Craticula cuspidata (Kuumltzing) DGMann +Cyclotella iris Brun amp Heacuteribaud +Cyclotella meneghiniana Kuumltzing +Cymatopleura elliptica (Breacutebisson) WSmith +Cymatopleura solea (Breacutebisson) WSmith +Cymatopleura solea var apiculata (WSmith) Ralfs +

506

Cymbella affiniformis Krammer +Cymbella alpestris Krammer +Cymbella aspera (Ehrenberg) Peragallo + +Cymbella compacta Oslashstrup + +Cymbella cymbiformis CAgardh +Cymbella dorsenotata Oslashstrup +Cymbella excisa Kuumltzing +Cymbella excisiformis Krammer +Cymbella laevis Naegeli +Cymbella lanceolata (Ehrenberg) Kirchner + +Cymbella lancetulla (Krammer) Krammer +Cymbella lange-bertalotii Krammer + +Cymbella neocistula Krammer +Cymbella neoleptoceros Krammer +Cymbella parva (WSmith) Kirchner + +Cymbella peraspera Krammer +Cymbella proxima Reimer +Cymbella subcistula Krammer +Cymbella tumida (Breacutebisson) Van Heurck +Cymbella vulgata Krammer +Cymbopleura amphicephala (Naegeli) Krammer +Cymbopleura anglica (Lagerstedt) Krammer +Cymbopleura apiculata Krammer +Cymbopleura austriaca (Grunow) Krammer +Cymbopleura citrus (Carter amp Bailey-Watts) Krammer +Cymbopleura laeviformis Krammer +Cymbopleura korana (Grunow) Krammer +Cymbopleura naviculiformis (Auerswald) Krammer + +Cymbopleura peranglica Krammer +Cymbopleura rhomboidea Krammer +Cymbopleura subaequalis (Grunow) Krammer +Cymbopleura subapiculata Krammer +Cymbopleura subaustriaca Krammer +Cymbopleura sublanceolata Krammer +Cymbopleura sp1 +Decussata hexagona (Torka) Lange-Bertalot + +Delicata delicatula (Kuumltzing) Krammer +Denticula elegans Kuumltzing +Denticula tenuis Kuumltzing +Diadesmis contenta var biceps (Grunow) Hamilton +Diadesmis gallica var perpusilla (Grunow) Lange-Bertalot +Diatoma ehrenbergii Kuumltzing + +Diatoma hyemalis (Roth) Heiberg + +Diatoma mesodon (Ehrenberg) Kuumltzing + +Diatoma moniliformis Kuumltzing +Diatoma vulgaris Bory +Diatomella balfouriana Greville +Diploneis elliptica (Kuumltzing) Cleve + +Diploneis interrupta (Kuumltzing) Cleve +Diploneis oblongella (Naegeli) ACleve + +Diploneis oculata (Breacutebisson) Cleve + +Diploneis ovalis (Hilse) Cleve + +Diploneis petersenii Hustedt + +Ellerbeckia arenaria (Moore) Crawford +Encyonema alpiniforme Krammer +Encyonema caespitosum Kuumltzing +Encyonema elginensis (Krammer) DGMann +

507

Encyonema hebridicum (Gregory) Grunow in Cleve amp Moller +Encyonema neogracile Krammer + +Encyonema mesianum (Cholnoky) DGMann +Encyonema minutum (Hilse) DGMann + +Encyonema norvegicum var alpinum Krammer + +Encyonema perpusillum (A Cleve) DGMann +Encyonema reichardtii (Krammer) DGMann +Encyonema prostratum (Berkeley) Kuumltzing +Encyonema silesiacum (Bleisch) DGMann + +Encyonema sp1 +Encyonopsis cesatii (Rabenhorst) Krammer +Encyonopsis falaisensis (Grunow) Krammer + +Encyonopsis grunowii Krammer +Encyonopsis microcephala (Grunow) Krammer +Epithemia adnata (Kuumltzing) Breacutebisson + +Epithemia argus (Ehrenberg) Kuumltzing +Epithemia argus var alpestris (WSmith) Grunow +Epithemia sorex Kuumltzing +Epithemia sorex var gracilis Hustedt +Epithemia turgida (Ehrenberg) Kuumltzing +Epithemia turgida var granulata (Ehrenberg) Brun + +Eucocconeis alpestris (Brun) Lange-Bertalot +Eucocconeis flexella (Kuumltzing) Cleve +Eucocconeis laevis (Oslashstrup) Lange-Bertalot +Eunotia arcus Ehrenberg +Eunotia bilunaris (Ehrenberg) Mills + +Eunotia diodon Ehrenberg + +Eunotia exigua (Breacutebisson) Rabenhorst +Eunotia flexuosa (Breacutebisson) Kuumltzing + +Eunotia glacialis Meister + +Eunotia incisa Gregory + +Eunotia inflata (Grunow) Noumlrpel-Schempp amp Lange-Bertalot +Eunotia minor (Kuumltzing) Grunow + +Eunotia monodon Ehrenberg +Eunotia monodon var bidens (Gregory) WSmith +Eunotia pectinalis (Dillwyn) Rabenhorst + +Eunotia pectinalis var undulata (Ralfs) Rabenhorst + +Eunotia praerupta Ehrenberg + +Eunotia soleriolii (Kuumltzing) Rabenhorst +Eunotia sudetica O Muumlller +Eunotia tetraodon Ehrenberg in Van Heurck + +Fallacia insociabilis (Krasske) DGMann +Fallacia naumannii (Hustedt) DGMann +Fallacia pygmaea (Kuumltzing) Stickle amp DGMann +Fragilaria capucina Desmaziegraveres + +Fragilaria capucina var amphicephala (Grunow) Lange-Bertalot +Fragilaria capucina var capitellata (Kuumltzing) Lange Bertalot +Fragilaria capucina var rumpens (Kuumltzing) Lange-Bertalot +Fragilaria capucina var vaucheriae (Kuumltzing) Lange-Bertalot +Fragilaria crotonensis Kitton +Fragilaria heidenii Oslashstrup +Fragilariforma bicapitata (Mayer) DMWilliams amp Round +Fragilariforma virescens (Ralfs) DMWilliams amp Round + +Frustulia amphipleuroides (Grunow) ACleve + +Frustulia crassinervia (Breacutebisson) Lange-Bertalot amp Krammer +Frustulia krammeri Lange-Bertalot amp Metzeltin +Frustulia saxonica (Rabenhorst) De Toni + +

508

Frustulia vulgaris (Thwaites) De Toni + +Geissleria decussis (Oslashstrup) Lange-Bertalot amp Metzeltin + +Geissleria schoenfeldii (Hustedt) Lange-Bertalot amp Metzeltin +Geissleria similis (Krasske) Lange-Bertalot amp Metzeltin +Gomphoneis transsilvanica (Pantocsek) Krammer +Gomphonema acuminatum Ehrenberg +Gomphonema acutiusculum (OMuumlller) ACleve +Gomphonema angustum CAgardh + +Gomphonema clavatum Ehrenberg + +Gomphonema dichotomum Kuumltzing +Gomphonema gracile Ehrenberg + +Gomphonema hebridense Gregory +Gomphonema micropus Kuumltzing +Gomphonema minutum (CAgardh) CAgardh +Gomphonema olivaceum (Hornemann) Breacutebisson + +G olivaceum var olivaceolacuum Lange-Bertalot amp Reichardt +Gomphonema parvulum Kuumltzing +Gomphonema parvulum var exilissimum Grunow +Gomphonema pumilum (Grunow) Reichardt amp Lange-Bertalot +Gomphonema sarcophagus Gregory +Gomphonema truncatum Ehrenberg + +Gomphonema vibrio Ehrenberg + +Gomphonema olivaceum var1 +Gomphonema olivaceoides Hustedt var1 +Gomphonema olivaceoides Hustedt var2 +Gomphonema sp +Gyrosigma acuminatum (Kuumltzing) Rabenhorst + +Gyrosigma attenuatum (Kuumltzing) Rabenhorst +Gyrosigma obtusatum (Sullivant amp Wormley) Boyer +Gyrosigma sciotoense (Sullivant amp Wormley) Cleve +Hannaea arcus (Ehrenberg) Patrick + +Hantzschia abundans Lange-Bertalot +Hantzschia amphioxys (Ehrenberg) Grunow + +Hantzschia elongata (Hantzsch) Grunow + +Hygropetra balfouriana (Grunow) Krammer amp Lange-Bertalot +Hygropetra capitata (Ehrenberg) Lange-Bertalot Metzeltin amp Witkowski +Karayevia laterostrata (Hustedt) Round amp Bukhtiyarova +Lemnicola hungarica (Grunow) Round amp Basson +Luticola cohnii (Hilse) DGMann +Luticola mutica (Kuumltzing) DGMann + +Luticola nivalis (Ehrenberg) DGMann +Luticola ventricosa (Kuumltzing) DGMann +Martyana martyi (Heribaud) Round +Mastogloia grevillei W Smith +Mastogloia elliptica CAgardh +Melosira lineate (Dillwyn) CAgardh +Melosira varians CAgardh +Meridion circulare (Greville) CAgardh + +Meridion circulare var constrictum (Ralfs) Van Heurck + +Muelleria gibbula (Cleve) Spaulding amp Stoermer +Navicula angusta Grunow + +Navicula antonii Lange-Bertalot +Navicula aquaedurae Lange-Bertalot +Navicula cantonatii Lange-BertalotNavicula cataracta-rheni Lange-Bertalot +Navicula catalonogermanica Lange-Bertalot amp Hofmann +Navicula concentrica Carter +

509

Navicula cryptocephala Kuumltzing + +Navicula cryptotenella Lange-Bertalot + +Navicula densilineolata (Lange-Bertalot) Lange-Bertalot +Navicula detenta Hustedt +Navicula expecta Van Landingham +Navicula gregaria Donkin +Navicula hambergii Hustedt +Navicula jakovljevicii Hustedt +Navicula lanceolata (CAgardh) Ehrenberg + +Navicula lapidosa Krasske +Navicula oligotraphenta Lange-Bertalot amp Hofmann +Navicula protracta (Grunow) Cleve +Navicula pseudolanceolata Lange-Bertalot + +Navicula radiosa Kuumltzing + +Navicula reinhardtii (Grunow) Grunow +Navicula rhynchocephala Kuumltzing +Navicula rostellata Kuumltzing +Navicula stankovicii Hustedt +Navicula subtilissima Cleve +Navicula tridentula Krasske +Navicula tripunctata (OFMuumlller) Bory + +Navicula trivialis Lange-Bertalot +Navicula viridula (Kuumltzing) Ehrenberg +Navicula viridulacalcis Lange-Bertalot +Navicula vulpina Kuumltzing +Naviculadicta brockmannii (Hustedt) Lange-Bertalot +Naviculadicta pseudosilicula (Hustedt) Lange-Bertalot +Neidium affine (Ehrenberg) Pfitzer +Neidium affine var longiceps (Gregory) Cleve +Neidium alpinum Hustedt +Neidium ampliatum (Ehrenberg) Krammer + +Neidium bergii (ACleve) Krammer +Neidium binodeforme Krammer +Neidium binodis (Ehrenberg) Hustedt + +Neidium bisulcatum (Lagerstedt) Cleve + +Neidium dubium (Ehrenberg) Cleve +Neidium iridis (Ehrenberg) Cleve +Neidium productum (WSmith) Cleve + +Neidium septentrionale ACleve +Nitzschia alpina Hustedt + +Nitzschia angustata Grunow +Nitzschia angustatula Lange-Bertalot +Nitzschia capitellata Hustedt +Nitzschia dissipata (Kuumltzing) Grunow + +Nitzschia dissipata var media (Hantzsch) Grunow + +Nitzschia fonticola Grunow + +Nitzschia linearis (CAgardh) WSmith + +Nitzschia palea (Kuumltzing)W Smith +Nitzschia recta Hantzsch + +Nitzschia sigmoidea (Nitzsch) WSmith +Nitzschia sinuata (Thwaites) Grunow in Cleve amp Grunow + +Nitzschia sinuata var delognei (Grunow) Lange-Bertalot +Nitzschia sinuata var tabellaria (Grunow) Grunow +Nupela lapidosa (Krasske) Lange-Bertalot +Orthoseira roseana (Rabenhorst) Orsquo Meara +Pinnularia acrosphaeria Rabenhorst + +Pinnularia acrosphaeria var parva Krammer +

510

Pinnularia anglica Krammer +Pinnularia angulosa Krammer +Pinnularia biceps Gregory +Pinnularia borealis Ehrenberg + +Pinnularia borealis var islandica Krammer +Pinnularia borealis var scalaris (Ehrenberg) Rabenhorst +Pinnularia borealis var subislandica Krammer +Pinnularia borealis var sublinearis Krammer + +Pinnularia brandeliformis Krammer +Pinnularia brandelii Cleve +Pinnularia brebissonii (Kuumltzing) Rabenhorst +Pinnularia crucifera ACleve +Pinnularia cuneola Reichardt +Pinnularia decrescens Grunow +Pinnularia divergentisima Grunow +Pinnularia diversa Oslashstrup +Pinnularia dubitabilis var minor Krammer +Pinnularia esoxiformis Fusey +Pinnularia frequentis Krammer +Pinnularia gibba Ehrenberg + +Pinnularia gigas Ehrenberg +Pinnularia grunowii Krammer +Pinnularia inconstans Mayer + +Pinnularia infirma Krammer +Pinnularia intermedia (Lagerstedt) Cleve + +Pinnularia julma Krammer amp Metzeltin +Pinnularia krammeri Metzeltin +Pinnularia lange-bertalotii Krammer +Pinnularia lata (Breacutebisson) Rabenhorst +Pinnularia macilenta Ehrenberg +Pinnularia microstauron (Ehrenberg) Cleve + +Pinnularia microstauron var nonfasciata Krammer +Pinnularia microstauron var rostrata Krammer +Pinnularia neomaior Krammer +Pinnularia neomaior var inflata Krammer +Pinnularia nobilis var regularis Ehrenberg +Pinnularia nodosa (Ehrenberg) WSmith +Pinnularia obscura Krasske + +Pinnularia obscuriformis Krammer +Pinnularia peracuminata Krammer +Pinnularia persudetica Krammer +Pinnularia pisciculus Ehrenberg +Pinnularia rabenhorstii (Grunow) Krammer + +Pinnularia reichardtii Krammer +Pinnularia rhombarea Krammer + +Pinnularia rhombarea var variarea Krammer +Pinnularia rupestris Hantzsch + +Pinnularia septentrionalis Krammer +Pinnularia sinistra Krammer + +Pinnularia stomatophora (Grunow) Cleve + +Pinnularia streptoraphe Cleve +Pinnularia subcapitata var subrostrata Krammer +Pinnularia subcommutata Krammer +Pinnularia subcommutata var nonfasciata Krammer +Pinnularia subgibba Krammer +Pinnularia subrupestris Krammer + +Pinnularia sudetica (Hilse) Hilse +

511

Pinnularia tirolensis Krammer +Pinnularia undula Krammer +Pinnularia viridiformis Krammer + +Pinnularia viridis (Nitzsch) Ehrenberg + +Pinnularia sp1 +Pinnularia sp2 +Pinnularia sp3 +Pinnularia rabenhorstii var1 +Placoneis clementoides (Hustedt) EJCox +Placoneis constans var symmetrica (Hustedt) EJCox +Placoneis elginensis (Gregory) EJCox +Placoneis ignorata (Schimanski) Lange-Bertalot +Placoneis paraelginensis Lange-Bertalot +Placoneis placentula (Ehrenberg) Heinzerling +P placentula var rostrata (Mayer) Andresen Stoermer amp Kreis +Placoneis porifera (Hustedt) EJCox +Planothidium dubium (Grunow) Round amp Bukhtiyarova +Planothidium ellipticum (ACleve) Round amp Bukhtiyarova +Planothidium frequentissimum (Lange-Bertalot) Round amp Bukhtiyarova + +Planothidium lanceolatum (Breacutebisson) Round amp Bukhtiyarova + +Planothidinium oestrupii (ACleve) Round amp Bukhtiyarova +Planothidium peragallii (Brun amp Heacuteribaud) Round amp Bukhtiyarova +Planothidium rostratum (Oslashstrup) Round amp Bukhtiyarova +Psammothidium bioretii (Germain) Bukhtiyarova amp Round + +Psammothidium helveticum (Hustedt) Bukhtiyarova amp Round + +Pseudostaurosira brevistriata (Grunow) DMWilliams amp Round +Puncticulata comta (Ehrenberg) Haringkansson +Puncticulata pratermissa (Lund) Haringkansson +Reimeria sinuata (Gregory) Kociolek amp Stoermer +Rhoicosphenia abbreviata (CAgardh) Lange-Bertalot + +Rhopalodia gibba (Ehrenberg) OMuumlller + +Rhopalodia gibba var parallela (Grunow) HampMPeragallo +Rhopalodia gibberula (Ehrenberg) OMuumlller + +Sellaphora bacillum (Ehrenberg) DGMann + +Sellaphora laevissima Kuumltzing +Sellaphora pupula (Kuumltzing) Mereschkowsky + +Sellaphora rectangularis (Gregory) Lange-Bertalot +Stauroneis acuta WSmith +Stauroneis anceps Ehrenberg + +Stauroneis gracilior (Rabenhorst) Reichardt +Stauroneis gracilis Ehrenberg +Stauroneis obtusa Lagerstedt +Stauroneis phoenicenteron (Nitzsch) Ehrenberg + +Stauroneis producta Grunow +Stauroneis schimanskii Krammer +Stauroneis smithii Grunow + +Stauroneis sp1 +Staurosira construens (Ehrenberg) DMWilliams amp Round +S construens var binodis (Ehrenberg) DMWilliams amp Round +Staurosira construens var venter (Ehrenberg) Hamilton +Staurosirella leptostauron (Ehrenberg) DMWilliams amp Round + +Staurosirella leptostauron var dubia (Grunow) Bukhtiyarova +Staurosirella pinnata (Ehrenberg) DMWilliams amp Round + +Stenopterobia delicatissima (Lewis) Breacutebisson +Stephanodiscus hantzschii Grunow +Stephanodiscus rotula (Kuumltzing) Hendey +Surirella angusta Kuumltzing + +

512

Surirella bifrons Ehrenberg +Surirella biseriata Breacutebisson +Surirella brebissonii var kuetzingii Krammer amp Lange-Bertalot +Surirella gracilis Grunow +Surirella linearis WSmith +Surirella linearis var helvetica (Brun) Meister +Surirella minuta Breacutebisson in Kuumltzing +Surirella robusta Ehrenberg +Surirella spiralis Kuumltzing +Surirella splendida (Ehrenberg) Kuumltzing + +Surirella turgida WSmith +Synedra acus Kuumltzing +Synedra capitata Ehrenberg +Synedra parasitica (WSmith) Hustedt +Synedra parasitica var subconstricta (Grunow) Hustedt +Synedra tabulata (CAgardh) Kuumltzing +Synedra ulna (Nitzsch) Ehrenberg + +Synedra ulna var biceps (Kuumltzing) Kirchner +Tabellaria fenestrata (Lyngbye) Kuumltzing +Tabellaria flocculosa (Roth) Kuumltzing + +Tabellaria ventricosa Kuumltzing + +

Ehrenberg (21) Cymbella CAgardh sensu stricto (20) and Eunotia Ehrenberg (17)These taxa corresponded to those thought to contribute to high biodiversity inoligotrophic lakes (Lange-Bertalot amp Metzeltin 1996) According to Levkov et al(1998) Navicula sensu lato and Cymbella sensu lato dominate in aquatic habitats onShara Mountain We also found 11 taxa whose taxonomic position could not beprecisely established according to available literature so their relationship to closelyrelated taxa is also discussed below

Pinnularia sp1 Figs 4-7

Description Valve outline linear with parallel to slightly convex or concave marginsValve ends are broadly truncated to obtusely rounded length 67-110 microm breadth205-240 microm lengthbreadth ratio 32-46 Raphe moderately lateral Outer fissures(proximal raphe ends) are curved central pores large and rounded terminal fissure(distal raphe ends) are sickle shaped Axial area linear 15 to 14 of the valve breadthornamented with irregular structures Central area rounded to asymmetrical 12 ofthe valve breadth Striae broad moderately widely separated 3-410 microm radiate inthe middle parallel to weakly convergent towards the ends

Diagnostic characters outline breadth and presence of irregular structure in theaxial area

Taxonomic remarks This taxon is closely related to Pinnularia splendida Hustedt Prabenhorstii (Grunow) Krammer and P lata (Breacutebisson) Rabenhorst Pinnularia sp1has a more linear outline and smaller axial area than P splendida Both taxa haveirregular structures present in axial area This taxon is similar in valve outline to Prabenhorstii but the latter has narrower valves (16-18 microm) The valve outline differsfrom P lata in being more linear but also by the presence of the irregular structures inthe axial area

513

Slide No 46A ZLC Diatom Collection at Institute of Biology Faculty of NaturalSciences Skopje Republic of Macedonia

Distribution The taxon is known only from Lake Crno Ezero and occurs as rarevalves in the sediment

Fig 2 Scatter-plot of analysed specimens of P rabenhorstii in the space of the first and the seconddiscriminant axes

Fig 3 Scatter-plot of centroids of analysed groups of P rabenhorstii in the space of the first and thesecond discriminant axes

514

Figs 4-7 Pinnularia sp1 Figs 8-13 Pinnularia rabenhorstii var1 Scale bar =10 microm

515

Pinnularia rabenhorstii (Grunow) Krammer var1 Figs 8-13

Description Valve outline linear with parallel to moderately convex margins Valveends are bluntly rounded not protraced Length 38-71 microm breadth 12-17 microm lengthbreadth ratio 32 (in smaller specimens) to 51 (in larger specimens) Raphe moderatelylateral Proximal raphe ends with distinct rounded central pores Terminal rapheends large sickle shaped Axial area narrow central area large elliptical to transapicallywidened Striae broad 4-710 microm radiate in the middle to weakly convergent towardsthe ends

Diagnostic characters outline valve ends length breadth

Taxonomic remarks According to Krammer (2000) this complex consists of fourvarieties two of which are mentioned for oligotrophic waters in Europe Prabenhorstii (Grunow) Krammer var rabenhorstii and P rabenhorstii var franconicaKrammer The varieties are often found in mixed populations and the main differentialcharacter for these two taxa is breadth 16-18 microm and 120-135 microm respectivelyKrammer (2000) gives a clear distinction between the two European varieties of Prabenhorstii without recognizing forms with intermediate morphometriccharacteristics

The populations of P rabenhorstii were observed in three separate localities on twodifferent mountains in the western region of Macedonia (Shara Mountain and BistraMountain) Our observations show that there is a marked infraspecific variationregarding morphology and cell size Populations observed on Shara Mountain formtwo natural groups that correspond to the sampling localities and clearly differ in sizeand morphology The population living in Crno Ezero (Shara Mountain) has largerspecimens that most closely resemble the nominate variety (length 57-71 microm breadth155-170 microm) Considering the valve outline and breadth the population found onBistra mountain belongs to P rabenhorstii var franconica (length 53-71 microm breadth12-14 microm) A population from lakes near the spring area of River Pena (Shara Mountain)shows a difference in morphology regarding the outline breadth length and striadensity The valves are linear to linear-elliptic with rounded ends (whereas Prabenhorstii var rabenhorstii has moderately subrostrate ends) The length is 38-71microm (40-60 microm being the most abundant size range in the population) breadth 12-17microm (usually 13-15 microm) lengthbreadth ratio 32-51 (usually 35-45)

Smaller specimens usually have an average lengthbreadth ratio of circa 35 whilelarger ones circa 45 Stria density is also variable and is 4-7 depending on valvedimensions

The highest multivariate variability (Fig 2) was observed in the sample of group 3(populations from lakes near river Pena springs Shara Mountain) since somespecimens of this group overlapped with the specimens of group 1 (populationsfrom Lake Crno Ezero Shara Mountain) Discriminant analysis of centroids (multi-variate mean values) of the groups analysed (Fig 3) showed a clear discriminationof difference between the samples The first discriminant axis clearly separated allthree groups analysed The second axis additionally separated group 3 (populationsfrom lakes near river Pena springs Shara Mountain)

516

Nevertheless both analyses (of specimens and centroids) showed that at the multivariatelevel there were clear and statistically significant differences among the samplesanalysed All three taxa show differences at the multivariate level and the differentialdiagnosis for these taxa should be based on a combination of quantitative andqualitative characters eg length breadth lengthbreadth ratio and outline

Slide No 54A95 Shara ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Distribution Known only from lakes near river Penarsquos springs where it is abundantin the sediment


Description Valve outline linear with parallel to slightly convex margins narrowingtoward the acutely rounded ends Length 85-175 microm breadth 22-28 microm lengthbreadth ratio 38-63 Raphe semi-complex outer raphe fissure laterally bent Centralpores small rounded terminal raphe fissures as in P viridis (Nitzsch) EhrenbergAxial area linear 15-14 of the valve breadth tapering at the ends Central arealarge rounded to asymmetric frac12 of the valve breadth Striae 55-6010 microm radiatein the middle convergent at the ends crossed by wide longitudinal bands

Diagnostic characters outline size shape of central area and striae density

Taxonomic remarks This taxon is similar to Pinnularia stidolphii Krammer (Krammer2000 fig 1833) According to Krammer (2000) there is no other taxon with similarcharacters but he also mentioned that there is a small difference in the shape of thecentral area between European populations and populations from New Zealand Thepopulation found in Crno Ezero Shara Mountain has a greater valve breadth andfewer striae in 10 microm than P stidolphii This taxon differs from P viridis by thelengthbreadth ratio shape of the central area and striae density and from Pviridiformis Krammer by the valve breadth and shape of the central area

Slide No 47A Shara ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Distribution known only from Lake Crno Ezero common in the sediment

Encyonema sp1 Figs 19-23

Description Valves moderately dorsi-ventral dorsal margin arched ventral marginslightly swollen in the middle Ends not set off from the valve body bluntly roundedLength 60-83 microm breadth 14-18 microm lengthbreadth ratio 39-55 Axial areamoderately wide ventrally displaced opening in small central area Raphe filiformproximal raphe ends dorsally deflected distal raphe ends prolonged dorsally deflectednot reaching the dorsal valve edge Dorsal striae parallel in the middle 7-810 microm toslightly radiate at the ends 1010 microm Ventral striae radiate in the middle 810 microm toconvergent at the ends 11-1310 microm 18-22 puncta in 10 microm

Diagnostic characters outline length breadth shape of axial area

517

Figs 14-18 Pinnularia sp2 Scale bar = 10 microm

518

Taxonomic remarks According to length and breadth this species is close to Eperelginense Krammer but it differs by the outline Encyonema perelginense hasstrongly arched dorsal margin and acutely rounded ends It is distinguished from Evulgare Krammer and closely related taxa by being more elongate in valve outlineand valve length and breadth

Slide No 26A Nidze ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Distribution Known only from a small wetland near the Prava Reka River NidzeMountain with 12 abundance

Cymbopleura sp1 Figs 24-31

Description Valves moderately dorsiventral broadly lanceolate dorsal marginstrongly arched ventral margin slightly arched to linear in the middle Ends slightlyprotracted and acutely rounded Length 75-145 microm breadth 28-36 microm lengthbreadthratio 26-39 Axial area narrow to moderately broad sublinear broadening towardsthe valve center Central area elliptical to slightly asymmetrical to the ventral sideRaphe lateral narrowing towards the proximal and distal ends Proximal raphe endsexpanded in large central pores Striae radiate distinctly punctated 5-610 microm to10-11 at the ends puncta 18-2210 microm

Diagnostic characters outline length breadth shape of axial and central area

Differential diagnosis The morphology of this taxon is comparable to Cymbopleurainaequalis (Ehrenberg) Krammer C inaequaliformis Krammer and C albanicaKrammer amp Miho (Table 2) The taxon differs from mentioned taxa by thecombination of quantitative and qualitative characters from C inaequalis by smallerbreadth length and shape of the central area from C inaequaliformis by dimensionsvalve ends and shape of the central area while from C albanica it is distingushed bybreadth valve ends striae density

Slide No 5495 Shara ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Distribution Known from epipelic assemblages of Lakes Bogovinsko Ezero whereit is present with 8 abundance and Belo Ezero and lakes near river Penarsquos springswhere it is rare (less than 01) in the epipelon

Gomphoneis transsilvanica (Pantocsek) Krammer sensu lato Figs 32-34

Taxonomic remarks There is a prolonged controversy regarding the genusGomphoneis Cleve in relation to its separation from Gomphonema Ehrenbergresulting in numerous transfers of taxa between these two genera Dawson (1973)on the basis of striae with double rows of areloae states that the species complex Golivaceum should be transferred to the genus Gomphoneis Krammer (1982) proposed

Figs 19-23 Encyonema sp1 Figs 24-26 Cymbopleura sp1 Scale bar =10 microm

519

520

that G transsilvanica should be a member of Gomphoneis because of its non-differentiated apical area and 3-5 rows of areolae in one stria Cladistic analysis ofthe members of the Gomphoneis group (Kociolek amp Stoermer 1989 1993) indicatesthat they are mostly related to Gomphopleura Reichelt ex Tempegravere and the Gomphoneiselegans (Grunow) Cleve group because of the presence of marginal lamina Analysisof the recent populations from Lake Ohrid Lake Prespa (Levkov et al 2003a) andNidze Mountain reveal that the specimens have an overall morphology that is quitedifferent and needs additional investigations for proper taxonomic revision TheGomphoneis taxa found in Nidze Mountain as well as taxa found in Lakes Prespaand Ohrid possess two rows of areolae in one stria but also different shape of centralarea to G transsilvanica

Distribution in the study it was found in a tributary of the Trnovcica river NidzeMountain 820 m asl in the sediment

Slide No 4202 Nidze ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Gomphonema olivaceum (Hornemann) Breacutebisson var1 Figs 35-37

Description Valve clavate with broadly rounded head pole and narrowly roundedfoot pole Widest part near the middle portion of the valve Length 40-70 microm breadth12-14 microm Raphe filiform proximal and terminal fissures small and distinct Axialarea linear very narrow Central area elliptical occupying half of the valve breadthStriae radiate throughout the valve indistinctly punctated 12-1310 microm

Taxonomic remarks Investigations of the G olivaceum group in the area studied ledto separation of four different taxa In epiphytic communities on Nidze Mountain a

Table 2 Morphometric characteristics of Cymbopleura ianequalis complex

Taxa Outline Ends Central area LengthBreadth Striaepuncta(microm) (in 10 microm)

Cymbopleura broadly slightly truncate slightly 90-190 32-441 5-714-181

inaequalis rhomboid and ellipticallanceolate bluntly rounded

Cymbopleura rhomboid not protracted slightly 57-12023-28 6-924-30inaequaliformis lanceolate ovoid

Cymbopleura lanceolate slightly apiculate rounded or 57-6318-191 9-1028-301

albanica elliptical 55-105 24-282 70-8518-242

Cymbopleura broadly slightly truncate to elliptical or 75-14528-36 5-618-22sp1 lanceolate acutely rounded asymmetric

1 Data taken from Krammer (2003) table on page 182 Data taken from Krammer (2003) description on pages 24-25

Figs 27-31 Cymbopleura sp1 light microscopy Scale bar =10 microm

521

522

Figs 32-34 Gomphoneis transsilvanica (Pantocsek) Krammer sensu lato Figs 35-37 Gomphonemaolivaceum (Hornemann) Breacutebisson var1 Figs 38-41 Gomphonema olivaceoides Hustedt var1Figs 42-48 Gomphonema olivaceoides Hustedt var2 Scale bar =10 microm

523

taxon close to G olivaceum var olivaceolacuum Lange-Bertalot amp Reichardt wasobserved but this has somewhat different morphometric features from the statedvariety especially as regards the shape and size of the central area and overalldimensions The same taxon has also been observed in Lake Prespa (Levkov et al2003b) It is a transitional form between G olivaceum var olivaceolacuum andGomphoneis transsilvanica From the last it is best distinguished by smaller dimensionsand shape of the central area

Distribution in the study it was found in tributary of Trnovcica river Nidze Mountain820 m asl in the sediment

Slide No 42A02 Nidze ZLC Diatom Collection at Institute of Biology Faculty ofNatural Sciences Skopje Republic of Macedonia

Gomphonema olivaceoides Hustedt var1 Figs 38-41

Description Valve clavate with broadly rounded head pole and narrowly roundedfoot pole widest part near the middle portion of the valve Length 15-28 microm breadth7-9 microm Raphe filiform not extending in the central area Proximal and terminalfissures small and distinct Axial area linear very narrow Central area transverse ofthe valve breadth Four stigmoids are present in the central area opposite to theproximal raphe fissures Striae radiate throughout the valve 16-2010 microm indistinctlypunctated Pseudoseptum was observed




Valve clavate with broadly rounded slightly truncated head pole and narrowly roundedprotracted foot pole Widest part is the middle portion of the valve Length 25-37microm breadth 6-10 microm Raphe filiform not extending in the central area Proximaland terminal fissures small and distinct Axial area linear very narrow Central areatransverse of the valve breadth Four stigmoids are present in the central area oppositethe proximal raphe fissures Striae radiate throughout the valve 14-1610 micromindistinctly punctate Pseudoseptum was observed

Distribution river Chaushica Shara Mountain 1650 m asl on stones

Slide No 22-2496 Shara ZLC Diatom Collection at Institute of Biology Facultyof Natural Sciences Skopje Republic of Macedonia

Taxonomic remarks Hustedt (1950) describes G olivaceoides from Kirchensee as18-35 microm long 5-6 microm wide and has 10 striae10 microm Patrick amp Reimer (1975)found 10-12 striae10 microm sometimes 13-14 at the valve ends Foged (1971a)mentioned Gomphonema olivaceoides var densestriatum Foged and that it differsfrom the nominate variety by the finer striae 18-2110 microm The same author mentionedG olivaceum var spitsbergense Foged that differs in ecological preferences (pHbelow 7) and valve morphology

524

According to Foged (1971a) G olivaceoides is widely distributed (Europe NorthAmerica Central Asia) and possesses great variation in morphology and size withfew recognized varieties ie var cochleariforme Manguin [later transferred toGomphoneis as G quadripunctata var cochleariformis Manguin ex Kociolek ampStoermer by Kociolek amp Stoermer (1991)] var lanceolatum Manguin varspitsbergense Foged and var densestriatum Foged Kociolek amp Stoermer (1988)also found this taxon in Lake Baikal together with Gomphoneis quadripunctatum(Oslashstrup) Dawson ex Ross amp Sims The species was transferred to Gomphoneis dueto the possession of biserate striae (Carter amp Bailey-Watts 1981) as in case of otherspecies from G olivaceum group (Dawson 1974) but Krammer (1982) rejected thisconcept of the genus Gomphoneis Here we are following the species concept ofKrammer amp Lange-Bertalot (1986-1991)

Both identified populations in Macedonia have different valve shape and differentstria density than nominate species and recognized varieties The var1 is similar to Golivaceoides sensu Foged (fig II 10) but it differs by the greater stria density andlarger breadth This taxon has previously been recorded in Lake Ohrid (Jurilj 1954)and in Lake Prespa (Levkov et al 2003a) According to Jurilj (1954) the taxon ismore closely related to Gomphonema elegans var quadripunctatum Skvortzov amp Mayer(syn G innatum Skvortzov) than to G olivaceum (Hornemann) Breacutebisson Accordingto Zabelina et al (1951) G innatum is 52 microm long and 13 microm wide with 12 striae10microm The second population (var 2) from Shara Mountain has quite different valveoutline in comparison to the figures presented in Foged (1971a) and Hustedt (1950)

Navicula jakovljevicii Hustedt Figs 49-55

Taxonomic remarks According to Hustedt (1945) N jakovljevicii possesses featuresthat are not typical for Navicula sensu stricto since the morphology of the striaeplaces it closer to Caloneis Cleve Reichardt (1992) states that this taxon has occludedareolae on the inside part of the valve creating a longitudinal line similar to CaloneisPinnularia (pro parte) and Gomphoneis (pro parte) Specimens with similar striaepattern have been identified in fossil deposits from Transylvania (Lange-Bertalot2001) Hustedt (1945) states that the valve is 32-60 microm long 8-11 microm wide and has16-18 striae10 microm Reichardt (1992) found populations in Lake Zug (Zugersee)that are smaller (length 329-530 microm breadth 84-97) and have coarser striae (140-15510 microm) while populations from the river Krka had a length of 39-85 microm and abreadth of 88-117 microm Finally Lange-Bertalot (2001) presents populations withcells of 26-85 microm long and with 14-17 striae10 microm

Analysed morphometric features of N jakovljevicii populations from Nidze Mountainand Lake Ohrid (Levkov et al 2003a) suggest that there are at least two differentpopulations separated by two characters The first N jakovljevicii morphotype 1 issimilar to Hustedtrsquos (1945) description as regards the number of striae in 10 microm (16-

Figs 49-55 Navicula jakovljevicii Hustedt Figs 56-64 Navicula reinhardtii (Grunow) GrunowFigs 56-58 N reinhardtii morphotype 1 Figs 59-61 N reinhardtii morphotype 2 Figs 62-64 Nreinhardtii morphotype 3 Scale bar =10 microm

525

526

18) and the presence of a longitudinal band but has a greater length (50-80 microm)The second N jakovljevicii morphotype 2 has smaller forms (30-50 microm) an indistinctlongitudinal band and considerably fewer striae in 10 microm (12-14) Additionalcomparative investigations will be made on populations from Lake Ohrid and NidzeMountain

Distribution in this study it was found in a tributary of the Trnovcica river NidzeMountain 820 m asl in the sediment


Navicula reinhardtii (Grunow) Grunow Figs 56-64

Taxonomic remarksAccording to Lange-Bertalot (2001) this taxon has not changedits morphology since the Upper Tertiary period with cells of 35-70 microm long and 11-18 microm wide In aquatic ecosystems on Nidze Mountain and Lake Prespa (Levkov et al2003b) there are at least three different morphotypes differentiated by valve outlineThe first morphotype (Figs 56-58) resembles the holotype the second (Figs 59-61)has typically rhomboid valves with acutely rounded valve ends broader (17-22 microm)cells and fewer striae (6-710 microm) the third (Figs 62-64) has a markedly ellipticaloutline and acutely rounded ends In Lake Ohrid Jurilj (1954 fig 37bc) identified Nreinhardtii with rhomboid valves but does not present their morphometric values Theoccurrence of a different valve morphology of this taxon has also been recorded inLake Hovsgol Mongolia (Edlund et al in press) and Lake Baikal (Mann 1999)



Discussion

Unicellular taxa show a greater relative local species richness compared to multicellulartaxa and their distribuiton is much less dependent on geographical distances (Hillebrandet al 2001) Several researchers confirm the existence of centers with a high diversityof diatoms (Jurilj 1954 Foged 1971b Stoermer 1975) mostly within oligotrophiclakes with a high buffering capacity for acidity (Lange-Bertalot amp Metzeltin 1996)Nevertheless diatom diversity also greately depends on the number of samplingsand number of analysed valves (Hillebrand et al 2001)

Application of different taxonomic concepts in specific taxa determination has ledto an increasing number of known taxa but has also created additional confusionwith so called ldquocriticalrdquo forms (Levkov et al 2003a) Mann (1999) states that Diploneisfusca (Gregory) Cleve - D smithii (Breacutebisson) Cleve complex ldquodemesrdquo show stablemorphometric characteristics over a long period implying that they are constantgenetic features In cases where prolonged sampling has been carried out theoccurrence of multiple size classes - either overlapping or not - has been determined

527

(Potapova amp Snoeijs 1997) This has led to the hypothesis that larger and smallerldquodemesrdquo might only be distinct life stages of just one taxon (Kociolek amp Stoermer1988)

Description of new taxa based on one or a very few cells has been justly criticizedalthough there are still such attempts (Krammer 2000) Since we also consider thisapproach to be unproductive and possibly erroneous (by omitting the whole range ofmorphological and other variation of the population in question) a certain numberof observed taxa in both mountains investigated have been put aside for futureinvestigations either because of their low abundance in the material or sporadic(spatial or temporal) occurrence in the samples The morphometric analysis of thetaxa described here is based on stable features observed during the whole 8 years ofinvestigation The descriptions of and comments on other diatoms were made aftertwo or more years of observation (on tens or hundreds of valves per taxa) of theirmorphological features In addition they were abundant over this period Some taxapreviously observed in Lake Ohrid or Lake Prespa will be described after futurecomparative analysis

A Shara mountain

The highest diatom diversity was observed in glacial lakes on Shara Mountainecosystems that are considerably better preserved from human influence than lowlandlotic environments (Muumlller et al 1998) Eutrophication is reported to be the mostimportant threat for these lakes depending on water volume latitude and geologicalsubstrate (Bjork 1988) On the other hand effects of acidification as the mostprominent problem of glacial lakes in North Europe (Arzet et al 1986 Morsell etal 1991 Lotter et al 1997 1999) have not yet been detected on Shara Mountain

In previous investigations on Shara Mountain aquatic ecosystems (Levkov et al2001) some of the observed taxa were either not identified or incorrectly identifiedas in the case of the species complexes Pinnularia borealis Ehrenberg P rupestrisHantzsch P viridis (Nitzsch) Ehrenberg Cymbella affinis Kuumltzing as a result of thelsquofitting into known speciesrsquo problem

Each of the investigated lakes presents a certain specificity in diatom assemblagemainly caused by a difference in environmental factors (Lange-Bertalot amp Metzeltin1996) In Crno Ezero Navicula pseudolanceolata Lange-Bertalot Pinnulariarabenhorstii P borealis Ehrenberg sensu lato P sp2 Cymbella proxima ReimerCymbopleura apiculata Krammer or oligotrophic alkaliphilous taxa (Van Dam etal 1994 Rott 1995) were dominant The presence of rare species makes this themost important lake on Shara Mountain (Levkov et al 2001)

Diatom assemblages of Bogovinsko Ezero are very similar to those reported forMittersee Austria (Lange-Bertalot amp Metzeltin 1996) Human impact is greater as aresult of shepherding (summer pastures and sheepfolds) but the littoral region isstill marked by the presence of oligotrophic taxa such as Pseudostaurosira brevistriata(Grunow) DMWilliams amp Round Eunotia diodon Ehrenberg Neidium dubium(Ehrenberg) Cleve Pinnularia decrescens Grunow (Lange-Bertalot amp Steindorf 1996Lange-Bertalot amp Metzeltin 1996) as well as the relatively abundant Cymbopleura sp1

528

Belo Ezero and lakes in the vicinity of the Pena River source waters are typicallyhumic lakes dominated by Cymatopleura sp1 Navicula pseudolanceolata Lange-Bertalot N concentrica Carter Stauroneis gracilis Ehrenberg Pinnularia borealissl P rabenhorstii var1

Lake Dolno Karanikolicko on the other hand is a typically oligotrophic calcium-poor lake without sediment on the bottom and few vascular plants on which epiphyticFragilaria taxa dominate (Morales 2001 Douglas amp Smol 1999) The higher elevation(2180 m asl) of this lake results in prolonged ice periods and low summertemperatures which stimulate the growth of these taxa and others such as Muelleriagibbula (Cleve) Spaulding amp Stoermer Gomphonema truncatum EhrenbergStauroneis acuta WSmith and Cymbella neocistula Krammer (Schmidt et al 1998Lotter amp Bigler 2000)

Shara Mountain also has numerous oligotrophic (or dystrophic) limnogenous andsoligenous peat bogs (Dammon amp French 1987) with a very rich diatom diversityprimarily dependent on the altitude and persistence of these aquatic habitats Indystrophic bogs (at lower altitude) Hantzschia amphyoxys and Pinnularia borealissl dominate as aerophils that can endure prolonged dry periods (Van de Vijver et al2003) while in stable permanent bogs an association of Cymbopleura subaequalis(Grunow) Krammer Pinnularia viridiformis Krammer P subcapitata GregoryFrustulia spp (Novakova 2002) dominates On higher altitudes (above 2000 m)Eunotia incisa Gregory Cavinula pseudoscutiformis (Hustedt) DGMann amp Stickleand Tabellaria flocculosa (Roth) Kuumltzing dominate the peat bogs but sometimesacidophilic taxa such as Eunotia exigua (Breacutebisson) Rabenhorst E glacialis MeisterE pectinalis (Dillwyn) Rabenhorst Frustulia saxonica (Rabenhorst) De ToniPinnularia obscura Krasske are also frequently found (Dickman et al 1984 DeNicola2000) The last three taxa are mostly found in late spring in streams created bymelting snow Usually in streams and rivulets under 2000 m elevation an assemblagedominated by Achnanthes oblongella Oslashstrup Cocconeis placentula EhrenbergCymbella aspera (Ehrenberg) Peragallo Encyonema neogracile Krammer Hannaeaarcus (Ehrenberg) Patrick is frequent while above 2000 m asl Tabellaria ventricosaKuumltzing Diatoma mesodon (Ehrenberg) Kuumltzing Diatoma hyemalis (Roth) HeibergCavinula pseudoscutiformis Cymbopleura subaequalis (Grunow) KrammerStaurosirella pinnata (Ehrenberg) DMWilliams amp Round and taxa known fromwater sources on silicate substrates dominate

B Nidze Mountain

Investigations of the diatom microflora on Nidze Mountain revealed significantdifferences at various elevations In particular based on the observed diatomassemblages the whole area can be roughly divided into three zones (i) refugial(ii) intermittent and (iii) alpine

(i) refugial zone - catchment area of the river Trnovcica (800-1400 m asl) epiphyticdiatom communities consist of Gomphoneis transsilvanica Gomphonema olivaceoidesvar1 Navicula jakovljevicii Navicula reinhardtii and Ellerbeckia arenaria (Moore)Crawford This kind of community was hitherto known only for sources of the riverCrni Drim at Lake Ohrid (Hustedt 1945 Jurilj 1954 Levkov et al 2003a) The only

529

possible conclusion is that this is a Tertiary refugial microflora once (during thePliocene epoch) dominant in a much wider area and which today apart from LakesOhrid and Prespa (as well-defined refugia) exists only in a few places such asNidze aquatic habitats

According to Manakovic amp Andonoski (1984) during the Pliocene the area aroundMariovo was covered by the so-called Lake Mariovo At the end of the Pliocene andthe start of the Pleistocene when the Aegean landmass decreased in size Lake Mariovowas drained and the recent river water network formed (river Crna Reka) Theformer Lake Mariovo was never connected to Lakes Ohrid or Prespa [these belongto the Lake Dessaret group (Cvijic 1906)] but nevertheless had the same floristicelements as Lake Ohrid Freshwater lake fossil deposits (diatomite) are abundantlyfound in Manastirec-Zovik-Besiste region characterized by the association of Naviculahasta Pantocsek N reinhardtii var gracilior Grunow E arenaria (Jenko ampGuzelkovski 1958) or taxa previously recorded also for fossil deposits in Transylvania(Pantocsek 1892) According to Kociolek amp Spaulding (2002) E arenaria is mostlytypical of oligotrophic waters (as interpreted from Krammer amp Lange-Bertalot 1991)These authors also state that Ellerbeckia valves are usually present inside theActinocyclus Ehrenberg valves (as interior valves) as a result of the gradual turnoverfrom marine to freshwater habitats Even more remarkably they claim that the twoforms are actually the same species (conspecific) Crawford (2004) disputes thesefindings showing that this association is purely physical fitting of the valves of thetwo species In the area investigated here we also find valves of A normanii (Gregoryex Greville) Hustedt but valve fitting between the two species was not observedCrawford (2004) also states that E arenaria is usually collected from subareal sitesas epiphytic on mosses in alkaline areas and our findings are in complete concordancewith Crawfordrsquos (2004) comments on the distribution of the species

Numerous authorities consider freshwater diatoms to be widely distributed (Hustedt1930 Krammer amp Lange-Bertalot 1986-1991 Krammer 2000 2002 Lange-Bertalot 2001) and that endemism is a rare event mostly confined to lakes (Hustedt1945 Jurilj 1949 1954 Martens 1997) and extremely rare in running waters(Hustedt 1945) This conclusion most definitely applies to aquatic environmentsin Europe which have repeatedly changed their characteristics due to geotectonicor climatic influences mostly over several glaciation periods (Kociolek amp Spaulding2000) In lower parts of Nidze Mountain these events (geotectonic in the Pleistoceneand climatic in the Pliocene) had a minor influence on the diatom microfloracomposition In the same period areas around sources of the Trnovcica Riverwere very sparsely inhabitated by humans also resulting in low human pressureThis contrasts for example with the situation in the Vardar River valley (alsopartly known as a refugia centre) where human impact has led to profoundmicroflora successions due to severe pollution (Krstic 1995 Krstic et al 1999Levkov 2001) These conclusions for Nidze Mountain are supported by studies onthe forest ecosystems (Em et al 1985)

(ii) intermittent zone - This begins from 1400 m asl onwards and is representedby a completely different diatom microflora dominated by other (but no lessinteresting) diatom taxa Running waters between 1400 and 1600 m elevation support

530

a flora dominated by Decussata hexagona (Torka) Lange-Bertalot Hygropetrabalfouriana (Grunow) Krammer amp Lange-Bertalot Placoneis ignorata (Schimanski)Lange-Bertalot Pinnularia spp and although changed by elevation and ecologicalconditions still display a markedly ldquorelictrdquo association in which Cocconeispseudothumensis Reichardt Placoneis clementoides (Hustedt) EJCox P placentula(Ehrenberg) Heinzerling can also be found known so far only for Lakes Ohrid andPrespa (Jurilj 1954 Levkov et al 2003ab)

Decussata hexagona in Macedonia was originally found only in one peat bog onShara Mountain (Levkov et al 2001) as epiphytic on aquatic mosses According toLange-Bertalot (2001) this taxon is rare and occurs in small populations but inbrooks on Nidze Mountain (near the Redir forest house) its abundance is more than20 in the samples Populations of the rare D hexagona on Nidze Mountain areexcellently preserved in Macedonia Significant populations (5) of the rareHygropetra balfouriana described for high Alpine regions (Krammer 2000) and theArctic (Van de Vijver et al 2003) as an epiphyte on vascular plants (Beyens amp Van deVijver 2000) were observed as epiphytes on mosses in a few streams between 1500and 1600 m asl Since its occurrence would have been expected in much higherelevation (above 2000 m) the only explanation would again be supportive for acomplex refugial zone on Nidze Mountain

(iii) alpine zone - altitudes above 1600 up to 2450 meters are inhabited by diatomtaxa common to all high mountains in Macedonia as a zone basically influencedgeomorphologically and biologically by glaciation events The diatom microflorais mostly represented by taxa of Pinnularia Ehrenberg Eunotia Ehrenberg TabellariaEhrenberg Frustulia Rabenhorst Cymbella CAgardh and Stauroneis Ehrenberg(Metzeltin amp Witkowski 1996 Cantonati 1998 Cameron et al 2002 Negro et al2003)

C Floristical analyses

Comparative floristic analysis of the diatom distribution in both mountain aquaticecosystems revealed only 2274 floristic similarity although Shara and NidzeMountains are only approximately 100 km apart with 123 taxa being common toboth investigated areas This is probably the result of several factors

ndash Presence of taxa that are widespread on both mountains like Achnanthidiumminutissimum (Kuumltzing) Czarnecki Meridion circulare (Greville) CAgardhDiatoma hyemalis D mesodon Cocconeis placentula Planothidium lanceolatum(Breacutebisson) Round amp Bukhtiyarova Amphora pediculus (Kuumltzing) Grunow (Tricket al 2002 Vilbaste amp Truu 2003)

ndash Presence of taxa normally distributed in high mountain habitats (boreal taxa) onboth mountains (Krammer amp Lange-Bertalot 1986-1991) such as Aulacoseiraalpigena (Grunow) Krammer Eunotia flexuosa E glacialis E incisa Frustuliaamphipleuroides (Grunow) ACleve Nitzschia alpina Hustedt

ndash Presence of oligotrophic taxa on both mountains Cymbopleura naviculiformis(Auerswald) Krammer Diploneis spp Encyonema minutum (Hilse) DGMann

531

Encyonopsis falaisensis (Grunow) Krammer Epithemia turgida var granulata(Ehrenberg) Brun Navicula pseudolanceolata (Lange-Bertalot amp Metzeltin 1996)

ndash Taxa found only in glacial lakes on Shara Mountain like Cymbella alpestrisKrammer C dorsenotata Oslashstrup C proxima Cymbopleura anglica (Lagerstedt)Krammer C apiculata Pinnularia neomaior Krammer P septentrionalisKrammer Stauroneis acuta

ndash Taxa typical only for the refugial habitats on Nidze Mountain such as Gomphoneistranssilvanica Gomphonema olivaceoides var1 N reinhardtii N jakovljeviciiCocconeis pseudothumensis Ellerbeckia arenaria

In particular the low diatom microflora similarity recorded is mainly the result ofthe last two factors Shara Mountain possesses glacial lakes with a unique microfloracomposition while Nidze Mountain does not have any such aquatic habitatsConversely the richest microflora on Nidze Mountain is determined in the refugialzone near the Trnovcica River consisting mostly of taxa so far known only forLakes Ohrid and Prespa in Macedonia These two diatom communities are almostcompletely different containing only a very few common ldquocosmopolitanrdquo taxa

Concluding remarks

In addition to the completely unexpected major difference in diatom microfloracommunity between the two geographically very close mountains investigated oneof the most prominent findings in this study was the diatom microflora refugiacenter on Nidze Mountain consisting of diatom assemblages very similar to thefamous Lake Ohrid forms These findings clearly indicate the necessity for futurefundamental investigations regarding both the recent and the fossil diatomcomposition and distribution in Macedonia since it is obvious that even after arelatively long period of research (Petkov 1910) the diatom microflora in this partof Europe remains poorly investigated

A valuable spin-off of our investigation has been the foundation of the very firstdiatom collection herbarium in Republic of Macedonia (at the Laboratory for AlgalSystematics) which hopefully will fundamentally support taxonomic diatom researchin the country

Acknowledgements

Authors wish to express the warmest gratitude to Ditmar Metzeltin for his full cooperation inpresented taxonomic research Sarah Spaulding and Frithjof Sterrenburg for the useful commentsand English revision of the manuscript thus profoundly increasing the quality of the paper For thefield studies and sampling authors are grateful to The Research Society of Biology Students ofMacedonia We also want to express our gratitude to Ljljana Tomovic for statistical analyses and toSlavco Hristovski for technical support

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532

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ROUND EF amp L BUKHTIYAROVA (1996) Four new genera based on Achnanthes(Achnanthidium) together with a re-definition of Achnanthidium - Diatom Res 11 345-361

ROUND FE RM CRAWFORD amp DG MANN (1990) The diatoms biology and morphologyof the genera 1-747 - Cambridge University Press Cambridge

SALA SE EA SAR amp ME FERRARIO (1998) Review of materials reported as containingAmphora coffeaeformis (Agardh) Kuumltzing in Argentina - Diatom Res 13 323-336

SCHMIDT A et al (1874-1959) Atlas der Diatomaceen-Kunde Heft 1-120 Tafeln 1-480 (Tafeln1-213 ASchmidt 213-216 MSchmidt 217-240 FFricke 241-244 HHeiden 245-246 OMuumlller247-256 FFricke 257-264 HHeiden 265-268 FFricke 269-480 FHustedt) - Aschersleben ampLeipzig

SCHMIDT R S WUNSAM U BROSCH J FOTT A LAMI H LOumlFFLER A MARCHETTOHW MUumlLLER M PRAZAacuteKOVAacute amp B SCHWAIGHOFER (1998) Late and post-glacial historyof meromictic Laumlngsee (Austria) in respect to climate change and anthropogenic impact - AquaticSci 60 56-88

SNOEIJS P amp M POTAPOVA (1998) Ecotypes or endemic species - a hypothesis on the evolutionof Diatoma taxa (Bacillariophyta) in the northern Baltic Sea - Nova Hedwigia 67 303-348

STATSOFT Inc (1995) STATISTICA for Windows [Computer program manual] - Tulsa

STERRENBURG FAS (1994) Studies on the genera Gyrosigma and Pleurosigma(Bacillariophyceae) [The species of Sullivant and Wormley 1859 synonomy and differentiationfrom other Gyrosigma taxa] - Proc Acad Nat Sci Philadelphia 146 217-236

STERRENBURG FAS (1995) Studies on the genera Gyrosigma and Pleurosigma(Bacillariophyceae) Gyrosigma acuminatum (Kuumltzing) Rabenhorst G spenceri (Quekett) Griffithand G rautenbachiae Cholnoky] - Proc Acad Nat Sci Philadelphia 146 467-480

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STERRENBURG FAS (2002) Good reasons for a marriage between taxonomy and ecologyIntroductory address International Workshop on Diatom Ecology and Taxonomy a marriage ofnecessity Szczecin-Kulice 23-27 Oct 2002 - Organizers Univ of Szczecin California Academy ofSciences Univ of Oldenburg

STOERMER EF (1975) Comparsion of the benthic diatom communities in Lake Michigan andLake Superior - Int Vereinigung Theor Limnol Verh 19 932-938

STOERMER FE (2001) Diatom taxonomy for paleolimnologists - J Paleolimnol 25 393-398

TRICK CG I F CREED MF HENRY amp DS JEFFRIES (2002) Distribution of diatoms in aforested stream containing a series of interconnected lakes - Water Air Soil Pollution Focus 2 103-128

VAN DAM H A MARTENS amp J SINKELDAM (1994) A coded checklist and ecologicalindicator values of freshwater diatoms from Netherlands - Netherlands J Aquatic Ecol 28 117-133

VAN DE VIJVER B A VAN KERCKVOORDE amp L BEYENS (2003) Freshwater and terrestrialmoss diatom assemblages of the Cambridge Bay area Victoria Island (Nuanavut Canada) - NovaHedwigia 76 225-244

VILBASTE S amp J TRUU (2003) Distribution of benthic diatoms in relation to environmentalvariables in lowland streams - Hydrobiologia 493 81-93

WASHINGTON HG (1984) Diversity biotic and similarity indices A review with special relevanceto aquatic ecosystems - Water Res 18 653-694

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ZABELINA MM IA KISSELEV AI PROSKINA-LAVRENKO amp VS SHESHUKOVA(1951) Inventory of freshwater algae of the USSR Vol 4 Diatoms 1-619 - Sov Nauka Moscow

Received 16 May 2004 accepted in revised form 18 January 2005

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as in the case of Navicula Bory sensu stricto (Lange-Bertalot 2001) or PinnulariaEhrenberg (Krammer 2000) Also Round et al (1990) triggered an intensive changein diatom nomenclature by the transfer of numerous taxa into an expanded number ofdiatom genera This activity has had a variety of results In some cases the result wasthat a taxon became assigned to a different more restricted genus (Round amp Bukhtiyarova1996 Kingston 2000) or to the same genus but by different authors (Andresen et al2000) This practice has been criticized (Kociolek 1996 Lange-Bertalot 2001) becauseof higher level transfers that are seen as simplistic and lack a proper investigation oftype specimens Type material revisions enable a much more realistic view ontaxonomy ecology and distribution of specific taxa (Sterrenburg 1994 1995) as wellas finding possibly new previously overlooked species (Cox 2003)

Revision of data on the distribution of certain cosmopolitan taxa such as Amphoracoffeaeformis (CAgardh) Kuumltzing in Argentina (Sala et al 1998) or StephanodiscusEhrenberg species (Haringkansson amp Locker 1981) brings to light possiblemisidentification mainly caused by the application of European flora keys to materialsfrom entirely different geographic areas and therefore fitting the species into knownspecies (Kociolek amp Spaulding 2000) The result has been severe criticism of thecosmopolitan distribution of diatoms (Mann amp Droop 1996 Snoeijs amp Potapova1998) and gradual abandonment of the concept in the latest revisions of specificgenera (Krammer 2000 2002 2003 Lange-Bertalot 2001) Cosmpolitian diatomdistribution can be expected in North Europe where aquatic ecosystems have repeatedlybeen influenced by glaciation events (Mann 1999 Kociolek amp Spaulding 2000)while the only centres of diatom speciation could be the old relict lakes (like LakesOhrid or Prespa in Macedonia) that have conserved their ancient conditions (Jurilj1949 1954 1956 Jerkovic 1971 Levkov et al 2003ab) Investigations of oligo-trophic lakes in Europe usually confirm the existence of new (or possibly new)diatom taxa (Lange-Bertalot amp Metzeltin 1996) and a significant diatom diversity(Hustedt 1950 Levkov et al 2001) However the number of natural ecosystems(with low anthropogenic impact) is constantly decreasing as various human impacts(Muumlller et al 1998) continue to impact environments and which consequently leadsto a decrease in the number and abundance of sensitive and rare diatom taxa (Lange-Bertalot amp Steindorf 1996) These ecosystems are therefore a subject of investigationsince diatoms are regarded as reliable indicators of acidification (Arzet et al 1986Marchetto amp Schmidt 1993 Niederhauser amp Schanz 1994) or changes in trophicstatus (Wunsam amp Schmidt 1995 Lotter et al 1998 Hall amp Smol 1999)

Besides the increasing applicability of diatoms to different types of environmentalstudies careful taxonomy becomes crucial in obtaining reliable data on diatom diversityand distribution As pointed out by Sterrenburg (2002) the basic shortcoming in thepublic concern about diversity loss is that ldquowe should first know what there is in anarea before we can determine what we are losingrdquo With particular attention to thechanges in taxonomic concepts of numerous genera and species and also the lack ofdetailed data on composition and distribution of diatom taxa in mountain aquaticecosystems in Macedonia a long-term survey (1995-2003) of these habitats on differentmountains in Macedonia has been conducted Hitherto published data on SharaMountain glacial lakes (Levkov et al 1998 2001) are scarce while there are no data

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Diatom assemblages on Shara and Nidze Mountains, Macedonia

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Transcript of Diatom assemblages on Shara and Nidze Mountains, Macedonia