Fottea, 9(1): 135–148, 2009 135

The desmids of the Swamp Nature Reserve (North Bohemia, Czech Republic) and a small neighbouring bog: composition and ecological condition of both sites

Jan Šť a s t n ý

Department of Botany, Charles University, Prague, Benátská 2, CZ – 128 01 Praha, Czech Republic

Abstract: The present study is a summary of recent findings concerning the desmid flora of the Swamp Nature Reserve and a small neighbouring bog. A total of 203 desmid taxa belonging to 22 genera were found, including 39 taxa that are considered very rare in the Czech Republic. Of this number, 46 taxa were found in the Swamp and 203 taxa in the neighbouring bog. The ecological condition of both localities was assessed on the basis of the species composition of the desmid communities encountered. The results indicate that the present overall ecological situation of the Swamp is favourable for the development of a characteristic, acidophilous desmid community. They also document quite an exceptional natural value of the second site studied, fully justifying its recent designation as part of the Swamp Nature Reserve. The results presented here may also be used in the future as a basis for comparative ecological studies tracing the development of both these sites.

Key words: bioindicators, desmids, ecology, species diversity, Swamp Nature Reserve

Introduction (1996). Despite being one of the dominant algal The Swamp Nature Reserve, designated as such groups, the desmids of the Swamp have not been in 1972, has been the subject of algological investigated in detail so far. However, among investigations since the beginning of the 20th the 195 algal taxa found Ma t t a u c h (1936) century. The scientific activities of the prominent also mentions 55 taxa of desmids. Ka l i n a & German algologist A. Pascher were closely Punčochářová (1996) found 25 desmid taxa, but connected with the Swamp; he published several since many taxa were obviously misidentified, studies focused on chrysophytes (Pa s c h e r their data should be considered with much 1909) and heterokontophytes (Pa s c h e r 1939). reserve. Pascher’s student, F. Mattauch, published a very To my knowledge the second site studied, a detailed and comprehensive study of the locality small bog adjacent to the Swamp, has not yet been (Ma t t a u c h 1936). The site is well- known due the object of any algological investigation so far. to the occurrence of two rare representatives of Desmids are particularly important Synurophyceae: Mallomonas ouradion K.Ha r r i s indicators of wetland ecosystems due to their et D.E.Br a d l e y and Mallomonas matvienkoae highly specific ecological demands (see e.g. B.As m u n d et Kr i s t i a n s e n (Ka l i n a 1969, As m u n d Co e s e l 1998, 2001, 2003, Šť a s t n ý 2005, Ha š l e r & Kr i s t i a n s e n 1986). It was a rich source of et al. 2008). The main aim of the present study material for taxonomic studies on dinophytes was therefore to summarize recent knowledge of (Pf i e s t e r & Po p o v s k ý 1979, Po p o v s k ý & Pf i e s t e r the desmid flora of both sites and to evaluate the 1990) and the green alga Dicranochaete bohemica current ecological condition of these ecosystems was also firstly described from here( No v á k o v á & on the basis of the species composition of the Po p o v s k ý 1972). The diatom flora of the locality desmid communities. Another aim of the study was investigated by St o j a n o v s k i & Ka l i n a (1989). was to evaluate the changes of the Swamp desmid The latest paper dealing with the algal flora of flora over the last 70 years by comparing my data the Swamp so far is by Ka l i n a & Punčochářová with the data of Ma t t a u c h (1936), in particular 136 Šť a s t n ý : Desmids of the Swamp with respect to possible eutrophication of the locality with alkaline lake water and the subsequent degradation of the sensitive acidophilous flora.

Locality

Both localities studied are situated in Northern Fig. 1. Location of the sampling sites in the territory of the Bohemia about 1.5 km northeast of the town of Doksy, Czech Republic and in the south-eastern part of the Máchovo in the south-eastern part of the Máchovo jezero lake jezero lake [(M) Máchovo jezero lake; (S) Swamp; (B) (Mácha´s lake) at an altitude of 265 meters a.s.l (see neighbouring bog]. Fig.1). They lie in small bays which are remnants of an original bank of a glacial lake. Their bottoms are covered with , in particular Sphagnum Material and methods auriculatum Sc h i mp . (No v á k o v á & Po p o v s k ý 1972). th Both localities developed in the 14 century due to a The algal material was collected in June 2005, May rise of groundwater level after the lake was founded. 2006 and in February and March 2007. Altogether, The Swamp Nature Reserve (total area of 1.45 ha, 72 samples were taken from both sites. The samples GPS 50°34’48.19’’ N, 14°40’4.77’’ E) is a transitional were collected by sucking off the from the bog and its water regime depends on the lake water sediment with a syringe and by squeezing out mosses level (Ka l i n a & Punčochářová 1996). Nearly the (in particular Sphagnum sp.) and dominant aquatics whole area of the Swamp is covered with Sphagnum (particularly Utricularia spp.). Shortly after sampling, sp. and the water is oligotrophic and strongly acidic part of the material was fixed with formaldehyde (pH = 3.5–4.6); only a small part of the locality is rather to a final concentration of about 4 % and later used oligomesotrophic and less acidic (pH = 5.1–5.4) due for semi-quantitative assessment according to the to overgrowing by Phragmites australis (Ca v .) Tr i n following relative scale: 1 – very rare; 2 – scattered; ex St e u d . In the southern part of the Swamp there are 3 – abundant; 4 – very abundant; 5 – mass occurrence. copious stands of Utricularia minor L. and Nymphaea Light microscopic observations were done mostly candida Pr e s l . The conductivity values measured on living material. Environmental variables (pH, -1 ranged between 88 and 174 μS.cm . The Swamp is conductivity) were measured using a Combo HI separated from the lake by a narrow reinforced dam 98129 portable instrument (HANNA, Germany), that should prevent disturbance of the locality by the microphotographs were taken using an Olympus BX51 eutrophic and alkaline (pH = 7.5–8.0) lake water. light microscope and Olympus Camedia C – 5050Z The second site studied (total area of ca 2.5 ha) digital microphotographic equipment. is situated about 350 m south-southeast of the Swamp, The relative rarity of the individual taxa within in the so-called Břehyňská Bay, north of the Břehyňský the Czech Republic was evaluated on the basis of the stream inflow into the lake (GPS 50°34’34.44’’ N, author’s extensive personal experience and classified 14°40’14.45’’ E). It is a transitional bog and its water according to the following scale: 1 – scattered regime also depends on the lake water level. This occurrence; 2 – rare taxon; 3 – very rare taxon. In locality, however, is more fragmented than the Swamp. general, taxa labelled as very rare in the Czech Republic There are distinct gradients of acidity and trophy. The are also rare in Central Europe (see e.g. Rů ž i č k a 1977, most extensive eastern part of the locality, covered 1981, Gu t o w s k i & Mo l l e n h a u e r 1996, Le n z e n w e g e r with Sphagnum sp., is oligotrophic and strongly acidic 2003). (pH = 3.4–4.2) and in one pool extensive stands of Detrended correspondence analysis (DCA) Utricularia minor have developed. Closer to the lake using Canoco for Windows 4.5 (Te r Br a a k & this part gradually changes into a mesotrophic littoral Šm i l a u e r 1998) was done for statistic evaluation. reed belt. The reaction of the water in this part is The evaluation focused on the differences in species slightly acidic (pH = 5.0–6.3), there are a number of composition between the sampling sites. The basic shallow pools filled in withUtricularia australis R.Br. detrending analysis was carried out with respect to and the littoral reed belt gradually merges into the lake. downweighting of rare species. The conductivity values measured at this site ranged from 54 to 156 μS.cm-1. This second site became a part of the newly- established Swamp Nature Reserve in June 2007. Results However, since the two localities have a somewhat different character and a completely different history A total of 203 desmid taxa belonging to 22 genera concerning algological investigations, they are were identified; 46 taxa were found in the Swamp discussed separately in this paper. and 203 taxa in the neighbouring bog (Table 1). Fottea, 9(1): 135–148, 2009 137

Eight taxa ( lineatum, oligotrophic and strongly acidic part basically margaritiferum, Cylindrocystis brebissonii, two main types of habitats may be distinguished. dissiliens, spirostriolatum, The very shallow, periodically desiccating pools teliferum, extensus are dominated by adaptive, frequently dividing var. isthmosus and armatum) were species (in particular Cylindrocystis brebissonii, found forming zygospores. laevis, cucurbita and Closterium striolatum). On the other hand, large The Swamp species with a much lower growth rate (Co e s e l The species composition of the desmid 1982), e.g. jenneri, M. oscitans, communities from the Swamp corresponds M. truncata, Cosmarium ralfsii or Xanthidium well with the very low pH of the water. Most armatum, require deeper pools with a more stable species found may be characterized as strictly water regime for their development. The species acidophilous; the dominant species (Cylindrocystis richness of the samples from this part of the locality brebissonii, Tetmemorus laevis, Closterium is rather poor, comprising mostly 10-20 desmid striolatum, Staurastrum punctulatum, St. simonyi) taxa (the highest number generally occurring in belong to the most common acidophilous desmids, deeper, stable pools). However, the abundance of frequently occurring in all kinds of strongly acidic those particular taxa is generally very high and the water bodies. The extreme milieu conditions were majority of the species mentioned occur locally in also reflected by the generally rather poor species great mass (see Fig. 2). richness of the site as a whole as well as by the The species richness on the transect from poor species richness of the individual samples, the oligotrophic, strongly acidic part of the site which ranged from 2 to 11 taxa per sample and to the lake sublittoral exhibits a characteristic slightly increased up to 18 taxa per sample only in pattern influenced by the pH and trophy gradient: the eutrophic part with overgrowing Phragmites transitional, oligomesotrophic zone with 20- australis. This local increase in the trophic state 30 taxa, slightly acidic, mesotrophic zone with was also reflected in the presence of several up to 90 taxa and eutrophic lake sublittoral taxa preferring oligomesotrophic (Closterium 32 taxa. However, with the increasing species baillyanum, Cl. juncidum, Cosmarium amoenum, richness the abundance of the individual taxa in C. pseudopyramidatum, Micrasterias truncata) or a sample decreases and hence also the number even mesotrophic conditions ( ansatum, of the characteristic taxa (see Fig. 2). Typical Staurastrum lapponicum, Cosmarium quadratum, representatives of the transitional, oligomesotro- Closterium parvulum). phic zone are e.g. Closterium baillyanum, Cosma- The taxa Cosmarium botrytis var. rium pyramidatum, C. pseudopyramidatum, C. mediolaeve and Cosmarium cf. tenue prefere tinctum, Penium cylindrus and Haplotaenium mesotrophic or slightly eutrophic conditions and rectum. The mesotrophic, slightly acidic littoral obviously are not autochtonous in the Swamp zone is characterized by the co-occurrence of and originate most likely from the lake littoral many generally rare taxa, and moreover in this zone. This hypothesis is also supported by the fact part of the locality it is even possible to find some that they have been found near the lake and in a representatives of the eutrophic lake tychoplankton minimal abundance (one of each species). (e.g. Closterium venus, Staurastrum eurycerum, The most rare and interesting findings are Cosmarium humile, C. wittrockii, C. reniforme, Micrasterias jenneri, Euastrum luetkemuelleri C. subgranatum, etc.). var. carniolicum, Cosmarium prominulum var. The ordination diagram (Fig. 2), which in- subundulatum, C. subquadrans var. minor and cludes the samples from 2005 and 2006, illustrates Actinotaenium perminutum. The finding of a the situation in the bog. The first two of the DCA robust form of the rather recently described and ordination axes explained 26.2 and 5.7% of the very rare Haplotaenium indentatum (Ko u w e t s variance of the species data, respectively. The first 1991) is also remarkable. ordination axis represents the pH gradient with acidic sites on the left. This is in accordance with The small bog the data published by Co e s e l (1998) and Šť a s t n ý The composition of the desmid communities from (2007), who consider the water pH the key factor the second site studied is distinctly determined influencing desmid distribution. The factor being by the pH and trophy gradients. In its eastern, responsible for the distribution of the samples 138 Šť a s t n ý : Desmids of the Swamp

Fig. 2. The results of DCA from site 2 (the bog) in the ordination space of the first two axes (circles = samples; triangles = species). The size of the circles correlates with the species richness of the samples. Only the statistically most relevant species are depicted. Taxon abbreviations: (ACCUC) Actinotaenium cucurbita; (CLBAI) Closterium baillyanum; (CLSTR) Cl. striolatum; (COCOT) Cosmarium contractum; (COPSP) C. pseudopyramidatum; (COPYR) C. pyramidatum; (CORAL) C. ralfsii; (COTIN) C. tinctum; (CYBRE) Cylindrocystis brebissonii; (HAIND) Haplotaenium indentatum; (HAREC) H. rectum; (MIJEN) Micrasterias jenneri; (MITRU) M. truncata; (NEDIG) Netrium digitus; (PECYL) Penium cylindrus; (STPOL) Staurastrum polymorphum; (TTGRA) Tetmemorus granulatus; (TTLAE) T. laevis; (XAARM) – Xanthidium armatum. with similar pH values is most likely the depth deterioration of previously rich desmid sites of the pools. The samples depicted in the lower have been reported as a result of processes like part of the ordination diagram were taken from eutrophication, acidification, desiccation or generally deeper pools than the samples depicted cultivation. The situation in the Netherlands in the upper part of the diagram. is documented by Co e s e l et al. (1978) and From the total of 203 taxa occurring in the Co e s e l (2003). Le n z e n w e g e r (1996) notes the bog, 102 taxa may be marked as more or less rare same phenomenon in Austria and Ši m e k (1997) in the Czech Republic, 39 of those even as very documents very convincingly the drastic influence rare (see Table 1). The most interesting findings of eutrophication on the desmid communities at are depicted below (Fig. 3, Fig. 4). the famous Czech algological site Řežabinec. The high susceptibility of desmids to milieu changes and their short generation time Discussion make them very suitable (in contrast to vascular ) to monitor short-term environmental Desmids are excellent bioindicators, not only changes. Particularly in the Netherlands, desmids with respect to their preference for certain have recently been used as bioindicators for the environmental conditions but also with respect assessment of water quality and, in a number of to the current ecological condition and stability nature restoration projects, for the assessment of of the given ecosystem (Co e s e l 1998, 2001). In nature conservation value (see e.g. Co e s e l 2003). particular due to their highly specific ecological With respect to the above-mentioned demands, desmids have recently become the most information, it was very interesting to compare endangered group of aquatic microorganisms, the current desmid flora of the Swamp with the at least in densely populated countries (Co e s e l data published by Ma t t a u c h (1936), in particular 1998). During the last decades, many cases of as regards the potential threat posed to the site by Fottea, 9(1): 135–148, 2008 139

Fig. 3. (a) Micrasterias oscitans; (b) M. jenneri; (c) M. apiculata; (d) Euastrum pinnatum; (e) E. turneri; (f) E. luetkemuelleri var. carniolicum; (g) simplicissimum; (h) Haplotaenium indentatum, morpha. Scale bar 30 μm; e, f 10 μm. the eutrophic lake water. (1936) are more typical of mesotrophic, slightly acidic water bodies. This was undoubtedly due The Swamp to the presence of a narrow littoral reed belt Whereas the vast majority of the 46 taxa that (see Ma t t a u c h 1936, p.386, 427), which offered have been recently found at the Swamp may be suitable conditions for the survival of the species characterized as strictly acidophilous, preferring preferring the above-mentioned mesotrophic strongly acidic, oligotrophic waters (see above), conditions. In addition, this fact may explain approximately 20 taxa mentioned by Ma t t a u c h the higher species diversity (55 taxa) found by 140 Šť a s t n ý : Desmids of the Swamp

Fig. 4. (a) baileyi var. caelatum; (b) aculeatum; (c) Cosmarium ralfsii; (d) C. carinthiacum; (e) C. obsoletum; (f) C. ocellatum var. notatum; (g) C. tetrachondrum; (h) C. paragranatoides. Scale bar 10 μm.

Ma t t a u c h (1936). Desmids show the highest sensitive species, characteristic of undisturbed, diversity in mesotrophic, slightly acidic water stable environment (Micrasterias oscitans, M. bodies (see e.g. Co e s e l 1982, 1998) and, in jenneri, Cosmarium ralfsii, Xanthidium armatum, general, at sites with distinct gradients in acidity Euastrum insigne, Staurastrum hystrix; see e.g. and nutrient content. Co e s e l 1998) have disappeared. In contrast, the However, even more interesting is the relative distribution of the adaptive taxa with higher qualitative aspect of the comparison of the species growth rates increased (see Table 1). This may composition: most of the ecologically highly produce evidence of lower stability of the biotope Fottea, 9(1): 135–148, 2009 141 that is most likely caused by occasional overflow is a more varied and better preserved copy of the of the alkaline, highly eutrophic lake water into the Swamp appears quite likely. This hypothesis is locality (see Ka l i n a & Punčochářová 1996). In also supported by the much higher desmid species the long-term aspect, it does not lead to significant richness of this site as well as by the occurrence increases in the pH and trophic state of the site of most of the ecologically highly sensitive taxa due to the buffering capacity of the Sphagnum. that have already presumably disappeared from However, it is very likely that such occasional the Swamp (see above). Whereas only 18 of the disturbances may have caused the extinction of 55 desmid taxa reported by Ma t t a u c h (1936) the most sensitive acidophilous species. have recently been found in the Swamp (see Table Despite the above-mentioned facts, the 1), at the second site the number of shared taxa is present ecological condition of the Swamp (with much higher (42). respect to the occurrence of desmids) seems to be With respect to the occurrence of desmids, rather favourable, and particularly the occurrence the above-mentioned facts clearly illustrate the of some very rare taxa (see above) makes it a very uniqueness of the second site studied. But given valuable biotope. the fact that a high nature value of a particular site, indicated by the composition of desmid The small bog communities, is usually a good indicator of its Looking at the composition of the desmid general biological value (Co e s e l 1998), I consider communities at the second site, the first remarkable the second site one of the most valuable wetland feature is its very high species richness, which habitats in the Czech Republic, deserving further is particularly due to the large pH and trophy detailed investigations. In addition, the present gradient and the high number of microhabitats results fully justify the recent designation of the available (see above). There is only one place discussed site as a part of the Swamp Nature among recently studied localities in the Czech Reserve. Republic which has higher species richness: the In the future, the results presented here Břehyně-Pecopala Nature Reserve (319 taxa, may be used as a basis for comparative ecological Šť a s t n ý , unpublished data). Also remarkable is studies aimed at the future development of both the richness of the locality in rare taxa (102 taxa, these sites. i.e. 50 %, see Table 1). Moreover, the majority of the rare taxa found are characteristic and indicative of well-preserved, stable ecosystems (see Co e s e l Acknowledgement 1998). Yvonne Němcová is acknowledged for information on The rarest findings include Micrasterias occurrence of representatives of the class Synurophy- oscitans, Pleurotaenium simplicissimum and ceae at the sites discussed. The study was supported by the research project no. 21620828 of the Czech Euastrum pinnatum. All these taxa are considered Ministry of Education, and by the Charles University to be extremely rare in Central Europe (Rů ž i č k a Science Foundation B BIO 164/2006 grant. 1977, 1981). In the existing desmid checklists of various European countries (Gu t o w s k i & Mo l l e n h a u e r 1996, Co e s e l 1998, Ma r h o l d & References Hi n d á k 1998, Ko u w e t s 1999, Abd e l a h a d et al. 2003, Le n z e n w e g e r 2003, Po u l í č k o v á et al. 2004) Abd e l a h a d , N., Ba z z i c h e l l i , G. & D‘Ar c h i n o , these taxa are either missing or their findings date R. (2003): Catalogo delle from the 19th or the early 20th century, which (, ) segnalate makes it very likely that they are already extinct in Italia. (A checklist of Desmids (Chlorophyta, or close to extinction. Zygnematophyceae) reported in Italy). – 120 Another very remarkable fact is that pp., Academia Nazionale delle Science detta all the 46 desmid taxa found at the Swamp dei XL, Roma. As m u n d , B. & Kr i s t i a n s e n , J. (1986): The were recorded also from the second site. In Mallomonas (Chrysophyceae). – Opera botanica addition, the distribution of the representatives 85: 5–128. of Synurophyceae at both these sites (Ně m c o v á , Co e s e l , P.F.M. (1982): Structural characteristics and pers. com.) is also very similar. This supports the adaptations of desmid communities. – J. Ecol. impression that the two sites are very similar in 70: 163–177. character, and the hypothesis that the second site Co e s e l , P.F.M. (1998): Sieralgen en Natuurwaarden. 142 Šť a s t n ý : Desmids of the Swamp

Table 1. List of all taxa found with data indicating their rarity and frequency of their occurrence at the localities studied [(S) Swamp; (B) neighbouring bog; (R) rarity within the Czech Republic; * species found by Mattauch (1936)].

S B R Actinotaenium cruciferum (De Ba r y ) Te i l i n g *

Actinotaenium cucurbita (Br é b .) Te i l i n g 2* 4

Actinotaenium diplosporum (P.Lu n d e l l ) Te i l i n g 1 1

Actinotaenium inconspicuum (W.et G.S.We s t ) Te i l i n g 1 2

Actinotaenium perminutum (G.S.We s t ) Te i l i n g 1 1 3 Actinotaenium silvae – nigrae (Ra b a n u s ) Ko u w e t s et Co e s e l 2 1

Actinotaenium turgidum (Br é b .) Te i l i n g * 1 Actinotaenium sp.1 1 2 Actinotaenium sp.2 1

Bambusina brebissonii Kü t z i n g ex Kü t z i n g * 2

Closterium abruptum W.We s t 2 2 1 Closterium acutum Br é b . 3 2

Closterium angustatum Ra l f s 2 1

Closterium archerianum Cl e v e 1 2

Closterium attenuatum Ra l f s 1

Closterium baillyanum Br é b . 2 2

Closterium baillyanum var. alpinum (Vi r e t ) Gr ö n b l a d 2

Closterium calosporum var. brasiliense Bø r g e s . 2 3

Closterium closterioides var. intermedium (J.Ro y et Bi s s e t ) Ru z i c k a * 2 3 Closterium cornu Ra l f s 2 1

Closterium cornu var. upsaliense No r d s t . 2 2

Closterium costatum Ra l f s 2

Closterium cynthia De No t . 1

Closterium delpontei (G.A.Kl e b s ) Wo l l e 1 3

Closterium dianae Ra l f s * 2

Closterium dianae var. arcuatum (Br é b .) Ra b e n h o r s t 2

Closterium didymotocum Ra l f s 2 2

Closterium directum W.Ar c h e r 1 2

Closterium gracile Ra l f s 3

Closterium intermedium Ra l f s 2

Closterium juncidum Ra l f s 2 3

Closterium kuetzingii Br é b . * 2

Closterium limneticum Le mm e r m . 1

Closterium lineatum var. elongatum (Ro s a ) Cr o a s d a l e 3

Closterium lunula Ra l f s 3

Closterium navicula (Br é b .) Lü t k e m . 1

Closterium nematodes var. proboscideum W.B.Tu r n e r 1 3

Closterium parvulum Nä g e l i 2* 2

Closterium pronum Br é b . 1 2

Closterium ralfsii var. hybridum Ra b e n h o r s t 2 Fottea, 9(1): 135–148, 2009 143

Table 1 Cont.

Closterium rostratum Ra l f s *

Closterium setaceum Ra l f s 2 1

Closterium striolatum Ra l f s 4* 3

Closterium tumidum var. nylandicum Gr ö n b l a d 2 2 1 Closterium turgidum Ra l f s 1

Closterium venus Ra l f s 1

Closterium cf. macilentum Br é b . 1

Closterium cf. pusillum var. monolithum Wi t t r o c k 1

Cosmarium amoenum Br é b . 2 2 1 Cosmarium angulosum Br é b . 2 2

Cosmarium blytii var. novae – sylvae W.et G.S.We s t 2 2

Cosmarium boeckii Wi l l e 1

Cosmarium botrytis var. mediolaeve W.We s t 1 1

Cosmarium carinthiacum Lü t k e m . 2 3

Cosmarium connatum Br é b . * 2

Cosmarium conspersum var. latum (Br é b .) W.et G.S.We s t 1 2

Cosmarium contractum Ki r c h n e r * 2 2 Cosmarium contractum var. ellipsoideum (El f v i n g ) W.et G.S.We s t 2 2

Cosmarium contractum var. minutum (De l p o n t e ) W.et G.S.We s t 1

Cosmarium contractum var. retusum (W.et G.S.We s t ) Wi l l i Kr i e g . et Ge r l o f f 1 2

Cosmarium crenulatum Nä g e l i 1

Cosmarium debaryi W.Ar c h e r 2 1

Cosmarium difficileL ü t k e m . 1 1

Cosmarium formosulum Ho f f 1

Cosmarium goniodes var. subturgidum W.et.G.S.We s t 1 1 1 Cosmarium granatum Br é b . 1

Cosmarium humile (F.Ga y ) No r d s t . 2

Cosmarium isthmochondrum No r d s t . *

Cosmarium jaoi Ko u w e t s 1 3

Cosmarium margaritaceum Me s s i k . *

Cosmarium margaritiferum Ra l f s 3 1

Cosmarium obsoletum (Ha n t z s c h ) Re i n s c h 2 3

Cosmarium obtusatum Sc h m i d l e 1

Cosmarium ocellatum B.Ei c h l e r et Gu t w . *

Cosmarium ocellatum var. notatum (No r d s t .) Wi l l i Kr i e g . et Ge r l o f f 1 3

Cosmarium ordinatum (Bö r g e s .) W.et.G.S.We s t 2 3

Cosmarium ornatum Ra l f s 2 1

Cosmarium pachydermum P.Lu n d e l l 1

Cosmarium paragranatoides Sk u j a 2 3

Cosmarium perforatum P.Lu n d e l l 1 3

Cosmarium prominulum var. subundulatum W.et.G.S.We s t 2* 1 3 Cosmarium pseudoornatum B.Ei c h l e r et Gu t w . 2 2 144 Šť a s t n ý : Desmids of the Swamp

Table 1 Cont.

Cosmarium pseudopyramidatum P.Lu n d e l l 2* 2 2 Cosmarium punctulatum var. subpunctulatum (No r d s t .) Bö r g e s . 1

Cosmarium pyramidatum Br é b . * 3 2 Cosmarium quadratulum var. boldtii (Me s s i k .) Wi l l i Kr i e g . et Ge r l o f f 1

Cosmarium quadratum Ra l f s 2 2

Cosmarium quadrum P.Lu n d e l l 1 1

Cosmarium quadrum var. sublatum (No r d s t .) W.et.G.S.We s t 1 3

Cosmarium ralfsii Br é b . * 3 3 Cosmarium rectangulare Gr u n o w *

Cosmarium regnellii Wi l l e 2

Cosmarium reniforme (Ra l f s ) W.Ar c h e r * 1

Cosmarium retusum (Pe r t y ) Ra b e n h o r s t 1 3

Cosmarium sexnotatum var. tristriatum (Lü t k e m .) Sc h m i d l e 2 3

Cosmarium simplicius (W.et.G.S.We s t ) Gr ö n b l a d 1 3

Cosmarium subcostatum var. minus (W.et.G.S.We s t ) Fö r s t e r 1

Cosmarium subcucumis Sc h m i d l e 1 1

Cosmarium subgranatum (No r d s t .) Lü t k e m . 1

Cosmarium subquadrans var. minor Sy m o e n s ex Co e s e l 1 1 3 Cosmarium subtumidum No r d s t e d t 2 2

Cosmarium tetrachondrum P.Lu n d e l l 1 3

Cosmarium tetraophthalmum Br é b . 2

Cosmarium tinctum Ra l f s 2* 2

Cosmarium vogesiacum Le m a i r e 2 3

Cosmarium wittrockii P.Lu n d e l l 2 2

Cosmarium cf. pseudoretusum Du c e l l . 1

Cosmarium cf. tenue W.Ar c h e r 1 1 Cosmarium sp.1 1 2 Cosmarium sp.2 2

Cylindrocystis brebissonii (Ra l f s ) De Ba r y 4* 4

Cylindrocystis crassa De Ba r y *

Cylindrocystis gracilis I. Hi r n 2 2

Desmidium aptogonum Kü t z i n g 2 2

Desmidium baileyi var. caelatum (Ki r c h n e r ) No r d s t . 2 3

Desmidium swartzii Ra l f s 2

Euastrum ansatum Ra l f s 3 2

Euastrum ansatum var. rhomboidale F. Du c e l l . 1 2

Euastrum binale var. gutwinskii (Sc h m i d l e ) Ho m f e l d 2* 2

Euastrum biscrobiculatum (Wo l o s z .) Co e s e l 1 3

Euastrum denticulatum F.Ga y 2

Euastrum dubium var. ornatum Wo l o s z . 2 1

Euastrum elegans Ra l f s * 2 1 Euastrum gayanum De To n i 2 Fottea, 9(1): 135–148, 2009 145

Table 1 Cont.

Euastrum germanicum (Sc h m i d l e ) Wi l l i Kr i e g . 1 2

Euastrum humerosum Ra l f s 2

Euastrum insigne Ra l f s *

Euastrum insulare var. silesiacum (Gr ö n b l a d ) Wi l l i Kr i e g . 1 2

Euastrum luetkemuelleri var. carniolicum (Lü t k e m .) Wi l l i Kr i e g . 1 2 3 Euastrum oblongum Ra l f s * 2

Euastrum pectinatum Ra l f s 2 1

Euastrum pinnatum Ra l f s 1 3

Euastrum pulchellum Br é b . 2 3

Euastrum subalpinum Me s s i k . 1 1 1 Euastrum turneri W.We s t 1 3

Euastrum verrucosum Ra l f s * 1

Gonatozygon aculeatum Ha s t i n g s 2 3

Gonatozygon brebissonii De Ba r y 2

Haplotaenium indentatum Ko u w e t s , morpha 3 3 3 Haplotaenium minutum (Ra l f s ) Ba n d o * 2 1 Haplotaenium rectum (De l p o n t e ) Ba n d o 2 2

Hyalotheca dissiliens Ra l f s 1* 2

Hyalotheca dissiliens var. tatrica Ra c i b o r s k i 2 1

Mesotaenium macrococcum (A.Br a u n ) J.Ro y et Bi s s e t 1

Micrasterias apiculata Ra l f s 2 3

Micrasterias crux – melitensis Ra l f s 1

Micrasterias denticulata var. angulosa (Ha n t z s c h ) W.et G.S.We s t 2 2

Micrasterias fimbriata Ra l f s 1 2

Micrasterias jenneri Ra l f s 3* 3 3 Micrasterias oscitans Ra l f s * 1 3 Micrasterias papillifera Br é b . 1

Micrasterias rotata Ra l f s 2

Micrasterias thomasiana var. notata (No r d s t .) Gr ö n b l a d 2

Micrasterias truncata Br é b . 2* 2

Micrasterias truncata var. quadrata Bu l n h . 2 2

Netrium digitus (Br é b .) It z i g s . et Ro t h e 2* 2

Netrium interruptum (Br é b .) Lü t k e m . 2 2

Netrium oblongum (De Ba r y ) Lü t k e m . *

Penium cylindrus Br é b . 1* 2

Penium exiguum W.We s t 1 3

Penium spirostriolatum J.Ba r k e r 1 1 Pleurotaenium archeri De l p o n t e 1 2 Pleurotaenium ehrenbergii (Ra l f s ) De Ba r y * 1

Pleurotaenium simplicissimum Gr ö n b l a d 1 3

Pleurotaenium trabecula Nä g e l i * 1

Pleurotaenium truncatum (Br é b .) Nä g e l i 1 146 Šť a s t n ý : Desmids of the Swamp

Table 1 Cont.

Pleurotaenium sp. 1

Sphaerozosma filiforme Ra l f s 1 3

Spirotaenia cf. erythrocephala It z i g s . 1

Staurastrum aculeatum Ra l f s 2 2

Staurastrum alternans Br é b . in Ra l f s 1

Staurastrum brachiatum Ra l f s 2 2

Staurastrum controversum Ra l f s * 1 1 Staurastrum dilatatum Ra l f s 2 2

Staurastrum eurycerum Sk u j a 1 2

Staurastrum gracile Ra l f s *

Staurastrum hirsutum Br é b . 2

Staurastrum hirsutum var. arnellii (Bo l d t ) Co e s e l *

Staurastrum hirsutum var. muricatum (Br é b . ex Ra l f s ) Fö r s t e r 2 2

Staurastrum hystrix Ra l f s * 2 3 Staurastrum inflexum Br é b . 2

Staurastrum lapponicum (Sc h m i d l e ) Gr ö n b l a d 2 1

Staurastrum margaritaceum Ra l f s 1 1

Staurastrum minimum Co e s e l 2 3

Staurastrum muticum Ra l f s 1

Staurastrum orbiculare var. depressum J.Ro y et Bi s s e t 1

Staurastrum pelagicum W.et G.S.We s t *

Staurastrum polymorphum var. pygmaeum Gr ö n b l a d 3

Staurastrum polytrichum (Pe r t y ) Ra b e n h o r s t 1 1

Staurastrum punctulatum Br é b . 3* 2

Staurastrum reinschii Ro y *

Staurastrum scabrum Br é b . 2 2 2 Staurastrum simonyi He i m e r l 3* 2

Staurastrum teliferum Ra l f s * 2

Staurastrum tetracerum Ra l f s 1

Staurastrum vestitum Ra l f s 2 3

Staurastrum cf. boreale W.et G.S.We s t 1

Staurastrum cf. crenulatum var. continentale Me s s i k . 2

Staurodesmus controversus (W.et G.S.We s t ) Te i l i n g 2 2 2 Staurodesmus convergens (Ra l f s ) S.Li l l . 1 1

Staurodesmus cuspidatus (Ra l f s ) Te i l i n g 1

Staurodesmus dejectus var. apiculatus (Br é b .) Te i l i n g * 1

Staurodesmus extensus var. isthmosus (He i m e r l ) Co e s e l 1 1 2 Staurodesmus glaber (Ra l f s ) Te i l i n g 1

Staurodesmus incus (Br é b .) Te i l i n g * 2 2 Staurodesmus omearae (W.Ar c h e r ) Te i l i n g 1 1

Staurodesmus triangularis var. subparallelus (Sm i t h ) Th o m . 1 1

Teilingia excavata var. subquadrata (W.et G.S.We s t ) St e i n 2 1 Fottea, 9(1): 135–148, 2009 147

Table 1 Cont.

Teilingia granulata (J.Ro y et Bi s s e t ) Bo u r r e l y 2

Tetmemorus granulatus Ra l f s 2* 3

Tetmemorus laevis Ra l f s 4* 4

Tetmemorus laevis var. minutus (De Ba r y ) Pl a y f a i r 2* 3 1 Xanthidium antilopaeum (Br é b .) Kü t z i n g 1

Xanthidium antilopaeum var. planum Ro l l 1

Xanthidium armatum Ra l f s * 3 2 Xanthidium cristatum Br é b . 1 3

Xanthidium octocorne Ra l f s 1 1

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© Czech Phycological Society Received June 24, 2008 Accepted October 15, 2008