Biologia 63/6: 901—906, 2008 Section Botany DOI: 10.2478/s11756-008-0140-x

Diversity of desmids in three Thai peat swamps*

Neti Ngearnpat1, Peter F.M. Coesel2 &YuwadeePeerapornpisal1

1Department of Biology, Faculty of Science, Chiang Mai University,Chiang Mai 50200, Thailand; e-mail: [email protected], [email protected] 2Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Kruislaan 318,NL-1098 SM Amsterdam, The Netherlands; e-mail: [email protected]

Abstract: Three peat swamps situated in the southern part of Thailand were investigated for their desmid flora in relation to a number of physical and chemical habitat parameters. Altogether, 99 were encountered belonging to 22 genera. 30 species are new records for the Thai desmid flora. Laempagarung peat swamp showed the highest diversity (45 species), followed by Maikhao peat swamp (32 species) and Jud peat swamp (25 species). Despite its relatively low species richness, Jud swamp appeared to house a number of rare taxa, e.g., subdenticulata var. ornata, M. suboblonga var. tecta and M. tetraptera var. siamensis which can be considered Indo-Malaysian endemics. Differences in composition of the desmid flora between the three peat swamps are discussed in relation to environmental conditions. Key words: desmids; ecology; peat swamps; Indo-Malaysian region; Thailand

Introduction

The desmid flora of Thailand has been investigated by foreign scientists for over a hundred years. The first records of desmids were published by West & West (1901). After that there were reports by Hirano (1967, 1975, 1992), Yamagishi & Kanetsuna (1987), Coesel (2000) and Kanetsuna (2002). The checklist of algae in Thailand (Wongrat 1995) mentions 296 desmid species plus varieties, belonging to 22 different genera. How- ever, when compared with many other tropical coun- tries, the Thai desmid flora is less well documented in terms of diversity, distribution patterns and related eco- logical data. From 2005 on, the first author aims to fill this scientific gap by investigating as many potential desmid sites as possible, distributed all over the Thai country. The present paper reports the results from some three peat swamps, a habitat that is very rare in Thailand.

Material and methods

Study sites The peat swamps in question are situated in the south- ◦  ◦  Fig. 1. Map showing location of the three peat swamps investi- ern part of Thailand, between 18 47 and 19 52 North and gated. 98◦59 to 99◦50 East (Fig. 1). The climate in this region is tropical, with well demarcated rainy and dry seasons. Nor- mally, the dry season is from November to April and the Laempagarung peat swamp (LP) rainy season from May to October. Samples were collected Located in the province of Phung-Nga, 9 m above sea level. in the year 2006. The samples were taken from a pool which was 1–2 m in

* Presented at the International Symposium Biology and of Green Algae V, Smolenice, June 26–29, 2007, Slovakia.

c 2008 Institute of Botany, Slovak Academy of Sciences 902 N. Ngearnpat et al.

Table 1. Physical and chemical features and trophic status of the 5.6–6.1). Conductivity values in Laempagarung and three swamps investigated. LP – Laempagarung peat swamp; MP Maikhao swamps were slightly above 300 µScm−1, but – Maikhao peat swamp; JP – Jud peat swamp. distinctly lower in Jud swamp (81 µScm−1). Laempa- Sampling site LP MP JP garung swamp appeared to be lowest in nitrate concen- tration but highest in ammonium concentration. Temperature ( ◦C) 29.1 32.2 28.8 Concentration of soluble reactive phosphorus in pH 5.9 5.6 6.1 Jud swamp was about ten times higher than in Laempa- Conductivity (µScm−1) 315 327 81 DO (mg L−1)84.24garung and Maikhao swamps. The trophic status of all −1 Total Alkalinity (mg CaCO3 L ) 11 11.5 19 three swamps was determined to be oligo-mesotrophic, − −1 NO3 (mg L ) 0.06 2.4 1.8 in combination with a moderate water quality. + −1 NH4 (mg L ) 0.13 0.04 0.04 A total of 99 species and nine additional vari- + −1 PO4 (mg L ) 0.09 0.07 0.85 eties were found (Table 2). They belong to 22 genera, i.e., Actinotaenium (2 species), Bambusina (1 species), Trophic status meso meso meso Closterium (14 species, 3 varieties), (24 species, 1 variety), Cylindrocystis (1 species, 1 vari- ety), Desmidium (5 species), Euastrum (11 species), depth. Most of the bottom was beset with Hydrilla verticil- Gonatozygon (1 species),Hyalotheca(2 species),Mi- lata and Utricularia sp.. crasterias (8 species), Netrium (1 species, 2 vari- eties),Penium(2 species), Pleurotaenium (6 species, Maikhao peat swamp (MP) Located in the province of Phaket, near the village of 1variety), Phymatodocis (1 species), Spondylosium (1 Maikhao, 5 m above sea level. The samples were taken species), Staurastrum (12 species, 1 variety),Stau- from a small pool covered with Salvinia spec., Pistia sp. rodesmus (1 species),Teilingia(1 species), Tetmemorus and Utricularia sp. (1 species), Triploceras (1 species) and Xanthidium (2 species). The distribution of the taxa over the three Jud peat swamp (JP) peat swamps is listed in Table 2. Located in the province of Trung, 15 m above sea level. The Of the three swamps investigated, Laempagarung, swamp, surrounded by agricultural areas, is subjected to various human activities. The samples were collected from with 13 genera, 45 species and 2 additional varieties, ap- a small drainage-canal with mats of filamentous algae and peared to house the most diverse desmid flora. The sec- Utricularia sp. along the banks. ond richest was Maikhao swamp (11 genera, 32 species and 2 additional varieties) whereas the lowest diversity Sampling and analytical methods was found in Jud swamp (11 genera, 28 species and two Water samples were collected in polyethylene bottles and additional varieties). kept in a cool box (5–7 ◦C). Planktonic desmids were col- µ lected by filtering 20 liters of water through 10 mmeshsize Discussion plankton net. Periphytic desmids were collected by suck- ing 12 mL of water by a cylindrical syringe set from the surface of any bottom sediment or submerged . This The total number of 99 species collected from the peat procedure was repeated at 5 different locations on each sam- swamps investigated is not particularly high when com- pling site. Subsequently, the 5 samples were put together as pared with species numbers found in European peat a 60 mL mixed sample (Watanabe & al. 2000). The algal lands (e.g. Coesel 1981). Likely, this may be related to samples were preserved with Lugol’s iodine solution. The the relatively poor development (in size and internal desmid species were identified at a magnification of 400× × differentiation) of the three Thai peat swamps under and 1000 . Drawings were made with the help of a drawing discussion. tube (camera lucida). Water temperature, conductivity, pH and dissolved When comparing the species composition of Laem- oxygen (DO) were measured in the field using portable pagarung, Maikhao and Jud swamps, it is striking meters. Alkalinity, ammonium nitrogen, nitrate nitrogen that they have not a single species in common. Only and soluble reactive phosphorus (SRP) were determined in four species, i.e. Bambusina borreri, Cosmarium con- the laboratory by phenolphthalein methyl orange indicator natum, Micrasterias foliacea and Pleurotaenium tra- method, nesslerization method, cadmium reduction method becula) were encountered in two of the three swamps, and ascorbic method (APHA, AWWA, WPCF, 1998), re- all the other ones were found in but one of the three spectively. swamps (Table 2). Assuming that the swamps have The trophic status of the water was classified according been sampled in an adequate way, it might indi- to the methods of Wetzel (2001) and Lorraine & Vollenwei- der (1981). The parameters DO, conductivity and the con- cate that the ecological conditions in the swamps are − + 3− centrations of NO3 ,NH4 and PO4 were used to calculate different. the water quality index according to the modified method Laempagarung swamp is characterized by a (rel- of Peerapornpisal et al. (2004). atively) large number of Cosmarium and Staurastrum species (Fig. 2). This might be attributed to the nature Results of the water body sampled, i.e., a relatively deep, large pool in which planktic species are to be expected. Many Results of the chemical analyses are given in Table 1. of the Cosmarium and Staurastrum species encountered Water at all sampling sites was slightly acidic (pH in this pool are adapted to a planktic way of life, either Desmids of Thai peat swamps 903

Table 2. Desmid taxa encountered in the three swamps investigated. LP – Laempagarung peat swamp; MP – Maikhao peat swamp; JP – Jud peat swamp. The symbol (*) indicates a new record for Thailand.

Mesotaeniaceae Cylindrocystis brebissonii (Ralfs) De Bary var. brebissonii JP C. brebissonii var. minor W. et G.S.West * JP Netrium digitus (Bréb.) Itzigs. & Rothe var. digitus MP N. digitus var. lamellosum (Bréb.) Gr¨onblad * MP N. digitus var. parvum (Borge) Willi Krieg.* MP Actinotaenium adelochondrum (Elfving) Teiling forma * MP A. curtum (Bréb.) Teiling MP A. turgidum (Bréb.) Teiling var. ovatum (Nordst.) Teiling MP Bambusina borreri (Ralfs) Cleve MP, JP Closterium cf. angustatum Ralfs MP C. baillyanum Bréb. * MP C. cf. calosporum Wittr. MP C. cynthia De Not. MP C. dianae Ralfs var. dianae JP C. dianae var. arcutum (Bréb.) Rabenh. MP C. dianae var. compressum G.A.Klebs * JP C. didymotocum Ralfs * JP C. directum W.Archer * MP C. gracile Ralfs var. gracile JP C. gracile var. striolatum Willi Krieg. * JP C. jenneri Ralfs MP C. leibleinii Ralfs var. minimum Schmidle * JP C. lineatum Ralfs JP C. navicula (Bréb.) L¨utkem. * M C. nematodes Joshua MP C. striolatum Rafts * JP Cosmarium angulosum Bréb. * LP C. askenasyi Schmidle MP C. binum Nordst. LP C. bioculatum Ralfs LP C. bireme Nordst. var. huzelii Kurt F¨orst. * LP C. connatum Bréb. MP, LP C. contractum Kirchn. var. ellipsoideum (Eflving) W. et G.S.West LP C. contractum var. incrassatum A.M.Scott et Prescott * LP C. cuneatum Joshua LP C. impressulum Elfving LP C. medioscrobiculatum W. et G.S.West var. egranulatum Gutw. LP C. obsoletum (Hantzsch) Reinsch var. sitvense Gutw. LP C. ocellatum B.Eichler et Gutw. LP C. pachydermum P.Lundell f. intermedium Maskell * LP C. portianum W.Archer f. extensum Prescott LP C. pseudamoenum Wille LP C. quadratulum (F.Gay) De Toni LP C. quinarium P.Lundell LP C. regnesii Reinsch LP C. reniforme (Ralfs) W.Archer * LP C. thailandicum Hirano LP C. trilobulatum Reinsch LP C. sp.1 (MP702) MP C. sp.2 (LP30) LP Desmidium aptogonum Bréb. var. tetragonum W. et G.S.West JP D. baileyi (Ralfs) Nordst. var. undulatum (Maskell) Nordst. LP D. coarctatum Nordst. JP D. occidentale W. et G.S.West MP D. quadratum Nordst. JP Euastrum ansatum Ralfs var. pyxidatum Delponte * MP E. cf. intermedium Cleve MP E. gnatophorum W. et G.S.West * MP E. insulare (Wittr.) J.Roy MP E. longicolle Nordst. * MP E. moebii (Borge) W. et G.S.West var. burmense W. et G.S.West MP E. praemorsum (Nordst.) Schmidle JP E. sinuosum W.Archer var. germanicum (Racib.) Willi Krieg. * JP E. spinulosum Delponte var. inermius Nordst. LP E. turneri W. et G.S.West * JP E. sp.1 (JP311) JP Gonatozygon aculeatum Hastings JP Groenbladia undulate (Nordst.) Kurt F¨orst. JP 904 N. Ngearnpat et al.

Table 2. (continued).

Hyalotheca dissiliens Ralfs var. tatrica Racib. * LP H. mucosa Ralfs LP Micrasterias alata G.C.Wall. LP M. ceylanica F.E.Fritsch MP M. foliacea Ralfs LP, MP M. radians W.B.Turner var. siamensis Hirano JP M. rotata Ralfs JP M. subdenticulata (Nordst.) Willi Krieg. var. ornata Willi Krieg. JP M. suboblonga Nordst. var. tecta Willi Krieg. JP M. tetraptera W. et G.S.West var. siamensis var. nov. * JP Penium cylindrus Bréb. * MP P. margaritaceum Bréb. * MP Phymatodocis irregulare Schmidle * JP Pleurotaenium ehrenbergii (Bréb.) De Bary LP P. kayei (W.Archer) Rabenh. MP P. subcoronulatum (W.B.Turner) W. et G.S.West JP P. trabecula N¨ageli var. trabecula LP, MP P. trabecula N¨ageli var. thailandicum Hirano JP P. truncatum (Bréb.) N¨ageli JP P. verrucosum (Bailey) P.Lundell MP Spondylosium planum (Wolle) W. et G.S.West LP Staurastrum avicula Bréb. * JP S. disputatum West et G.S.West LP S. ensiferum W.B.Turner LP S. excavatum W. & G.S.West LP S. furcatum (Ralfs) Bréb. LP S. inflexum Bréb. LP S. longibrachiatum (Borge) Gutw. LP S. sexangulare (Bulnh.) P.Lundell var. sexangulare LP S. sexangulare P.Lundell var. hasperum Playfair LP S. sp. 1 (LP03) LP S. sp. 2 (LP42) LP S. sp. 3 (LP44) LP S. sp. 4 (JP309) JP Staurodesmus phimus (W.B.Turner) Thomasson * LP Teilingia excavata (Ralfs) Bourr. LP Tetmemorus laevis Ralfs MP Triploceras gracile Bailey LP Xanthidium hastiferum W.B.Turner var. javanicum (Nordst.) Turner LP X. spinosum (Joshua) W. et G.S.West * LP

Fig. 2. Diagram showing the genera contribution (in number of different species) to the desmid flora of the three peat swamps. Desmids of Thai peat swamps 905

Figs 3–6. Four rare desmid taxa, endemic to the Indo-Malaysian biogeographical region, encountered in Jud peat swamp. 3 – Euastrum gnatophorum;4–Micrasterias subdenticulata var. ornate;5–M. suboblonga var. tecta;6–M. tetraptera var. siamensis;scale10µm. by small cell dimensions or by cell processes increasing to that, any intramarginal spines are lacking. the cell surface to content ratio. Holotypus: Preserved material has been deposited in Maikhao swamp and Jud swamp, where dimen- the Culture Collection of Algae of Applied Algal Re- sions of the sampled water bodies were smaller, have serched Laboratory (AARL), Faculty of Science ,Chi- a lower desmid species diversity marked by a relatively ang Mai University, Thailand as specimen No. PRE high portion of large-sized Closterium, Euastrum, Mi- AARL-G001 crasterias and Pleurotaenium species (Fig. 2). Remark- Type locality: M. tetraptera var. siamensis was col- ably, Jud swamp, with the lowest diversity of the three lected in Jud peat swamp, a mesotrophic, slightly acidic swamps, appeared to house the highest number of rare, (pH 6.1) fen, where it occurred in a shallow drainage- peculiar species. Taxa such as Euastrum gnatophorum, canal with floating mats of bladderworts and filamen- Micrasterias subdenticulata var. ornata and Micraste- tous algae. rias. suboblonga var. tecta. (Figs 3–5) are only rarely Collector: Neti Ngearnpat. reported and can be considered endemics of the Indo- Date of collecting: January, 6, 2006 Malaysian biogeographical region (e.g. Krieger 1937, 1939). Of Micrasterias tetraptera W. et G.S. West a deviating form was encountered, described as var. sia- Acknowledgements mensis var. nov., see below (Fig. 6). Their exclusive occurrence in Jud peat swamp might be linked up with The authors like to thank the members of the Applied Algal Research Laboratory in Chiang Mai for their kind help in the low conductivity of the habitat (as compared to collecting samples. Many thanks are also due to the Faculty Laempagarung and Maikhao swamps, Table 1) for it of Science and the Graduate School of Chiang Mai Univer- is well known that conductivity is one of the most sity for supporting this research. The research project was important parameters determining the distribution of funded by the Thailand Research Fund by way of the Royal desmid species (Coesel 1975, 1982). Obviously, the high Golden Jubilee Ph.D Program (Grant No. PHD/ 0186/ concentration of orthophosphate in this habitat (Ta- 2546). ble 1) which, in general, is thought to be unfavourable to oligo-mesotrophic desmid species (Spijkerman & Co- esel 1998 ) does not hinder the occurrence of these rare References desmid taxa (presumably because there is another fac- Coesel P.F.M. 1975. The relevance of desmids in the biological tor limiting the primary production, therefore the com- typology and evaluation of fresh waters. Hydrobiol. Bull. 9: petition by other algal species). 93–101. Coesel P.F.M. 1981. Classification of desmid assemblies in a Description of new taxon Dutch broads area. Arch. Hydrobiol. 91: 56–81. Coesel P.F.M. 1982. Structural characteristics and adaptations of desmid communities. J. Ecol. 70: 163–177. Micrasterias tetraptera W. et G.S. West var. siamensis Coesel P.F.M. 2000. Desmids (, Desmidiaceae) from Ngearnpat et Peerapornpisal var. nov. Thale Noi (Thailand). Nord. J. Bot. 20: 369–383. Descriptio: A varietate nominata differt lobis lateralibus Hirano M. 1967. Freshwater algae collected by the Joint Thai- incisuris ordine secundo tantum (in semicellulae parte Japanese Biological Expedition to Southeast Asia 1961–1962, pp. 1–71. In: Kira T. & Iwata K. (eds), Nature and Life in inferiore) vel tertio (in semicellulae parte superiore); Southeast Asia. Flora and Fauna Research Society, Kyoto. spinis intramarginalibus absentibus. Cellularum longi- Hirano M. 1975. Phytoplankton from Lake Boraphet in the Cen- tudo 190–195 µm, latitudo 155–163 µm. Typus: figura tral Plain of Thailand. Contr. Biol. Lab. Kyoto Univ. 24: nostra 6. Differs from the nominate variety in that the 187–203. lateral lobes in the lower part of the semicell are only Hirano M. 1992. Desmids from Thailand and Malaysia. Contr. Biol. Lab. Kyoto Univ. 28: 1–98. differentiated to the second order (instead of the third Kanetsuna Y. 2002. New and interesting desmids (Zygnematales, order), those in the upper part of the semicell only to Chlorophyceae) collected from Asia. Phycol. Res. 50: 101– the third order (instead of the fourth order). In addition 113. 906 N. Ngearnpat et al.

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