Periphyton and Sphagnicolous Protists of Dystrophic Bog Lakes (Brandenburg, Germany) II
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Limnologica 29 (1999)407424 LIMNOLOGICA http://www.urbanfiscber.de/j ournals/limno © by Urban & Fischer Verlag 1) Institut ftir Zoologie der Freien Universitiit Berlin, AG Protozoologie, Berlin, Germany; 2) Institut ftir 0kologie der Friedrich-Schiller-Universit~it Jena, AG Limnologie, Jena, Germany Periphyton and Sphagnicolous Protists of Dystrophic Bog Lakes (Brandenburg, Germany) II. Characteristic Species and Trophy of the Lakes MICHAELA C. STRUDER-KYPKE 1) • WILFRIED SCHONBORN 2) With 8 Figures and one Table Key words: Bog lakes; artificial substrates; ciliates; flagellates; testate amoebae; eutrophication Abstract Glass-slides were exposed in tour different bog lakes located in great Weichselian stage of the last glaciation and deposited in the forest areas in Northern Germany, about 100 km north of Berlin, in glacial drift or outwash plain. Those ice blocks often took order to examine the protist community of these dystrophic, acid several hundred years to melt completely, finally resulting lakes. Species composition and succession of the periphyton were in the formation of kettle lakes (Fig. 1). About 11,000 years studied during one year. The four investigated bog lakes exhibit simi- ago, the paludification began and mesotrophic-subneutral lar species compositions, most of the species were present in each of the four lakes. The periphyton was composed of a great variety of pro- circumstances prevailed. This process stopped approximate- tist species. Specimens of all major flagellated protists and of most of ly 9,000 years ago. The ice blocks were melted, the climate the ciliate taxa were found on the slides. Many of these species are changed and it became dry and warmer. Approximately ubiquitous. Some species, however, like the flagellates Spongomonas 4,000-2,000 years ago, a secondary turf layer started to sacculus, Rhipidodendron huxIeyi, Cyathobodo sp., and Pseudoden- grow on these paludificated lakes. Water was supplied only dromonas vlkii as well as the ciliates Ophrydium hyaIinum, Lep- by precipitation and the run-off from surrounding land topharynx costatus, Uroleptus caudatus, and Platyophrya sphagni are (CASPER & SCHEFFLER 1990; SUCCOW 1987; SUCCOW & found more frequently in acid waters. A regular and common occur- JESCHKE 1990). rence of these species partly characterizes the acid Sphagnum bogs. The larger bogs possess a residual lake which shows al- Besides that, common species like Bodo spp., Cryptomonas spp., Go- most no further terrestrialization, but reveals the excess niomonas truncata, Chilodonella uncinata, Vorticella sp., and Cyr- tolophosis mucicola are represented in the periphyton. Examinations water of the bog. Its edges are surrounded by Sphagnum of the testate amoebae in the Sphagnum mats show a very different mats that regularly tear off. In the center, kettle hole bogs are species composition compared to earlier studies. oligotrophic. The bog is surrounded by laggs (small water- The investigated bog lakes revealed mesotrophic conditions. In filled ditches) which collect the run-off water and are more the upper regions of the Sphagnum mats and in the center of the larg- mesotrophic (StJccow & JESCHKE 1990). er bog lakes, oligotrophic conditions still prevail. The lower regions With exception of the testate amoebae, protists of bogs of the Sphagnum mat and the smaller lakes, however, are influenced have always been neglegted in studies dealing with this habi- by the partly degradation of plants and the run-off water from sur- tat (e.g. HARNISCH 1925, 1929). Only few investigations rounding land. In comparison to earlier studies, the species composi- were carried out concerning the protist community of acid tion indicates a beginning eutrophication of the bog lakes. A direct anthropogenic impact can be excluded. bog lakes. The flagellates of bogs have not been examined or presented in detail up to now. SKUJA'S works (1948, 1956, 1964) described many planktonic flagellates, but from acid Introduction lakes. The ciliates of acid bogs were comprehensively stud- ied and many new species were described by GROLII~RE The bog lakes were formed by the melting of ice blocks (1975a, 1975b, 1977a, 1977b, 1978) and WENZEL (1953). which were broken from the withdrawing glaciers of the Besides that, only older literature exists about this habitat 0075-9511/99/29/04-407 $ 12.00/0 407 ~ 15,000 b. Chr. Deposited ice blocks in the glacial drift ~ 11,000 b. Chr. (late glacial) Melting of the ice blocks and formation of kettle lakes, begin of the paludification 8,000 b. Chr. (post glacial) End of the paludification i- and temporary stagnation in the peat formation Fig. 1. Development of the bog lakes (after JESCHKE 1987). (e.g. LEVANDER 1900; MERMOD 1914; SCHLENKER 1908; Material and Methods THIEBAUD &; FAVRE 1906). Testate amoebae are named tyrphobionts or sphagno- Sampling bionts and are considered as characteristic organisms of The sampling of the periphyton with glass slides as artificial sub- bogs. Therefore, extensive studies exist concerning their dis- strates according to WmBERT(1969) was performed as described in tribution, zoning, ecology, and systematics (e.g. HARNISCH STRI~DER-KYPKE(1999, this volume). Also the live counting of the 1925, 1927, 1929; MEISTERFELD 1977, 1979a, 1979b; protists is decribed there in detail. SCHONBORN 1962a, 1962b, 1963, 1990). Each region of the The sampling and counting of the testate amoebae of the Sphag- Sphagnum mats comprises its own community of testate num mats were done as described by SCn0NBORN(1963). To differ- amoebae. The horizontal distribution of these protists de- entiate the communities of the upper and lower horizons, the Sphag- pends on the degree of humidity; the vertical zoning is de- num cushion was cut - starting from the surface - into short pieces pendent on the light impact and the trophy (SCHONBORN of approximately 1 cm length and examined separately. 1962b, 1963). Characteristic genera of testate amoebae with- in the wet Sphagnum are: Hyalosphenia, Difflugia, Arcella, Light microscopy Centropyxis, Nebela, Euglypha, and Heleopera. The species Hyalosphenia papilio, Amphitrema flavum, A. stenostoma, Species determination was performed by means of an Axiophot mi- croscope with high magnification and - if necessary - with phase Heleopera sphagni, and Placocista spinosa harbour zoo- contrast or differential interference contrast. Additionally, protargol chlorellae and are very sensitive indicators for oligotrophic impregnation of whole slides was carried out in order to confirm conditions (ScHONBORN 1963). Reasons why the group of species determinations of the ciliates. testate amoebae in general are typical bog inhabitors, are the Species determination was made mainly with the help of the fol- relative low competitive pressure with other protists and the lowing literature: oligotrophic conditions in bogs (GOTTLICH 1990; SCHON- Flagellates: ETTL (1983), ETTL & GARTNER (1988), HUBER- BORN 1962a, 1962b). PESTALOZZI (1941, 1955, 1968), PATTERSON & LARSEN (1991), Our study deals with the species composition of the bog POPOVSKY & PFIESTER(1990), SKUJA(1948, 1956, 1964), and STAR- lakes, with main emphasis on the periphyton in the Sphag- MACH (1985). Ciliates: FOISSNERet al. (1991, 1992, 1994, 1995), KAnL(1930, num mat. Indicator species of acid, dystrophic lakes are fig- 1931, 1932, 1935). ured out and the environmental conditions (especially the trophy) of the lakes are compared to earlier studies (SCHON- BORN et al. 1965) and discussed. Protargol impregnation of whole slides The investigated bog lakes Himmelreichsee, Teufels-See, The protargol staining of the exposed slides followed the procedure Groger Barschsee and Kleiner Barschsee were described in described by WILBERT (1976). The cells were fixed in vapour of a detail in STRODER-KYPKE (1999, this volume), showing the 2% aqueous solution of Osmium tetroxide for 1-2 minutes. Subse- seasonal changes in the periphyton community and the peri- quently the liquid remaining on the slides was mixed with a drop of phyton structures of different habitats within the lakes. egg-albumen and the slides were allowed to dry at room temperature 408 Limnologica 29 (1999) 4 for 24 hours. The periphyton was post-fixed in Bouin's solution for sulfate solution for 1-2 minutes. After a last washing step in running 25 minutes and then transferred to 70% ethanol for 20 minutes, in tap water for 5 minutes, the slides were dehydrated in ethanol series order to remove the fixative. At this stage, the staining can be inter- (30, 50, 70, 95, 100%) with Xylene as final medium (5 minutes each) rupted, e. g. to collect the slides of several sampling dates. The slides and mounted with Entellan (Fa. Merck, Darmstadt). were washed for 5 minutes in running tap water. Subsequently, oxi- dation in a 5% Potassiumpermanganate solution for 1-2 seconds, fol- lowed by bleaching with 5% oxalic acid, also for 1-2 seconds, took place. Both steps were followed by a 10 minute washing step in run- Results and Discussion ning tap water. Then the slides were rinsed in Aqua dest. and placed in a 0.5% Protargol solution (= Albumosesilver of the Fa. Fluka, Species list of the flagellates and ciliates in Buchs/Switzerland) for 45 minutes at 45-50 °C. After this impregna- the four bog lakes tion, the slides were rinsed in Aqua dest. and transferred into a 1% Hydroquinone solution for 5 minutes. The slides were washed for 10 In Table 1 all species of flagellates and ciliates are listed minutes in running tap water and finally fixed in a 5% Sodiumthio- which were found in the lakes during the investigations from Table 1. Species list of the flagellates, ciliates and rhizopods of the periphyton and Sphagnum of the bog lakes in alphabetical order. Abbreviations: Bio - Biocoenosis; H/A - Lake Himmelreichsee, site A; H/B - Lake Himmelreichsee, site B; T - Lake Teufels-See; G/A - Lake Groger Barschsee, site A; G/B - Lake GroBer Barschsee, site B; K - Lake Kleiner Barschsee; uh - upper horizon; lh - lower horizon; As - Aufwuchs, sessile; Av - Aufwuchs, vagile; P - Plankton; o - single; + - rare (testate amoebae: present); ++ - regularly observed (testate amoebae: dominant); +++ - nearly always present.