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Brackish-water phytoplankton of the Flemish lowland Developments in Hydrobiology 18 Series editor H.J. Dumont H.J. Dumont and J. Green, eds., Rotatoria. Lake Vechten and Tjeukemeer, The Nether ISBN 90 6193 754 x. 1980. lands. ISBN 90 6193 762 O. 1982. J. Barica and L.R. Mur, eds., Hypertrophic F. Schiemer, ed., Limnology of Parakrama Ecosystems. ISBN 90 6193 752 3. 1980. Samudra - Sri Lanka. A case study of an M. Dokulil, H. Metz and O. Jewsen, eds., ancient man-made lake in the tropics. Shallow Lakes. Contributions to their lim ISBN 90 6193763 9. 1983. o nology. ISBN 90 6193 753 1. 1980. Curt Forsberg and Jan-Ake Johansson, eds., Michael Goulding, Man and Fisheries on an Forest Water Ecosystems. Amazon Frontier. ISBN 90 6193 755 8. 1981. ISBN 90 6193 764 7. 1983. W.O. Williams, eds., Salt Lakes. B. Pejler, R. Starkweather and Th. Nogrady, ISBN 90 61937566. 1981. eds., Biology of RoUfers. Ernest R. Schockaert and Ian R. Ball, eds., ISBN 90 6193 765 5. 1983. The Biology of the Turbellaria. J. Merilainen, P. HuUunen and ISBN 9061937574.1981. R.W. Battarbee, Paleolimnology. V.K.E. IImavirta, R.I. Jones and E.P.E. ISBN 90 6193 766 3. 1983. Persson, eds., Lakes and Water Manage U.T. Hammen, ed., Saline Lakes. ment. ISBN 90 6193 758 2. ISBN 90 6193 767 1. 1983. O.M. Harper and J.A. Bullock, eds., Rutland R.G. Wetzel, ed., Periphyton of Freshwater Water-Decade of Change. Ecosystems. ISBN 90 6193 768 x. 1983. ISBN 90 6193 759 O. 1982. A.G. Caljon, ed., Brackish-water Phytoplankton Peter G. Sly, ed., Sediment/Freshwater Interac of the Flemish Lowland. ISBN 90 6193 769 8. tion. ISBN 90 6193 760 4. 1982. 1983. Mark D. Morgan, ed., Ecology of Mysidacea. Anton McLachlan and Theuns Erasmus, eds., ISBN 90 6193 761 2. 1982. Sandy Beaches as Ecosystems. R.D. Gulati and S. Parma, eds., Studies on ISBN 90 6193 770 1. 1983. Brackish-water phytoplankton of the Flemish lowland by A. Caljon 1983 DR W. JUNK PUBLISHERS ~. a member of the KLUWER ACADEMIC PUBLISHERS GROUP ,- THE HAGUE / BOSTON / LANCASTER ~ Distributors jar the United States and Canada: Kluwer Boston, Inc., 190 Old Derby Street, Hingham, MA 02043, USA jar all other countries: Kluwer Academic Publishers Group, Distribution Center, P.O.Box 322, 3300 AH Dordrecht, The Netherlands Library of Congress Cataloging in Publication Data Ce.J.Jon, A. Brackish-water phytoplankton o~ the Flemish lowland. (Developments in hydrobiology ; 18) Bibliography: p. Includes index. 1. Algae--Belgium--East Flanders. 2. Phytoplankton- Belgium--East Flanders. 3. Brackish water ~lora-­ Be~~~--East Flanders. 4. Coastal flora--Belgium- East 'Flanders. I. Title. II. Series. QK574.B4C34 1983 589.39493'1 83-13579 ISBN-13: 978-94-009-6556-0 e-ISBN-13: 978-94-009-6554-6 DOl: 10.1007/978-94-009-6554-6 ISBN 90-6193-751-5 (series) Copyright © 1983 by Dr W. Junk Publishers, The Hague All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publishers, Dr W. Junk Publishers, P.O. Box 13713,2501 ES The Hague, The Netherlands Preface Numerous papers have been published on the animal and algal communities of fresh-water and marine biotopes, and their relation to the physical and chemical factors of their environment. Brackish-water biotopes have been studied much less intensely, presumably due to their limited distribution and to the complexity of their bioco enoses, composed of fresh-water, brackish-water and marine organisms. The reduced size of the species present forms a supplementary difficulty, which is especially pronounced in the algal communities, a major part of which are composed of nanno-phytoplankters. The University of Ghent considered a detailed survey sufficiently attractive to set up because of the high species diversity, the gaps in the knowledge of brackish-water nannophytoplankton at the species and population levels, and the convenient proximity of brackish-water biotopes. Moreover, the brackish-water biotopes proved to be unique, and consist of a series of landlocked creeks, con nected to the sea by means of an underground salt water supply. VII Contents Chapter 1 Introduction..................... . ................ . ............... . . 1 Chapter 2 Study area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2.1. Topographical situation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2.2. Historical aspects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2.3. Edaphic factors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2.4. Hydrology. ................. . .................................. 5 2.4.1. Salinity fluctuations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2.4.2. Origin of the brownish colour of certain creeks. . . . . . . . . . . . . . . . . . . . . . . . 6 2.4.3. Hydrography.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2.5. Biotopes................. . ................... . .......... . . .... 7 2.6. Climatic features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10 Chapter 3 Methods............................................. . ...... . .. . . . 11 3.1. Sampling procedure. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11 3.2. Physical properties, water chemistry, chlorophyll a and phaeophytin a . . . . . . . . . . . .. 11 3.3. Phytoplankton samples. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12 3.4. Periphyton and benthos samples. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Chapter 4 Data analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 4.1. Classification techniques .......................... . .. . ...... ... ... " 15 4.2. Ordination techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 4.2.1. Principal component analysis (P.C.A.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 4.2.2. Detrended correspondence analysis (D.C.A.). . . . . . . . . . . . . . . . . . . . . . . . .. 17 4.3. Transformations....................... . . . ......... . . . ..... .. .... 17 4.4. Selection of the number of the variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17 Chapter 5 Physical properties, water chemistry and phytoplankton production. . . . . . . . . . . . . . . . . .. 19 5.1. Temperature.......... . ....... . ................. . .... . ......... . 20 5.2. Conductivity................................................... 20 5.3. Chlorides...................................................... 23 5.4. pH.................................................. . ....... 24 5.5. Carbonate, bicarbonate and alkalinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25 5.6. Calcium, magnesium, total hardness. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 5.7. Ammonia.............................................. . .. . ... . 32 5.8. Nitrite.............. . . . ............................. . ....... . . 34 5.9. Nitrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34 5.10. Orthophosphate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 36 5.11. Dissolved oxygen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 36 5.12. Active chlorophyll a of the total phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . .. 38 5.13. Phaeophytin a of the total phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 41 5.14. Active chlorophyll a of the net phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 41 5.15. Phaeophytin a of the net phytoplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44 5.l6. Surface area of the total phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44 VIII Chapter 6 Phytoplankton groups. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45 6.1. Seasonal phytoplankton groups. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45 6.1.1. Seasonal phytoplankton groups in the Grote Geule (1). . . . . . . . . . . . . . . . . .. 45 6.1.2. Seasonal phytoplankton groups in the Rode Geule (2) . . . . . . . . . . . . . . . . . .. 45 6.1.3. Seasonal phytoplankton groups in the Kapellen-poldcrkreek (5). . . . . . . . . . . .. 48 6.104. Seasonal phytoplankton groups in the Boerenkreek (10). . . . . . . . . . . . . . . . .. 48 6.1.5. Conclusions.............................. . . . . . . . . . . . . . . . .. 49 6.2. Synopsis of the seasonal phytoplankton groups. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54 6.2.1. Chlamydomonas umbonata-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54 6.2.2. Monoraphidium contortum-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 58 6.2.3. Anabaena spiroides var. longicellularis-group. . . . . . . . . . . . . . . . . . . . . . . . .. 58 6.204. Hemiselmis simplex-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 59 6.2.5. Gymnodinium splendens-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60 6.2.6. Chaetoceras orientalis-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60 6.2.7. Prymnesium spec. I-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60 6.2.8. Comparison of the synoptic seasonal groups. . . . . . . . . . . . . . . . . . . . . . . . . . 61 6.2.9. Salinity preferences of the synoptic seasonal groups. . . . . . . . . . . . . . . . . . . .. 61 6.3. Spatial phytoplankton distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 62 6.3.1. Spatial phytoplankton groups in the sample set of January 1973 . . . . . . . . . . .. 62 6.3.2. Spatial phytoplankton groups in the sample set of November 1976. . . . . . . . . .. 67 6.4. Discussion.. . ........... . ...................................... 73 Chapter 7 Typification of algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77 7.1. Brackish-water species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77 7.1.1. Subdivisions of brackish water. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77 7.1.2. The origin of brackish water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 78 7.1.3. Ecological features of brackish-water species. . . . . . . . . . . . . . . . . . . . . . . . .. 78 7.104. Types of brackish-water species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79 7.104.1. Holeuryhaline species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79 7.104.2. Euryhaline limnobionts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79 7.104.3. Euryhaline marine species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80 7.1.404. Brackish-water species -Hyphalmyrobionts . . . . . . . . . . . . . . . . . .. 80 7.2. Trophic preferences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80 7.3. Saprobity preferences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80 7 A. pH preferences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81 7.5. Littoral species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81 7.6. Plankton species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81 7.7. Benthic species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81 7.8. Periphytic species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 82 7.9. Rheophiles..................................................... 82 7.10. Thermal preferences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 82 Chapter 8 Systematic account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83 Taxonomic index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 259 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 265 Chapter 1 Introduction From a systematic and ecological point of view, the and De Raeve (1975) on the vegetation; Dumont & algae of brackish waters are of great interest. The Gysels (1971), and Verstraeten (1978) on the fauna; algal community in these biotopes is composed of Van Meel (1960) on the algae;De Ridder{1956)onthe holeuryhaline, euryhaline marine and euryhaline rotifers and on some physical and chemical aspects. brackish-water organisms and euryhaline and weakly Most of these papers contain physical and chemical euryhaline limnobionts. The relative proportions of data, and these make it possible to study the these different components is closely related to evolution of several factors in time. We could also salinity. In Flanders, the algological richness of the compare our algological observations with those by brackish waters in the coastal region had been Van Meel (1960), although this author examined only documented by the work of Conrad & Kufferath net phytoplankton, while the main biomass and algal (1954) who studied some brackish waters in the diversity is due to the nanno phytoplankton. vicinity of the township of Lillo 0 , near Antwerp. Many studies on the taxonomy of brackish-water These interesting biotopes were later to be destroyed algae have been published, but till now no truly by the extension of the harbour of Antwerp, but a systematic survey is available. Papers such as Conrad comparable area which is at least as varied is still to & Kufferath (1954) give an adequate impression of be found in the north of East Flanders. It is composed the species diversity of certain brackish waters and of a series of landlocked creeks. describe interesting new taxa, but they offer few We selected this area to carry out an in-depth clues for the identification of algal taxa. Diatoms are algological study. It is a relic of the repeated trans the most intensely studied algal group. They were gressions and regressions of the North sea and of studied, among others, by Hustedt (1925, 1930 a, b, many dike-bursts which occurred during the last 1931, 1932, 1933 a, b, 1937, 1959, 1961, 1962, millennia. It forms a geographical unity with the 1964, 1966) and by Cho1noky (1968) in brackish creeks of Zealand Flanders (The Netherlands). Its water, limnetic and marine biotopes. Brackish-water creeks are landlocked and are connected to the sea by species are also dealt with in general works such as means of an underground supply of salt water only. "Rabenhorst's Kryptogamen Flora", but not all This supply tends to keep the salinity of most creeks algal groups are treated here: Flagellatae (Gemeinhardt at a reasonably constant level. Depending on the rate & Schiller 1930), Cyanophyta (Geitler 1932), of salt water supply, a gradient from nearly limnetic Dinoflagellatae (Schiller 1933, 1937), Conjugatae to strongly mesohaline waters exists. This gradient (Krieger 1937), Heterocontae (Pascher 1939), even leads to polyhaline waters in Zealand Flanders. Oedogoniales (Gemeinhardt 1939), Zygnemales In some creeks, however, a gradual increase in salinity (Kolkwitz & Krieger 1941) and the earlier cited has been observed during the last decennia. Many are Diatoms (Hustedt 1930 a, 1931, 1932, 1933 a, b, in the process of silting up, and some are so shallow 1937, 1959, 1961, 1962, 1964, 1966). For these that they desiccate or near-desiccate during dry groups in part, and for all algal groups not cited, summers. This phenomenon is responsible for the identification of brackish-water taxa requires a pronounced fluctuations in salinity found in some perusal of the specialized literature on this subject creeks. The whole range of adaptations to these (a.o. Butcher 1959, 1961, 1967; Caljon 1976, 1979, periodically and spatially changing osmotic con 1983; Carter 1937; Conrad 1926 a, b, 1938, 1939; ditions, make the flora and the fauna of this Conrad & Kufferath 1954; Dragesco 1965; Heynig landlocked creek system an extremely diverse one. 1967; Hulburt 1965; Koster 1966; Ruinen 1938; The complexity of its biocenoses is intriguing and has Schiller 1953, 1957; etc.). Because fresh-water, challenged many a liminologist. brackish-water and marine organisms co-occur in Previous investigations which have been carried brackishcwater, one should also consult the extensive out in this area are: Van Hecke & Rammeloo (1971) literature on the systematics of fresh-water and 2 marine algae. For this reason, we decided to classification techniques are nowadays widely applied accentuate the systematic part of this study and to in many scientific disciplines (psychology, sociology, provide an extensive iconographic part, thus hoping biological sciences, etc.). A host of papers deal with to facilitate the future systematic study of brackish the application of these methods in the treatment of water algae. Moreover, we add a short ecological phytoplankton data: Allen & Koonce (1973), Estrada typification to the description of each taxon. (1972), Symons (1970, 1973), Reyssac & Roux Once the data on algae and environmental factors (1972), Holland & Claflin (1973), Thurlow et aI. are collected, they need to be analyzed. We used two (1975), Bartell et al. (1978) and Caljon (1978 b). types of ordination methods (principal component Levandowsky (1972) first applied these methods to analysis and detrended correspondence analysis) and the phytoplankton of brackish-water biotopes, but one type of classification technique (group average without very positive conclusions, unlike the results cluster analysis) to identify objectively the different of the present study. algal communities in our samples. Ordination and 3 Chapter 2 Study area 2.1. Topographical situation - Klein Geulken (17) - Kapellenpolderkreek (5). The landlocked creek area is situated in the West of this first concentration of creeks we also northern part of East Flanders (Belgium), in a sampled the Noorddijkkreek (6) on the territory of territory which is limited in the North, the East and the municipality of Watervliet. the West by the Belgian-Dutch border and in the The second group of landlocked creeks is situated South by the limit Polders·Flemish sandy region, i.e. further west, on the territory of the municipalities of approximately the line Sint·Laureins-Boekhoute St.·Jan·in-Eremo, St.·Margriete and Waterland Assenede-Sas-van-Gent. Two concentrations of Oudeman, with Lambert·coordinates 61 (W), 97 (E), landlocked creeks occur in this area. The first group is 220 (N) and 215 (S) on survey-maps 13/3-4 and situated in its eastern part, on the territory of the 5/7-8 of Belgium (Fig. 2). We surveyed the follow municipalities of Assenede and Boekhoute, with ing creeks in this area: Lambert·coordinates 105 (W), 110 (E), 219 (N) and Molenkreek (7,8,9) 214 (S) on survey·maps 14/1-2 and 6/5-6 of Boerenkreek (10) Belgium (Fig. 1). We studied five creeks in this area: Roeselarekreek (II, 19) Grote Geule (1) Hollandersgat (12) Rode Geule (2, 3, 4) Blokkreek (13) Grote Kil (16) Vrouwkenshoekkreek (14) Fig. 1. Topographic map of the creek area near Assenede and Watervliet with locations of sampling stations 1-6, 16, 17.

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Numerous papers have been published on the animal and algal communities of fresh-water and marine biotopes, and their relation to the physical and chemical factors of their environment. Brackish-water biotopes have been studied much less intensely, presumably due to their limited distribution and to
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