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Lake Chad: Ecology and Productivity of a Shallow Tropical Ecosystem PDF

583 Pages·1983·15.325 MB·English
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LAKE CHAD MONOGRAPHIAE BIOLOGICAE VOLUME 53 Series editor H. J. DUMONT 1983 Dr W. JUNK PUBLISHERS a member of the KLUWER ACADEMIC PUBLISHERS GROUP THE HAGUE / BOSTON / LANCASTER LAKECHAD Ecology and Productivity of a Shallow Tropical Ecosystem Edited by J .-P. CARMOUZE, J .-R. DURAND and C. LEVEQUE 1983 Dr W. JUNK PUBLISHERS a member of the KLUWER ACADEMIC PUBLISHERS GROUP THE HAGUE / BOSTON / LANCASTER Distributors jor the United States and Canada: Kluwer Boston, Ine., 190 Old Derby Street, Hingham, MA 02043, USA jorall othercountries: Kluwer Aeademic Publishers Group, Distribution Center, P .O.Box 322, 3300 AH Dordreeht, The Netherlands Library of Congress Cataloging in Publication Data Hain entry under title: Lake Chad. (Monographiae biologicae ; v. 53) Includes bibliographies and index. 1. Lake ecology--Chad. Lake. 2. Biological productivity--Chad. Lake. 3. Chad. Lake. I. Carmouze, Jean-Pierre. H. Durand, Jean Rene. IH. LW@que, C. IV. Series. QP1.P37 vol.53 5748 [574.5'26322'096743] 83-4288 [QH195.C46] ISBN-13: 978-94-009-7268-1 e-ISBN-13: 978-94-009-7266-7 DOI: 10.1007/978-94-009-7266-7 Cover design: Max Velthuijs Copyright © 1983 by Dr W. Junk Publishers, The Hague. Softcover reprint ofthe hardcover 1st edition 1983 All rights reserved. No part of this publieation may be reprodueed, stored in a retrieval system, or transmitted in any form or by any means, meehanical, photoeopying, reeording, or otherwise, without the prior written permission of the publishers, Dr W. Junk Publishers, P.O. Box 13713, 2501 ES The Hague, The Netherlands. Contents Preface (J. Daget) VII Foreword (The Editors) XI Historical background (J.-P. Carmouze) I. The lacustrine environment and its evolution 1. Paleolimnology of an upper quaternary endorheic lake in Chad Basin (M. Servant and S. Servant) 11 2. The lacustrine environment (J.-P. Carmouze and J. Lemoalle) 27 3. Physical and chemical characteristics of the waters (J.-P. Carmouze, J.M. Chantraine and J. Lemoalle) 65 4. Hydrochemical regulation of the lake (J.-P. Carmouze) 95 11. The main types of communities and their evolution during a drought period 5. The aquatic vegetation of Lake Chad (A. Iltis and J. Lemoalle) 125 6. The phytoplankton (P. Compere and A. Iltis) 145 7. The zooplankton (L. Saint-Jean) 199 8. The benthic fauna: ecology, biomass and communities (C. Leveque, C. Dejoux and 1. Lauzanne) 233 9. The fauna associated with the aquatic vegetation (C. Dejoux) 273 10. Fish communities of Lake Chad and associated rivers and flood plains (V. Benech, J.-R. Durand and J. Quensiere) 293 111. The balance of a lacustrine ecosystem during 'Normal Chad' and aperiod of drought 11. Phytoplankton production (J. Lemoalle) 357 12. Secondary production (zooplankton and benthos) (C. Leveque and 1. Saint-Jean) 385 13. The exploitation of fish stocks in the Lake Chad region (J.-R. Durand) 425 IV. Trophic relations 14. Trophic celations between the phytoplankton and the zooplankton (A. Iltis and 1. Saint-Jean) 483 15. Trophic relations of fishes in Lake Chad (1. Lauzanne) 489 16. The impact of birds on the lacustrine ecosystem (C. Dejoux) 519 V 17. The lacustrine ecosystem during the 'Normal Chad' period and the drying phase (J.-P. Carmouze, J.-R. Durand and C. Leveque) 527 Systematic index 561 General index 571 VI Preface In 1963 the director general of the 'Office de la Recherche Scientifique et Technique Outre-Mer' (ORSTOM) asked me to form a well structured team of hydrobiologists based at Fort-Lamy (now N'Djamena) where a laboratory could be built as a branch of the already existing ORSTOM Center. The International Biological Program, IBP, had at this time recommended an integrated study of ecosystems selected in proportion to their representation of a particular zone or ecological conditions. I took this opportunity to propose Lake Chad as the model of a tropicallake in a semi-arid c1imate as part of the activity of the section Productivity of freshwater communities (PF). With the efficient help ofDr B. Dussart a research program was established involving the establishment of a permanent team of ten researchers in Chad. Apart from myself, ORSTOM only had three hydrobiologists: Mr R. Gras, Mr G. Loubens and Mr A. Iltis. These valuable researchers who had been working for the 'Centre Technique et Forestier Tropical' and had acquired considerable experience in African freshwaters, unhesitatingly accepted to join me at Fort-Lamy. It was necessary to enlarge this team with so me young and enthusiastic collaborators, namely, Mr C. Leveque, Mr C. Dejoux, Miss S. Duwat, Mr L. Lauzanne and Mr. J. R. Durand, later joined by Mr J. P. Carmouze and Mr J. Lemoalle. At first we stayed briefly at the laboratory of the 'Institut de Medecine Veterinaire et Tropicale' (IMVT) at Farcha until the construction of the hydrobiology laboratory at Fort-Lamy was completed. It is with pleasure that I pay respect not only to the scientific value of this first team which established itself by producing seven Ph.D's but especially to the good atmosphere and friendship established between the researchers and technicians at Fort-Lamy as well as in the field. Due to this exceptional atmosphere, there were only scientific problems to solve and the two years that I spent at Chad have left me with unforgettable memories. With the installation of the team in its laboratory, opened in 1965 and fairly weIl equipped, the research program was well on its way and I returned to Paris. Here, with the help of Dr B. Dussart and until1975, I supervised the scientific Photo 1 The ORSTOM Hydrobiological team in 1966. coordination of research and the recruitment of new scientists and technicians to ensure the maintenance or reinforcements of the first team. This experience proved to be extremely beneficial for the training of men who have become highly qualified specialists as weil as for the importance of their scientific results. The director general of ORSTOM took a certain risk by regrouping all his hydrobiologists at the same time on the same program. What appeared at first to be achallenge soon became a total success as the reader will find out for himse\f when reading this book. It should, however, be recognized that the interest and the scientific scope of the study presented here partly depends on a happy conjuncture: the presence of a team of scientists having had the time to study the state of the lake called 'Normal Chad' before and during the time when years of drought led to the state of 'Lesser Chad'. This indeed appeared to be a cyclic phenomenon that had already occurred at the beginning of the century as shown by Tilho's observations, and which would recur again after a new period of high waters. However, a researcher would only have the chance to observe this once during his career. This complete team of hydrobiologists then, was able to follow in detail the lowering of the lake level. If this event had occurred ten years earlier, there VIII would have been no one to study it; if it had occurred ten years later, it would have passed unobserved as research in Chad would have been impossible. When I first became interested in Lake Chad, I was struck by two factors in particular. In spite of the approximately 20000 km2 of area then known, Hutchinson had not considered it worthy to be included in the list of African lakes considered in his classical treatise of Limnology. Furthermore as far as the fish fauna was known, there were no true lacustrine species present. Alestes dageti, described as endemic to the lake by Blache and Miton, only appeared to me as an ecophenotypic form of Alestes nurse characterized by its dwarfism, a poorly known deterministic phenomenon, although it is relatively frequent in some fish populations. In 1967, I wrote that 'Ecologically Lake Chad is closer to the flood zones of large tropical rivers than true lakes .,. it thus appears probable that a slight worsening of climate could involve a complete drying of the lake'. The exceptional drought of the years 1972-73 which was dramatic for the Sahelian populations showed that it did not behave as a simple system, as drought first affected the north, i.e. the deepest part. It will be interesting to read how the major types of population changed over the course of the drying phase. Finally, it is necessary to recall that the fisheries in Lake Chad were developing around the sixties in relation to the introduction of nylon gill nets and especially the development of transport facilities to carry the dried and smoked fish towards Nigeria and the southern territories. Now, paradoxically, the driest years turned the production rocketing to 220000 tons in 1974, causing a huge concentration of fishermen in the lake. In addition the productivity of these tropical waters and the resilience of their species was seen to be much higher than provisionally estimated by comparison with other environments. Following several notes and papers published in specialized journals, the summary of 15 years of research presented here in an easily accessible form appeared welcome to me. It is not premature as the field-work had to be interrupted in 1979 nor is it too late because the problems of management and the improvement of the Chad basin are not yet solved. I agreed and accepted the invitation to preface this scientific work with real pleasure for it is a factual document. I would like to address my personal thanks to the authors for their collaboration as weIl as to the editors who agreed to publish it and distribute it to the international scientific community. I am sure that it will obtain from the latter all the success that it deserves. Jacques Daget Professor, National Museum of Natural History, Paris IX Foreword Lake Chad is a large, endorheic, shallow basin south of the Sahel zone. Although it is subject to important annual variations of area and volume, it is one of the largest lakes of the African continent, following those in the Rift valley (Victoria, Tanganiyka and Malawi). The biotopes of Lake Chad are similar to those found in lakes in the central Niger delta, southern Sudan, Lake Chilwa, Okavango delta and a number of other less important aquatic ecosystems. They are characterized primarily by the instability of hydrological conditions and the abundance of macrophytes (Cyperus, Typha and Phragmites, in particular). Thus the interest of studies conducted on Lake Chad extends beyond regional limits because many problems are common to all these regions even if each has its own specificity. Since the beginning, the research pro gram on Lake Chad was part of the French contribution to the International Biological Program, entitled 'Study of a Tropical Lake in a Semi-Arid Climate'. Its principal objective was to describe the lake hydroclimate and evaluate the biomass and production of the main groups of organisms in order to establish a balance of matter and energy and of their cycles on the different trophic levels. In addition to the fundamental value of this research for biological phenomena in a climatic zone that was fairly unknown until now, this study could also have economic consequences for the development projects for Chad basin, and particularly for the fisheries. Indeed Lake Chad is a very productive environment due to its morphology. Its average fisheries catch, estimated at more than 100 000 tons per year, provides an important economic activity for the nearby communities. Between 1964 and 1978, a team of several limnologists worked at the ORSTOM Center at N'Djamena, where a modern laboratory of 220 m2 was built in 1965. The first studies were limited to the east zone of the lake where it was possible to make preliminary tests for sampling methods and analysis with the Nausicaa, a 10 m long boat specially adapted to these shallow environ ments. It was soon apparent that a general view of the lake was essential for the proper interpretation of several observed phenomena. Thus, a study of large

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