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TROPICAL AFRICAN INLAND WATERS 3

© The Freshwater Biological Association 2006 Freshwater Forum 26 (2006), 3–37

A BRIEF HISTORY OF THE SCIENTIFIC STUDY OF TROPICAL AFRICAN INLAND WATERS

JACK TALLING

Dr J.F. Talling, Freshwater Biological Association, Far Sawrey,

Ambleside, Cumbria LA22 0LP, UK Introduction

The inland waters of Africa, and especially its large rivers, determined much of the original geographical exploration of the continent. Subsequently, their scientific study has contributed to the science of inland waters in general and of tropical inland waters in particular. The history of this tropical study is outlined below. It draws upon personal experience in several African universities and a research institute, vivid impressions of aquatic habitats and communities within African landscapes, and a desire to move from individual descriptions to wider comparisons and the underlying fundamentals.

In any history of scientific endeavour in Africa, the human factor deserves special attention. In colonial times much science, like trade, followed the flag. In Africa, unlike South America, this phase extended over – and benefited from – a period of rapid growth in freshwater science. Later, after the 1950s and widespread political independence, the science continued from both pre-established and new institutions – mainly universities and fishery institutes. These gave new opportunities to indigenous staff, and often hosted relatively brief expeditions from abroad. International and inter-governmental projects grew in importance. They were planned with regard to perceived needs, technical and socio-economic, of a region; but they could also enrich fundamental insights within the subject.

The wide coverage has necessarily involved the exclusion of detail on the contributions of individuals, although examples appear in the numerous references. In this I follow the pattern adopted in a history of British contributions to freshwater science244. The situation in 1900

By this date the disposition of the main rivers and lakes (Fig. 1) was reasonably well known. It was evident that most of the rivers had a strongly variable – indeed rhythmic – discharge, linked to the alternation of rainy and dry seasons. That of the Nile had been followed, by water level in Egypt, from ancient times; the crucial connection of its floodwater to

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Freshwater Forum 26 (2006)

FIG. 1. Map of Africa with named waterbodies (shallow lakes underlined)243. rains in Ethiopia was proposed in 1668 by the Jesuit Father Lobo236. Lakes (Fig. 2) were numerous, some very large and influenced by large-scale tectonic events like crustal warping (e.g. Victoria, Chad) and rifting (e.g. Tanganyika). Systems of both open and closed drainage were involved, with a corresponding wide range of water salinity. Only in a few cases were there analyses of the chemical composition, such as that obtained by Henry Wellcome – subsequently an influential figure in tropical medicine – for a salt lake in western Uganda207.

A beginning had been made towards assessment of aquatic faunas and floras. Most collections were sporadic, as of an early sample of algae from Lake Nyasa57. Near the end of the century there were two examples of more sustained enquiry. The aquatic fauna of Lake Tanganyika had attracted attention by the presence of endemics and the resemblance of some species – especially thick-shelled molluscs – to marine forms. This



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was taken by Moore as evidence of a direct marine origin for the lake, leading to his expeditions of 1895 and 1898 and subsequent book192 that suggested derivation from a Jurassic sea. The second example concerned the water-mass and plankton of Lake Nyasa (later Malawi), investigated during a German expedition of 1899–1900. Using a simple sampler, the expedition laboriously discovered the first thermocline to be described from a tropical lake82. Net-samples of plankton, gathered throughout 1899 and later examined by Schmidle, were used to construct a first rudimentary account of the seasonal changes of phytoplankton in a tropical lake234. The period 1900–1925

This period saw some notable acquisitions of information regarding aquatic environments and their biota, although integrated ecological science had not yet arrived. The vertical configurations (bathymetry) of various large lakes, including Victoria and Tanganyika (Fig. 1), were mapped and found to range widely. The work of Stappers in 1911–13 on L. Tanganyika237 disclosed a considerable volume of water at depths exceeding 1000 m. Vertical relationships were also important in the regulation of flow in rivers. Thus a ‘stepped’ longitudinal section of the Nile167 included Lake Plateau, Rift Valley, savanna, Sudd swamp and desert cataracts (examples in Fig. 3).

Various expeditions and their collections made somewhat disconnected contributions to aquatic science. Examples relating to water chemistry are cited by Talling & Talling246 and others to freshwater algae by Brook et al.30. Zoological work is exemplified by the plankton collections examined by Daday48 from Lake Victoria. There was the remarkable discovery of aquatic animals from within the Sahara212.

Three examples of more sustained and systematic study are of note. Shortly after 1900 the Wellcome Laboratories were set up at Khartoum in the Gordon Memorial College; from them Beam10 made chemical analyses over several years of water from the Blue and White Niles that demonstrated considerable seasonal and longitudinal changes. Collections of fishes from the Congo basin and the Nile were available in Europe and used by Boulenger to produce major monographs on the two fish faunas26,

27. Lastly, the controversial ‘Tanganyika Problem’ of Moore was tested further by the expedition of Cunnington (1904–05). From its collections and with the cooperation of many group-specialists, Cunnington assembled a ‘comparative limnology’47 and rejected the original hypothesis of a direct marine origin – already doubted from other work on the gastropod molluscs213.



FIG. 3. Dissimilar river sectors down the Nile: (top left) savanna, southern Sudan; (bottom left) desert cataract, Nubia; (above right) floodplain swamps (‘Sudd’).

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The beginnings of integrated ecology: the years 1925–1945

The following 20 years were a period of advancing colonial administration in many African territories, in which research was stimulated by economic and social issues. Two major areas were hydrology and fisheries. The former involved estimation of water budgets and the flow gauging of many rivers. One was the Nile, on which the original Aswan Dam (Fig. 4) was created, and for which Hurst and his associates in the Physical Department of the Ministry of Public Works at Cairo gathered information from sources to delta in a series of hydrological monographs The Nile Basin (e.g. 124). The broad picture for the Nile was later summarised in a more popular book by Hurst123, and for the hydrology of tropical Africa in general by Balek2.

Initiatives that concerned inland fisheries had a greater influence on the development of aquatic ecology. The most significant began with the fisheries survey of Lake Victoria92 by Graham and Worthington in 1927–28. This extended beyond fish faunistics, distribution and abundance to problems of changing thermal stratification267 and day-night (diel) vertical migration of zooplankton268. All these subjects were taken up further in work on the nearby Lake Albert. The investigation of diel change in Victoria was the first such work for tropical fresh waters.

In the years immediately following, there were two further British expeditions that closely interrelated biological and environmental features of East African lakes. That of Penelope Jenkin, in 1929, was a one-woman expedition that developed from friendship with a member of the Leakey family166. It took in Kenyan lake waters lacking surface outflows, with

FIG. 4. Impoundments of man-made lakes: (left) original Aswan Dam, photo. 1955; (above right) Kariba Dam, photo. 1983.



differences that were dominated by a wide range of titration alkalinity due to bicarbonate and carbonate130. The biota were often very dense, and involved a carbon flow that Jenkin investigated by experiments on photosynthesis in situ (adapted from the then recent work of Marshall and Orr in Scotland) and observations on the filter-feeding mechanism of flamingos131. A second and larger Cambridge expedition, in 1930–31, was led by Worthington – clearly enthused by his earlier African experience on lakes Victoria and Albert. It too dealt comparatively with a range of lakes, and with communities from phytoplankton to fishes. One participant, Beadle, provided a wide range of information on water chemistry and functional relationships which included photosynthesis-depth distribution6 and ecology of swamp invertebrates5. Besides zooplankton studies271, Worthington took up the comparative faunistics and biology of the fish communities, and subsequently suggested interpretations of their evolutionary differentiation in Africa269 – a topic that later attracted many biologists76, 79, 95, 218. Post-expedition he was helped in the assessment of zooplankton abundance by a student, Kate Ricardo271, who later was to contribute in the field with investigation of fish communities in several large rift-lakes of Central Africa18, 224.

Until the 1930s there was but little knowledge of physical and chemical aspects of vertical stratification in the deeper rift lakes. By 1913, Stappers and others had shown that the deepest lake of all – Tanganyika – was vertically stratified. In 1935–36 another Belgian scientist, Damas, contributed a wider range of findings – physical, chemical and biological – on the stratification of lakes Kivu and Edward51. He showed that deoxygenation of their deeper water was accompanied by large increments of ionic concentration, suggestive of long continuation rather than annual interruption of density stratification. Shortly afterwards, Beauchamp11 made comparable physical and chemical measurements on L. Tanganyika with a longitudinal sequence of stations – and a hand-operated winch! He demonstrated the existence of a large anoxic water-mass below 100–200 m depth. The work was later, in 1939–40, followed up by measurements on changes of stratification in Lake Nyasa (later Malawi)13. Here Beauchamp found short-period oscillations in the depth of individual isotherms that he interpreted as evidence for longitudinal internal waves or seiches, as well as some seasonality in vertical entrainment. Confirmatory evidence of these features was obtained by others in the 1950s and later decades24, 64 when longer time-series of observations were made, some linked with work on fisheries65. There were pioneer studies of the fish stocks by Bertram (Ricardo) et al.18 and later by Lowe161. The last, and the work of Beauchamp and Worthington, were forerunners of much later involvement in African freshwater research from the Freshwater Biological Association80 in Britain.

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In Ethiopia the beginnings of freshwater research were unusual. Before 1935 only a few expeditions with collections of samples205 were recorded. Later the brief period of Italian rule contained several surveys of major rift lakes256 and the elevated Lake Tana193, as well as various small crater lakes.

To 1945 therefore, in tropical Africa overall, we can record important elements of progress with the compositional-structure and (in much lesser degree) dynamics of freshwater environments and their biological communities. In the biological sphere there were – too many to summarize here – further acquisitions of taxonomic, faunistic and floristic information for aquatic organisms from plankton to fishes. River hydrology excepted, there was a marked deficiency of systematic observations maintained over long periods. 1945–1965: era of later colonial administration and its disappearance

With the cessation of hostilities in 1945, initiatives for freshwater research came from several European countries with colonial or dependent territories in Africa. In the main they linked with the development of fisheries institutes and universities, and so with long-maintained operations.

Nevertheless, expeditions continued to be made. Two from Belgium were important for exploration of the large rift lakes of Tanganyika (1946–47) and Kivu, Edward and Albert (1952–54). Inter alia, they provided bathymetric maps of detail and precision from the use of echo-sounding36, structures of physical and chemical stratification37, 147, and comparative studies of chemical composition, plankton252, zoobenthos151, 257 and macrophytes251. Neither was primarily devoted to problems of fisheries, although information on fish faunas 219, 220 and their food258 accrued.

The prospect of more productive fisheries in major African lakes led to the founding of several institutes and mobile teams. Of high scientific distinction, and long persistence, was the East African Fisheries Research Organisation (EAFRO) set up in 1947 beside Lake Victoria at Jinja, Uganda. It was then unusual (and sometimes criticised!) for the wide range of freshwater science that was pursued, partly reflecting the broad interests of its first Director, Beauchamp. His early Annual Reports12 included some speculative portrayals of freshwater ecosystem science. In these early years the staff represented interests in tropical fish biology95, 163, water chemistry, hydrography and phytoplankton73, and aquatic insects168. The range of research was enhanced then and later by encouragement of visiting scientists, including Carter with work on swamp chemistry39, myself in 1956 and with my wife in 1960–61246. The later staff members included



Corbet43, Fryer77, Garrod83, Hamblyn101, Elder and van Someren253, and subsequently Jackson127, Mann174 and Welcomme261.

Another initiative within fisheries research for territories of Central Africa was founded in 1951 from the Colonial Office in London. This, the Joint Fisheries Research Organization (JFRO), operated during 1951–63 over a series of lakes in or bordering the then Northern Rhodesia and Nyasaland – Bangweulu, Malawi128, Kariba104 and Tanganyika44. Its scientific staff included Fryer, Harding, Iles, Jackson and later Bowmaker and Coulter; Jackson has recently given a lively account of its history127. Although fish biology, taxonomy and fisheries received most attention, there were also important contributions in associated chemical surveys104, lake hydrography44, plankton ecology44 and the general biology of aquatic invertebrates78. Some more local surveys of fisheries, also supported from the British Colonial Office, had productive collaborations with other hydrobiologists. Examples include the work of Holden with Green on the River Sokoto117, 118 and with Green and Evans on Lake Albert70, 93, and that on Lake Chad from fisheries-based projects in northern Nigeria119, 227.

The post-1945 years also saw a major development of universities or university colleges, at which the research attractions of African inland waters were recognised and new members of staff with freshwater expertise appeared. In Uganda, at Makerere College (later University), there were Beadle and Lind who had special interests in lakes8 and papyrus swamps that included ecophysiology7 and community floristics157. Hartland-Rowe drew attention to a lunar rhythm in insect emergence110. In the Sudan, at the University College (later University) of Khartoum, was Rzóska (an émigré Polish hydrobiologist) and later Brook, Prowse, Thornton and myself. The College set up a Hydrobiological Research Unit in 1953, equipped with research launch (Fig. 5) and truck that enabled some immense longitudinal surveys of the upper Nile system to be made20,

230, 241. The work included some pioneering estimations of photosynthetic productivity223, but much concentrated on the distribution and seasonality of plankton and associated water characteristics of the White and Blue Niles near Khartoum. There, one predominant influence was the seasonal retention of water in reservoirs upstream31, 223, 232. Another was the reverse wash-out in annual floodwater and reservoir discharge, as had been followed by Abdin1 for phytoplankton of the Aswan reservoir in upper Egypt. During the late 1950s there was a massive and troublesome invasion of the White Nile by the floating water hyacinth Eichhornia crassipes88 that was already established, and studied, in the Congo (Zaïre) system17, 228. The latter and the upper-middle Niger, waters with rich fish faunas221, were investigated by Belgian and French scientists22a, 49, 53, 175.

In the 1950s there was a Belgian initiative, not directly connected with fisheries or universities of Africa, that had no equivalent elsewhere. A

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hydrobiological station was established at Uvira near the northern end of Lake Tanganyika. Its scientific workers included Dubois, Marlier and Symoens. From here time-sequences of observations were possible, which contributed to knowledge of interlinked hydrography, water chemistry58, and planktonic production239; also of the benthic invertebrates176 of this most ancient and unique lake.

Much further south, in the 1960s, another Hydrobiology Research Unit was active in the University College of Rhodesia and Nyasaland (later University of Zimbabwe) at Salisbury (later Harare). Subsequent participants, including Marshall 177, 190, Mitchell 190 and Robarts226, explored conditions in a variety of reservoirs or ‘dams’ that included Lake McIlwaine (= Chivero). Later much attention shifted to the new man-made Lake Kariba. 1965–1985: development of large man-made lakes and international projects

In the years after 1960, large ‘man-made lakes’ 163, 202 were created in tropical Africa for water distribution and power generation. These altered river regimes, with over-year storage, and provided unique examples of succession in aquatic environments and biota. Thus Lake Kariba on the Zambezi (Fig. 4) began to fill in 1958; there followed Lake Nubia-Nasser21,

148 on the Nile, Lake Volta in Ghana149, Lake Kainji in Nigeria126, and Lake Cahora Bassa23 in Moçambique.

FIG. 5. Research on the White Nile, Sudan, 1954: cabin of the launch Malakal, with J. Rzóska (standing) and the author.



Of these, the events ensuing in Lake Kariba were among the most dramatic and best known. They were traced by a series of research groups that included universities at Salisbury (later Harare)171, 189 and Johannesburg (‘Wits’), organisations for fishery research (JFRO104, 127, Kariba Fisheries Institute178, F.A.O. Fisheries Institute4) and the Nuffield Lake Kariba Research Station at Sinamwenda29, 182. Early features included a transitory increase in concentration of plant nutrients (e.g. nitrate); deoxygenation with sulphide production in hypolimnetic water; the displacement of a rich bush fauna; the spread of a water-fern, Salvinia molesta, over a considerable part of the lake surface189; and the successful introduction from Lake Tanganyika of a small pelagic ‘sardine’ Limnothrissa miodon178 that later supported a vigorous fishery. Notable scientific contributions included the demonstration of successive mixing/stratification events in a chain of sub-basins4, the co-existence of small tidal and surface seiche oscillations in water level260, and the influence of diel vertical migrations for the pelagic food chain15. Long-term changes were documented and assembled by Coche, McLachlan and Marshall, and placed in a generalised African setting183.

Much less research effort was given to another man-made lake on the Zambezi, Cahora Bassa23. For this political and military hazards52 were largely responsible. Nevertheless there was a notable investigation of plankton dynamics by Gliwicz90. Favourable conditions with available expertise were available for work on Lake Nasser (Egypt) and lakes Volta and Kainji in West Africa. Both local and international resources were involved. Thus, on the Nile from Aswan and Cairo, studies were made of the radically altered hydrological regime and its consequences for water characteristics68, hydrobiology100 and fisheries148. Much of the research on Lake Volta was carried out by university-based workers from Ghana149 and Britain. The Food and Agricultural Organisation (FAO) made a major contribution with the work of Petr and others on zoobenthos216 and fish biology215. This organisation also supported research on the smaller Lake Kainji of the Niger19, 115, 136, with further continuation from Imevbore and others126 of the University of Ife and work on the main river up- and down-stream97. Also studied in Nigeria were the effects on phytoplankton of impoundment on a smaller river67. A reservoir in Tanzania was the subject of a many-sided investigation from Britain55. Within the new lakes time-successions were evident in communities of plankton, benthos and fish. Interesting interrelations emerged, some involving zoobenthos, periphyton/aufwuchs and plankton as alternative and changing sources of food for fish217 – a group in which seasonal biological rhythms, developed under riverine conditions, were liable to change adaptively with flow variation126a.

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Some hydrological projects involved canals rather than dams. The biggest of these, the Jonglei Canal on the Upper White Nile, was not completed but involved extensive surveys of the Sudd swamp region121a between 1950 and 1980.

During 1965 preparations were being made in both Britain and France for intensive team work on two shallow and productive African lakes. That on Lake George in Uganda was supported by the Royal Society of London; that on Lake Chad by an overseas research organisation (ORSTOM – Office de la Recherche scientifique et technique outre-mer) of the French Government. Work extended over 6–12 years; it was designed to be contributory to a global project, the International Biological Programme (IBP). Organic production was a central theme, with interrelated analyses of planktonic, benthic and fish communities, considered in relation to the physical and chemical environment. The two lakes were hydrological contrasts, with George equable and Chad extremely variable in extent. In consequence, there were numerous ecological contrasts. Individual contributions included work on Lake George by Burgis, Darlington, Dunn, Ganf, Greenwood, Gwahaba, McGowan, Moriarty, Tevlin and Viner, and on Lake Chad by Bénech, Carmouze, Dejoux, Durand, Duwat, Gac, Gras, Iltis, Lauzanne, Lemoalle, Lévêque, Loubens, Quensière and Saint-Jean. Additional studies were made by visiting scientists. There are extended summaries of the overall scientific results33, 38, 96 that are major inputs to tropical freshwater science.

Also in the 1960s and 1970s, a number of smaller university-supported projects yielded results of high general interest. That from the University of Malawi involved a wide range of studies on Lake Chilwa, another hydrologically unstable lake, with events followed over a cycle of drying-up and re-flooding121, 135. From the University of Khartoum there was an exploration of chemical and plankton dynamics, seasonal and inter-annual, in the White Nile and over the river-reservoir cascade system of the Blue Nile103, 191, 235 where the creation of a new upstream reservoir had repercussions below. From Addis Ababa, Ethiopia, came work on crater lakes222, 265 and rift lakes16, 266; from Elizabethville (later Lumumbashi), Zaïre, studies of seasonality of reservoir stratification172, river chemistry240 and phytoplankton production75. A smaller river in Ghana was the base for a detailed examination of seasonal progression in benthic invertebrates125, and one in Gambia for an analysis of solute dynamics153. From Nigeria there was another seasonal study on a small lake influenced by the annual harmattan wind regime105, and which extended to day-night vertical migrations of Chaoborus larvae106. In Kenya, further work on time-sequences in environments and biota was made on lakes (many of them soda lakes) of the rift valley – especially Naivasha, Sonachi, Elmenteita and Nakuru. This involved university scientists from Nairobi, including



Coe42, Gaudet86, 87, Kalff134, Litterick181, Lind156, Mavuti181, Njuguna200 and Nogrady201; also expeditions, North American and German, from abroad that included Cerling40, Hecky114, Kilham142, MacIntyre170, Melack184, 185 and Vareschi254. These made wide-ranging studies, some of fundamental interest, involving chemical balance87, physical circulation170, primary production184, nutrient relationships134 and periodicity of phytoplankton200; also, in Nakuru, subsequent trophic transfer to a highly productive food-web255. From Britain, Green and his associates made contributions for other African water-bodies, including crater lakes in Cameroon94. American and German scientists examined the peculiar vertical stratification in Lake Kivu54, 129, 198, influenced by heating from below and methane accumulation at depth.

There was strong interest in the historical reconstruction of lake environments and biota over very long periods (palaeolimnology). Evidence of major changes of lake level (some in the rifts exceeding 100 m) had come from ancient strand lines199 and mineral deposits81 including fossil diatomites. Use was subsequently made of the detailed chemical and biological record in layered lake sediments. Pioneer studies in the 1960s on lakes Victoria, Naivasha and Tanganyika by Livingstone160 and his students Richardson225 and Kendall141 were followed by others in Ethiopia by Gasse84, 85. In Uganda there was work on Lake George111 and on swamps by Morrison195 and Hamilton102. Historical reconstruction could also be applied to problems of biogeography, as investigated by Dumont and his associates in waters of the Sahel region62, having regard to ‘the Sahara before the desert’ 122 for which French workers had found aquatic relics – including those of hippopotamus, crocodile and Nile perch – to be widespread50, 71, 212.

Inevitably, many had speculated on the nature of the nutrients most limiting for biological production. Based on chemical surveys, importance was attached to phosphorus, nitrogen246 and sulphur14, 116. More direct evidence was obtained from enrichment bioassays and element uptake; examples in East Africa included waters of lakes Victoria69, 72, Naivasha134,

214, Sonachi186 and a fishpond210, in Central Africa various reservoirs226 and Lake Chilwa196, and in West Africa the coastal Ebrié lagoon59.

Three projects on major water-bodies were carried out by larger research teams. That on Lake Victoria centred on the understanding and development of fisheries, and applied some novel fishing methods from abroad under the aegis of the FAO127. The new independence of Kenya and Tanzania was reflected in the development of two more bases for fisheries and associated research on the lake, at Kisumu203 and Mwanza263. Another fisheries-orientated project, on the less known and relatively remote Lake Turkana in Kenya120, was supported by the Overseas Development Agency of Britain. The third was executed by the equivalent French organisation ORSTOM and concerned the large, elongate, coastal Ebrié lagoon of the

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Ivory Coast, West Africa. Here there was a complex interplay of inputs from sea and freshwater plus pollutants from the adjacent city of Abidjan63. These had varied influences on the biota, as shown by studies ranging from bacteria to fish communities.

The period also witnessed a major project of the World Health Organisation (WHO) in West Africa. In parts of this region a disease (onchocerciasis) was rife due to a parasitic filarial nematode transmitted by an insect vector, the blackfly Simulium damnosum complex. The aquatic larvae of the blackfly form part of the zoobenthos of rapidly flowing streams, and were the target of geographically extensive chemical treatment involving the insecticide Abate. Efficacy depended on appropriate ecological knowledge obtained from numerous field studies238. Further freshwater work was related to the transmission of other parasites with a water-connected vector – mosquitoes for malaria91, snails for schistosomiasis32. 1985–2005: further initiatives, large and small

Special interest continued in the very large, deep and ancient lakes of Tanganyika and Malawi. Each was the object of a sizeable research group over several years; that to Tanganyika was set up from Finland159, that to Malawi from Britain188. The results ranged from time-changes of vertical stratification208, 209 to food-webs1a, 159, biodiversity164 and fish production229. There were, independently, other significant contributions by individual scientists41, 66, many attracted by evolutionary issues raised by the rich endemic diversity of cichlid fishes138, 146 and some invertebrate groups45. Here Japan and North America are well represented. Further international projects164 were concerned with biodiversity, cichlid evolution and fisheries.

Lake Victoria, the largest in area of all tropical lakes, was the scene of ecological change that had major socio-economic implications. The fishery was greatly altered in the 1980s by the spread of a large predator, the Nile perch (Lates niloticus), after its previous and controversial introduction77,

204 in the late 1950s. This led to radical change in the food-web and the loss of previously valued species like Oreochromis esculentus83, 174 and other endemic cichlids263. An active project developed on the ecology of haplochromine cichlids, HEST (Haplochromis Ecology Survey Team), based on Dutch-Tanzanian cooperation in the southern region of the lake263. Its members also contributed more widely on the Victoria fish stocks and fisheries264. Although the total fish yield increased after the spread of Lates, the change disadvantaged most local fishermen and much biodiversity was irretrievably lost3. Still more radical, and in part related to increase in the near-shore human population25, was clear evidence of



nutrient enrichment with enhanced planktonic productivity112, 203 and further depletion of oxygen in deep water113. Finally there was, in the 1990s, a troublesome invasion by the water hyacinth, Eichhornia crassipes. All these events and trends led to further international projects, in which the traditional domination of fisheries and fisheries management was followed by emphasis on ecosystem dynamics and wider socio-economic aspects. There was further attention to water movements as a controlling background factor in lakes, both in Victoria170a and the rift lakes of Malawi209 and Tanganyika45.

In two countries there was productive cooperation between Swedish and local scientists supported by the Swedish aid organisation SAREC (Department for Research Co-operation, within the Swedish International Development Agency). For Zimbabwe the focus was the natural economy of Lake Kariba after some 25 years of existence. Local inputs came from the University of Zimbabwe171, 179 and the Kariba Fisheries Institute169. Swedish contributions centred on chemical features158 and aspects of the planktonic and benthic communities46, 137. With Ethiopia the Swedish contribution was primarily by support to research on rift lakes and crater lakes already under investigation from the University of Addis Ababa139. It overlapped another overseas cooperation, involving biologists from the University of Waterloo in Canada. Lake Awassa was a principal research site89, 187, but there were also chemical and plankton surveys involving the comparison of many Ethiopian lakes140, 272 that extended earlier work in the 1960s266. The Austrian Academy of Sciences also gave training in limnology to numerous African professionals, and supported work on the functional ecology of streams in Kenya180. Also in Kenya there was long-continued and cooperative study of the rift lake Naivasha, initiated by Harper from Britain. This documented some major changes over time, mainly related to water balance and introductions of exotic and in part temperate species107, 108 to the relatively cool lake at moderate altitude.

Wider comparative studies were planned within an international project (‘International Decade of the East African Lakes’, IDEAL) that gathered pace in the 1980s and 1990s, and ranged latitudinally from Lake Turkana to Lake Malawi133. These included an emphasis on palaeolimnology, a subject to which others contributed99, 206, 259, and which included consequences of remote and recent climatic change150.

In West Africa, as well as in Chad, the French overseas research organisation ORSTOM (later IRD – Institut de Recherche pour le Développement) continued its work. This included further study of the Ebrié lagoon28, hydrological balance of catchments, lakes and rivers152, 173, stream faunas and anti-onchocerciasis assessment35, colonisation sequence of ponds98, biomass structure in small water-bodies247, and biodiversity of fish communities154, 155.

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Crater lakes continued to be objects of research throughout tropical Africa. From Cameroon there was in August 1986 a most unusual, sudden and damaging release of carbon dioxide of magmatic origin145; in Ethiopia the long-studied group near Bishoftu (Debre Zeit) showed chemical changes under human influence272; and in western Uganda another group provided material for the study of within-year changes of stratification and planktonic populations143, 144. Review and synthesis

Various researchers have chosen to bring together the information available on individual water-bodies. Lakes treated in this way include Kariba4, 194, Chilwa134, McIlwaine = Chivero248, Chad38, Tanganyika45 and Nasser21; rivers include the Nile231 and Niger97. In some cases the material surveyed was largely derived from a single extended research project. At the opposite extreme the monograph on the Nile river system, initiated by Rzóska, took in very diverse sources over a large geographical area and historical attribution. Regional surveys included one on West African waters132 and another on Ethiopian rift lakes250. African shallow water-bodies were treated in a cooperative international effort with components of directory34, bibliography and synthesis61, 243. Information from some individual countries, including Sudan60, Ethiopia249, Ghana74, Malawi197 and Zimbabwe233, has also been summarised.

Besides the obvious descriptive element, particular water-bodies can emphasize general topics or themes. For example, lakes Chilwa and Chad demonstrate time-successions arising from hydrological change, and man-made lakes like Kariba those from a recent origin. Such general topics have also been treated comparatively from wider experience over tropical Africa. There are examples for sequences of colonisation183, shallow lakes61, 243, water chemistry246, plankton seasonality109, 114, 242, aquatic vegetation56, floodplain dynamics262, biodiversity of fishes154, 270, fisheries research164 and conservation issues25, 165.

A single book, by Beadle9, has combined (up to 1980) regional descriptions and subject reviews over tropical Africa as a whole. Two others211, 245 have taken the still more extended view of general ecological science for tropical inland waters worldwide. Tropical fish communities have been similarly treated163. African experience then provided examples within a generalised scientific framework. This is particularly apt for lake developments in time, chemical control of aquatic biota, hydrodynamics of large warm lakes, temperature – production relationships, the determination of seasonality at low latitudes, and the localisation or adaptive radiation of genetic novelty.



Acknowledgements

I am indebted to Geoffrey Fryer, Peter Jackson, Jacques Lemoalle, Rosemary Lowe-McConnell and Jean-Jacques Symoens for helpful suggestions and criticism. Also to ‘Editions d’IRD’, for permission to use Fig. 1 originally published (in ref. 243) by ORSTOM, Paris. References

1. Abdin, G. (1948). The conditions of growth and periodicity of the algal flora of the Aswan reservoir (Upper Egypt). Bull. Fac. Sci. Egypt. Univ. 27, 157–175.

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