The history of fish communities, biodiversity and environment of Lake Victoria and the lessons learnt

The first fishery survey of Lake Victoria was conducted between 1927 and 1928 (Graham 1929). Atthat time, the lake had a diverse fish fauna and the fishery was dominated by two endemic tilapiine cichlids; Oreochromis esculentus (Graham 1929) and O. variabilis (Boulenger 1906). There were a number of other species such as Protopterus aethiopicus Heckel 1851, Bagrus docmac (Forsk.) 1775, Clarias gariepinus (Burchell), Barbus species, mormyrids, Synodontis spp, Schilbe intermedius (Linn.) 1762 and Rastrineobola argentea Pellegrin, 1904 that were also abundant in the lake most of which made a significant contribution to the fishery (Graham 1929, Worthington 1929, 1932, Kudhongania & Cordone 1974). Haplochromine cichlids were represented by at least 300 species more than 99% of them endemic (Greenwood, 1974; Witte et al., 1992 a & b). The fishery of Lake Victoria was similar to that of lakes Kyoga and Nabugabo (Worthington 1929; Trewavas 1933; Greenwood 1965, 1966; Beadle 1962, 1981). There were also important fisheries on the inflowing rivers of Lake Victoria, the most important of which were Labeo victorianus and Barbus altianalis (Cadwallader 1965). The small sized species notably Rastrineobola argentea and haplochromines cichlids were not originally commercially exploited.

Nutrient status and thermal stratification in Lake Victoria

Worldwide, human activity in the watershed has been found to induce lake responses at various levels, including at population and ecosystem scale. Recently, Carignan and Steedman (2000) reported on disruptions of biogeochemical cycles in temperate lakes following watershed deforestation and lor wildfire and Carignan et al., (2000 a, b) concluded that water quality and aquatic biota are strongly influenced by disturbances in the watershed. Similarly, Lake Victoria is no exception as people in its catchment have exploited it for the last hundred years or more, but have now begun to understand the extent to which they have thrown the lake into disorder and how their increasing activity in the watershed have driven some environmental changes within and around the lake.

The major aquatic systems of the Victoria and Kyoga lake basins

The Victoria and Kyoga lake basins form the major aquatic system of this study (Fig. I). The two lake basins share a common evolutionary history and have similar native fish faunas (Graham 1929, Worthington 1929). The two main lakes have also had similar impacts by introduction of Nile perch Lates niloticus and therefore these two lakes can be considered to be similar for ichiogeographical purposes. These lake basins have many satellite lakes isolated from one another and from the main lakes Victoria and Kyoga by swamps and other barriers. Some of these satellite lakes still possess stocks of endemic fish species which are almost extinct from the main water bodies. It was therefore considered that understanding of these lakes would contribute to the knowledge base required to solve some of the problems experienced in Lake Victoria and Kyoga especially the loss in trophic diversity arising. The study was carried out in these two main water bodies (Kyoga and Victoria) and on other satellite lakes e.g Wamala, Kachera, Mburo, Kayanja and Kayugi in the Victoria lake basin and lakes Nawampasa, Nyaguo, Agu, Gigate, Lemwa and Kawi in the Kyoga lake basin (Figs. 2, 3, 4, 5 & 6).

Physical and chemical characteristics of major aquatic ecosystems in the Victoria and Kyoga lake basins

The physical-chemical characteristics of any aquatic ecosystem include pH, conductivity, and temperature, water transparency, nutrient and the chlorophyll-a levels. Physical and chemical factors of any ecosystem determine the type and quality of flora present in it and these forms the basis on which the system operates. The elements required in largest amounts for plant productions are carbon, phosphorus, nitrogen, and silicon, which is important for diatoms as a major component of the cell wall. Nutrients may limit algal productivity in the tropics despite the high temperature there allowing rapid nutrient recycling. Nutrients most likely to be limiting African lakes are nitrogen (Talling & Talling 1965; Moss 1969; Lehman & Branstrator 1993, 1994) and phosphorus (Melack.et al l982; Kalff 1983) while silicon may limit diatom growth (Hecky & Kilham 1988). The objective of the study is to investigate the impact of physical-chemical characteristics on the distribution and abundance of organisms in the major aquatic ecosystems.

The diversity of aquatic flora and their importance in the Victoria and Kyoga lake basins

A great part of Uganda is endowed with water bodies in the forms of rivers and open water lakes. These bodies are never alone. They are either flanked or associated with plants, which are adapted to the wet conditions. They are so characteristic that they are part and parcel of the aquatic ecosystems. They occupy various positions depending on the amount of water in the relevant habitats.

The diversity and abundance of zooplankton in the Lake Victoria and Kyoga basins and their relationship to fish production

Biological diversity of an ecosystem is considered a reliable measure of the state of health of the ecosystem. In Uganda's large lakes, the Victoria and Kyoga, the past three decades have been characterized by profound changes in fish species composition following the introduction of the piscivorous Nile perch (Oguto-Ohwayo 1990). Over 300 haplochromine cichlid species comprising a wide range of trophic groups were lost along with a host of non-cichlid fishes which occupied virtually all available ecological niches and in the lakes (Witte 1992). A second major ecological event has been the gradual nutrient enrichment of the water bodies (eutrophication) from diffuse and point sources, while at the same time pollutants have also gained entrance into the water systems in pace with indusfrial development and human population increases in the lake basins. Eutrophication and pollution have drastically altered the physical and-chemical character of the water medium in which different fauna and flora thrive. In Lake Victoria these alterations have resulted in changes of algal species composition from pristine community dominated by chlorophytes and diatoms (Melosira etc) to one composed largely of blue-green algae or Cyanobacteria (Microcystis, Anabaena, Planktolyngbya etc) (Mugidde 1993, Hecky 1993).

Fish species diversity in the Victoria and Kyoga lake basins: their conservation and sustainable use

Introduction of exotic fish species especially the Nile perch Lates niloticus, is believed to be responsible for the decline of fish species diversity in lakes Victoria, Kyoga and Nabugabo.About 60% of the haplochromine cichlids are thought to have become extinct from L. Victoria due to predation by the Nile perch. However there are many lakes satelite to the lakes Victoria and Kyoga basins which still have fish fauna similar to that of the main lakes. many of the satellite lakes are separated from the main lakes in, which Nile perch was introduced by extensive swamps that provide a barrier to Nile perch .A survey was carried out in a number of these satelite lakes and an inventory made of existing fish species. Their distribution and relative abundances were also determined. The lakes studied included Nawampasa, Nakuwa,Kawi Lamwa Gigate, Nyaguo, Agu, Nabugabo. Kayanja, Kaytigi, Mburo, Kachera and Wamala.Some habitats within the main lakes Victoria and Kyoga, especially those with rocky outcrops· and macrophyte cover that provide refugia for endangered species from Nile perch,were also surveyed) Various stations along the River Nile were also sampled to quantify the fish species that are still resent. Kyoga minor lakes were found to have the highest number of fish species especially of haplochromine cichlids. Many haplochromine trophic groups that were thought to be extinct from 1. Victoria still occur in these lakes.!Some of the satellite lakes, especially lakes Kayugi, Mburo and Kachera still contain .healili populations of oreochromis. I esculentus that could be used as brood stock in fish farming. Many of these lakes should .I ( I therefore be protected for conservation offish species diversity

The diversity of amphibians, reptiles and mammals and their importance in the Victoria and Kyoga lake basins

The shore margins of Lakes in the Victoria basin are highly dented and mostly swampy, fringed by Papyrus and other wetland vegetation types important habitats for herpetofauna and wetland adapted mammals. Of recent, the extent of the 'wetland' has been extended in several places by the Water Hyacinth (Eichornia cryaseps). Ecologically, amphibians are important in many ways; they are mostly predators, acting as primary and secondary carnivores. Their prey consists mostly of insects, some of which are pests to crops or disease vectors. They are also inter-inked in food chains, often acting as food for other vertebrates, such as pigs, birds, snakes and sometimes man. Because of their ectothermic physiology, the life history and ecology of amphibians often differ markedly from that of birds or mammals (McCollough el ai, (992).Amphibians are known to be an easily recognisable taxon in given habitats; and populations are sometimes specialised within a narrow habitat. This makes it easy and practical to monitor changes in composition over time, given different onditions (Heyer el al 1994, Phillips 1990). Impacts on their habitat are reflected in changes in numbers and species diversity in a short time. These are some of the factors that have made amphibians to be recognised, nowadays, as good indicators of habitat change

Genetic status of selected fish taxa in relation to conservation of genetic and species diversity in the Victoria and Kyoga Lake basins

Cichlids are known for their explosive radiation especially in the African Great Lakes marked with a high level of lake endemism. These fishes have been characterized mainly along trophic and habitat differences, by variation in morphological structures such as teeth and jaws and by differences in body shape and coloration. Cichlids are important as a microcosm of macroevolution. The explosive radiation, young evolutionary scale, and the isolation of groups characterized with high levels of endemism and presence of living fossils makes the group important for evolutionary and genetic studies. Lake Victoria region cichlids which are isolated and relatively more recent in evolution were the last to be appreciated in their diversity. Recently Ole Seehausen has found scores of rock fishes in Lake Victoria which were up to then thought to be absent from the Lake and only known to occur in Lakes Malawi and Tanganyika. Greenwood put together the species groups of Lake Victoria, and later in the early 1980's revised the classification of haplochromine species to reflect the phyletic origin and interrelationship of the various groups in Lake Victoria region. Melan Stiassny has been interested in early evolution of cichlids while the likes of Paul Fuerst and Lees Kaufman and Axel Meyer have been interested and are working to explain the speciation mechanisms responsible for the explosive radiation and evolution of cichlids. Locally S.B Wandera and his student Getrude Narnulemo are spearheading the biodiversity and taxonomic studies of cichlids in Lake Victoria region

The biology, ecology and the fishery of Mukene, Rastrineobola argentea in Lakes Victoria, Kyoga and Nabugabo

Rastrineobola argentea, local name.s Mukene (Uganda) Omena (Kenya) and Dagaa (Tanzania) occurs in lakes Victoria, Kyoga and Nabugabo (Greenwood 1966). Until the decline of the native fishes especially the haplochromines, R. argentea was of little economic importance to the fisheries of these lakes. The stocks of this species have now increased and commercial catches on Lake Victoria account for over 30%. Fishing for R. argentea is currently still restricted to inshore areas and within the fringing islands of Lake Victoria. This fishery is not yet established on Lakes Kyoga and Nabugabo although the species is quite abundant in these lakes (Proude 1963). On moonless nights, kerosene pressure lamps are exposed on the lake to attract the fishes. These fishes are then fished out using fine meshed seine, lampara or lift nets

The biology and ecology of native non-cichlids in the Victoria and Kyoga lake basins

An overview of the biology and ecology of some of the constantly less important commercial species is given below. These included Bagrus docmac, Clarias gariepinus, Protopterus aethiopicus, Labeo victorianus, Barbus spp, Mormyrids, Synodontis spp, and Schilbe intermedius. The stocks of most of these species declined due to over-exploitation and introduction of non-native fishes especially Nile perch. A few of these taxa still survive in the main lake and others in satellite lakes. The current status of these species in the Victoria lake basin is not known but the available information provided some information on some habitat and other requirements of some of these originally important species of the Victoria lake basin.

Trophic interrelationships and food-webs among the fishes in ecosystems of the Victoria and Kyoga lake basin

The Victoria and Kyoga lake basins had a high fish species diversity with many fish species that were found only in these lakes. Two Tilapiines species Oreochromis esculentus and Oreochromis variabilis were the most important commercial species in these lakes and were found nowhere else on earth except in the Victoria and Kyoga lake basins (Graham 1929, Worthington 1929). Lakes Kyoga and Nabugabo also had endemic haplochromine species (Worthington 1929, Trewavas 1933, Greenwood 1965, 1966). As stocks of introduced species increased, stocks of most of the native species declined rapidly or disappeared altogether. The study was carried out on Lakes Victoria and Kyoga, River Nile, some selected satellite lakes from the two basins namely Lakes Mburo, Kachera, Wamala, Kayanja, Kayugi, Nabugabo, Victoria, Victoria nile and River Sio(Victoria lake basin). Lakes Kyoga (Iyingo), Nawampasa, Nakuwa, Gigati, Nyaguo, Agu, Kawi and Lemwa (Kyoga lake basin). Species composillon and relative abundance of fishes were estimated by detennining the overall average total number of each species encountered. A trophic consists of species using the same food category. Shannon-Weaver Index of diversity H (Pielou, 1969) and number of trophic groups, were used to estimate the Trophic diversity of various fish species in the lakes. Food analysis has been done on some fishes in some of the sampled lakes and is still going on, on remaining fishes and in some lakes. Generally fish ingested detritus, Spirulina, Melosira, filamentous algae, Planktolyngbya, Microcysists, Anabaena, Merismopedia, Spirogyra, higher plant material, rotifers, Ostracodes, Chironomid larvae and pupae, Choaborus larvae, Odonata, Povilla, Insect remains, Caridina, fish eggs and fish. Eight trophic groups were identified from thes food items ingestes. These included detritivores, algae eaters, higher plant eaters, zooplanktivores, insectivores, molluscivores, prawn eaters, paedophages and piscivores. Trophic diversity by number of trophic groups was highest in Lake Kyoga (6) followed by lakes Kayugi, Nabugabo, River Nile and Mburo (3) and the lowest number was recorded in kachera (2).

Fishing gear selectivity of Lates niloticus L. (Nile perch), Oreochromis niloticus L. (Nile tilapia) and Rastrineobola argentea Pellegrin (Mukene) in lakes Victoria, Kyoga and Nabugabo

Catch effort data on which fisheries management regulations are sometimes based are not available for most lakes in Uganda. However, failure to regulate fishing gears and methods has been a major cause of collapse of fisheries in the country. Fisheries have been damaged by destructive and non-selective fishing gears and methods such as trawling and beach seining, by use of gill nets of mesh size which crop immature fish and by introduction of mechanised fishing. Selectivity of the gears used to crop Lates niloticus 1. (Nile perch), Oreochromis niloticus 1. (Nile tilapia) and Rastrineobola argentea (Mukene) which are currently the most important commercial species in Uganda were examined in order to recommend the most suitable types, sizes and methods that should be used in exploiting these fisheries . Gill nets of less than 127 mm mainly cropped immature Nile ti1apia and Nile perch. To protect these fisheries, the minimum mesh size of gill nets should be set at 127 mm. Seine nets of 5 mm do catch high proportions of immature Mukene while those of 10 mm catch mainly mature Mukene. When operated inshore, both sizes catch immature Nile perch and Nile ti1apia as by-catch. To protect the Mukene fishery and avoid catching immature byecatch, a minimum mesh size of the Mukene net should have been 10 mm operated as Lampara type net offshore but since most fishennen have been using the 5 mm seine for over five years the minimum size should not be allowed to drop below 5 mm pending further thorough investigations. Beach seining, trawling and are destructive to fisheries and should be prohibited until data that may justify their use is available.

The biology and ecology of the Nile perch, Lates niloticus, and the future of its fishery in Lakes Victoria, Kyoga and Nabugabo

Lakes Victoria, Kyoga and Nabugabo had a similar native fish fauna of high species diversity. stocks of most of the native species declined rapidly and some completely disappeared after Nile perch was introduced and became well established. Although, overexploitation of the fish stocks, competition between introduced and native tilapiines and environmental degradation contributed to the reduction in fish stocks, predation by the Nile perch has contributed much to the recent drastic reductions in fish stock and could even drive the stocks to a total collapse. Nile perch is also currently the most important commercial species in Lakes victoria, Kyoga and Nabugabo and the stability of its stocks is important in the overall sustainability of the fisheries of these lakes. The question that was to be examined in this paper was whether the fisheries of Lakes Victoria, Kyogaand Nabugabo would stabilize and sustain production in the presence of high predation pressure by the Nile perch or whether the Nile perch would drive the fish stocks including itself to a collapse. I t was assumed that Nile perch driven changes in Lakes Victoria, Kyoga and Nabugabo would be driven to a level beyond which they would not change further. This would be followed by recovery and stability or the changes would continue to a point of collapse. It was assumed that Lake Albert represented the ideal stable state. The changes in the new habitats expected to be driven through a major change due to Nile perch predation to a stage where there would be no further changes. After this, a feedback mechanism would move the driven variable towards recovery. The variables would then stabilize and oscillate will an amplitude which approximates to what would be recorded in Lake Albert. Alternatively, the changes would proceed to a stage where the fishery would collapse. The specific hypothesis was that fish species composition and diversity, prey selection by the Nile perch and life history characteristics of the Nile perch in the new habitats would change and stabilize

The biology, ecology, distribution and conservation of surviving haplochromine cichlids in Lakes Victoria, Kyoga and Nabugabo

Haplochrmine cichlids were the most abundant taxa in Lakes Victoria, Kyoga and Nabugabo prior to introduction of the Nile perch. As stocks of the introduced predator increased, these taxa were depleted to such an extent that they are now virtually absent from the lake. The haplochromine cichlids played an important role in the ecology of Lakes Victoria, Kyoga and Nabugabo. They occupied virtually all trophic levels in the lake and facilitated an efficient flow of energy through the ecosystem. Their depletion seem to have left much organic matter whose decomposition has contributed to accumulation of dead organic matter which may be contributing to prolonged anoxia in Lake Victoria. The haplochromines formed an important small-scale fishery. Fishermen formerly subsisting on this fishery have been driven out of business because they cannot afford the expensive nets required for Nile perch fishery. In addition to providing a cheap source of fish protein to humans, the species were an important source of Scientific material for students of genetics antd adaptive radiation.