Until the 1970s, the fish fauna of Lake Victoria in East Africa was dominated by about 500 endemic haplochromine cichlid species, which comprised about 80% of the demersal fish mass. The cichlids were extremely diverse ecologically; however, the small diversity in gross morphology and the presence of intraspecific variation made it difficult to distinguish among species. In the first half of the 1980s, the Nile perch Lates niloticus, an introduced predator, suddenly boomed and cichlids declined dramatically. During the same period eutrophication increased strongly. With the decline of Nile perch catches in the 1990s, the cichlids showed some recovery. These events have triggered many studies and debates. Disagreements about the severity and causes of the decline often stemmed from considering the cichlid flock as a single unit owing to the lack of proper taxonomic and ecological knowledge. By studying cichlid communities, trophic groups, and individual species, researchers uncovered differential impacts that helped them to unravel the causes of the changes. It seems that lakewide Nile perch predation and eutrophication had the strongest impact and that the fishery only had a local effect. Knowledge of the differential decline and recovery of the haplochromine cichlids and the underlying causes is important for the proper management of biodiversity and the fishery in Lake Victoria. Consequently, knowledge of the systematics of cichlids is a key issue in managing the lake.
Lake Victoria had a fish fauna dominated by 500+ species of haplochromine cichlids that made up more than 80% of the fish mass. The five main trophic groups caught with bottom trawlers in the sub-littoral areas of the Mwanza Gulf were: detritivores, zooplanktivores, insectivores, molluscivores and piscivores. The detritivores (13+ species) formed the most important guild, making up 60-80% of the number of individuals, followed by the zooplanktivores (12+ species), which comprised 10-30%. In the 1980s the haplochromines from the sub-littoral and offshore areas (estimated at some 200 species) vanished almost completely. Commercial trawl fishery, the upsurge of the introduced Nile perch, and an increase of eutrophication were potential causes of this decline. In the 1990s, when Nile perch was heavily fished, a recovery of some haplochromine species was observed. We studied the decline and partial recovery of the different haplochromine trophic groups in the northern part of the Mwanza Gulf. The rate at which the trophic groups declined differed; the relatively large piscivores, insectivores and molluscivores were the first to disappear from the catches. The small detritivores and zooplanktivores declined at lower rates, especially the latter group. From the beginning of the 1990s a resurgence of both groups was observed. By 2001, the zooplanktivores had reached their previous level of abundance, but their diversity declined from more than 12 species to only three. Though four detritivorous species began being regularly caught again, they constituted only about 15% of the number of individuals, while the zooplanktivores made up more than 80%. The patterns of decline and recovery indicate that, though fishery played a role locally, predation by Nile perch and eutrophication were the main factors determining the fate of the haplochromines. However, it has so far been impossible to establish the causal relationship between the two, and the relative impact of each of these phenomena separately. The potential effects of the changed trophic dominance, and the importance of the haplochromines for the ecosystem and a sustainable fishery, are discussed.
Rapid morphological changes in response to fluctuating natural environments are a common phenomenon in species that undergo adaptive radiation. The dramatic ecological changes in Lake Victoria provide a unique opportunity to study environmental effects on cichlid morphology. This study shows how four haplochromine cichlids adapted their premaxilla to a changed diet over the past 30 years. Directly after the diet change toward larger and faster prey in the late 1980s, the premaxilla (upper jaw) changed in a way that is in agreement with a more food manipulating feeding style. During the 2000s, two zooplanktivorous species showed a reversal of morphological changes after returning to their original diet, whereas two other species showed no reversal of diet and morphology. These rapid changes indicate a potential for extremely fast adaptive responses to environmental fluctuations, which are likely inflicted by competition release and increase, and might have a bearing on the ability of haplochromines to cope with environmental changes. These responses could be due to rapid genetic change or phenotypic plasticity, for which there is ample evidence in cichlid fish structures associated with food capture and processing. These versatile adaptive responses are likely to have contributed to the fast adaptive radiation of haplochromines.
ABSTRACT. East Africa's Lake Victoria provides resources and services to millions of people on the lake's shores and abroad. In particular, the lake's fisheries are an important source of protein, employment, and international economic connections for the whole region. Nonetheless, stock dynamics are poorly understood and currently unpredictable. Furthermore, fishery dynamics are intricately connected to other supporting services of the lake as well as to lakeshore societies and economies. Much research has been carried out piecemeal on different aspects of Lake Victoria's system; e.g., societies, biodiversity, fisheries, and eutrophication. However, to disentangle drivers and dynamics of change in this complex system, we need to put these pieces together and analyze the system as a whole. We did so by first building a qualitative model of the lake's social-ecological system. We then investigated the model system through a qualitative loop analysis, and finally examined effects of changes on the system state and structure. The model and its contextual analysis allowed us to investigate system-wide chain reactions resulting from disturbances. Importantly, we built a tool that can be used to analyze the cascading effects of management options and establish the requirements for their success. We found that high connectedness of the system at the exploitation level, through fisheries having multiple target stocks, can increase the stocks' vulnerability to exploitation but reduce society's vulnerability to variability in individual stocks. We describe how there are multiple pathways to any change in the system, which makes it difficult to identify the root cause of changes but also broadens the management toolkit. Also, we illustrate how nutrient enrichment is not a self-regulating process, and that explicit management is necessary to halt or reverse eutrophication. This model is simple and usable to assess system-wide effects of management policies, and can serve as a paving stone for future quantitative analyses of system dynamics at local scales.
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