Mary A. Kishe-Machumu

Species Distinction and the Biodiversity Crisis in Lake Victoria

Abstract 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 ...

Differential decline and recovery of haplochromine trophic groups in the Mwanza Gulf of Lake Victoria

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.

Dietary shift in benthivorous cichlids after the ecological changes in Lake Victoria

Cichlid fishes of Lake Victoria are well known for their trophic specializations. In our study area, the sub-littoral waters of the Mwanza Gulf (Tanzania), at least 12 trophic groups coexisted in the 1970s. After the Nile perch upsurge and eutrophication in the 1980s, most cichlids disappeared from the area. During the 1990s, heavy exploitation of Nile perch reduced the abundance of this predator. Subsequently, some cichlid species, mainly zooplanktivores and small benthivores, recovered. To establish their ecological role in the changed environment, among other factors we studied their diet. Stomach contents of fish caught before and after the ecological changes were compared. The small benthivores shifted from a diet dominated by detritus and phytoplankton, supplemented with small quantities of midge larvae and zooplankton, to a diet of invertebrate preys of larger sizes. Currently, the diet includes zooplankton, midge larvae, shrimps and molluscs, and a very small amount of detritus and phytoplankton. Apparently, the recovering benthivores have a more carnivorous than a herbivorous diet. Concomitantly, the ratio of intestine length to standard length of the benthivores decreased with 30% and the average stomach fullness decreased as well. The observed dietary shifts may be due to (1) the changed availability of food types and a decline of the nutritious quality of detritus and phytoplankton due to the strong increase of cyanobacteria in the lake, (2) the decline of the former trophic specialists, and (3) the reduced water clarity. The diet of the former detritivores is currently very similar to that of the zooplanktivores. Consequently, if diet plays a role in the relatively slow recovery of the small benthivores compared to the zooplanktivores, this would only hold if they are relatively poorly adapted to their new diet.

The diet of Nile perch, Lates niloticus (L.) after resurgence of haplochromine cichlids in the Mwanza Gulf of Lake Victoria.

Haplochromine cichlids used to be the main prey of the introduced Nile perch, Lates niloticus, in Lake Victoria. After depletion of the haplochromine stocks at the end of the 1980s, Nile perch shifted to the shrimp Caridina nilotica and to a lesser degree to its own young and the cyprinid Rastrineobola argentea. In the present study, we investigated the Nile perch diet in the northern Mwanza Gulf after resurgence of some of the haplochromine species and compared it with data collected in the same area in 1988/1989. It became clear that haplochromines are again the major prey of Nile perch. The dietary shift from invertebrate feeding (shrimps) to feeding on fish (haplochromine cichlids) occurs at a smaller size than it did when Nile perch were taking primarily dagaa and juvenile Nile perch as their fish prey. The apparent preference for haplochromines as prey has reduced the degree of cannibalism considerably, which may have a positive impact on Nile perch recruitment.

Fast adaptive responses in the oral jaw of Lake Victoria cichlids

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.

Stable isotope evidence from formalin–ethanol-preserved specimens indicates dietary shifts and increasing diet overlap in Lake Victoria cichlids

Stable isotopes are increasingly being used to infer past and present trophic interactions in light of environmental changes. The Lake Victoria haplochromine cichlids have experienced severe environmental changes in the past decades that, amongst others, resulted in a dietary shift towards larger prey. We investigated how the changed environment and diet of the haplochromines influenced stable isotope values of formalin-then-ethanol-preserved cichlid specimens, and then investigated how these values differed among species before (1977–1982) and after substantial environmental changes (2005–2007). We found a small preservation effect on both δ13C and δ15N values, and significant differences in isotope values among haplochromine species collected before the environmental changes. In contrast, there was a remarkable similarity in δ13C and δ15N values among species collected from the contemporary ecosystem and two out three species showed significantly different stable isotope values compared to species of the historic ecosystem. In addition, we found a putative isotopic gradient effect along our 5-km-long research transect indicating that the studied demersal species are more stenotopic than previously thought. The environmental changes have resulted in dietary change and overlap of the haplochromines which provides insight into the trophic plasticity of these species, which are often considered trophic specialists.