Primary production rates, chlorophyll and phytoplankton biovolume were measured monthly from April 2003 to November 2004 in Lake Tana, a large tropical lake in the highlands of Ethiopia. The lake is characterised by low nutrient concentrations, and a low water transparency due to high silt load of the inflowing rivers during the rainy seasons (May-November) and daily resuspension of sediments in the inshore zone. The mean chlorophyll-a concentrations varied seasonally and ranged from 2.6 mg m -3 to 8.5 mg m -3 (mean: 4.5 mg m -3 ) in the offshore zone. Primary production was measured using the light-dark bottles technique. We incubated only at three depths, i.e. 0.6, 1.2 and 1.8 m. Therefore, we may have missed a substantial part of the depth production profile and probably also frequently missed P max . Gross primary production in the openwater averaged 2.43 g O 2 m -2 d -1 and ranged between 0.03 g O 2 m -2 d -1 and 10.2 g O 2 m -2 d -1 ; production was significantly higher in the inshore zone. The highest production rates were observed in the post-rainy season (Oct-Nov), which coincided with a bloom of Microcystis and higher chlorophyll levels. This seasonal high production is probably caused by a relatively high nutrient availability in combination with favourable light conditions. The gross primary production rates of L. Tana are among the lowest compared with other tropical lakes. This will be partly the result of our underestimation of gross primary production by often missing P max . Another cause is the oligotrophic nature of the lake in combination with its relatively low water transparency. The gross primary production per unit chlorophyll in the openwater zone was in the same range as in 30 other tropical lakes and reservoirs. The higher primary production in the inshore zone is probably the result of the daily water column mixing (Z mix ‡ Z t ) in this area, enhancing nutrient recycling. A large proportion of the annual primary production is realised in one of the four seasons only. This productive post-rainy season is relatively short (2 months) and therefore efficiency of transfer of matter between the first and second trophic level of the Lake ecosystem will be poor.
Studying species flocks (e.g. Darwin's finches, Caribbean anoline lizards, East African cichlid fishes) has proven to be highly successful in understanding the forces driving speciation. The only known, intact species flock of cyprinid fishes, the 15 Labeobarbus species in Lake Tana (Ethiopia), includes eight piscivorous species. Piscivory is a rare specialisation among the highly successful (>2000 species) but mostly benthivorous Cyprinidae. The extent and mechanisms of diversification of this remarkable Labeobarbus species flock, particularly among the unexpected piscivorous species, are still largely unknown. In the present study we demonstrate that all 15 Labeobarbus species are segregated to a great extent along spatial, trophic and/or temporal dimensions. The spatial distribution, diet (prey species but not prey size), time of active feeding and predation techniques differed significantly among the eight piscivores. Lake Tana's cyprinids displayed their retained potential for ecological diversification and speciation, including the uncommon specialisation of piscivory. The latter is probably a result of the absence of common African specialist piscivores in Lake Tana. We suggest that the evolution of Lake Tana's Labeobarbus species flock at this stage is predominantly structured by ecological selection models. The labeobarbs most likely underwent sequential stages of radiation and speciation: habitat divergence followed by trophic divergence.
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