1. This study focused on phytoplankton production in Lake Tanganyika. We provide new estimates of daily and annual primary production, as well as growth rates of phytoplankton, and we compare them with values published in former studies. 2. Chlorophyll-a (chl-a) in the mixed layer ranged from 5 to 120 mg chl-a m )2 and varied significantly between rainy and dry seasons. Particulate organic carbon concentrations were significantly higher in the south basin (with 196 and 166 mg C m )3 in the dry and the rainy season, respectively) than in the north basin (112 and 109 mg C m )3 , respectively). 3. Carbon : phosphorus (C : P) ratios varied according to season. Phosphorus limitation seemed to occur more frequently than nitrogen limitation, especially during the rainy season. Severe P deficiencies were rare. 4. Measured particulate daily primary production ranged from 110 to 1410 mg C m )2 day )1 ; seasonal contrasts were well marked in the north basin, but less in the south basin, where primary production peaks occurred also in the rainy season. Estimates of annual primary production, based on daily primary production calculated from chl-a and water transparency, gave values lower than those reported in previous studies. Picophytoplankton accounted on average for 56% of total particulate production in the south basin during the wet season of 2003. 5. Phytoplankton growth rates, calculated from primary production, ranged from 0.055 to 0.282 day )1 ; these are lower than previously published values for Lake Tanganyika.

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In keeping with the theme of this volume, the present article commemorates the 50 years of Hutchinson's (Am Nat 93:145-159, 1959) famous publication on the 'very general question of animal diversity', which obviously leads to the more important question regarding the driving forces of biodiversity and their limitation in various habitats. The study of phytoplankton in large lakes is a challenging task which requires the use of a wide variety of techniques to capture the range of spatial and temporal variations. The analysis of marker pigments may provide an adequate tool for phytoplankton surveys in large water bodies, thanks to automated analysis for processing numerous individual samples, and by achieving sufficient taxonomic resolution for ecological studies. Chlorophylls and carotenoids were analysed by HPLC in water column samples of Lake Tanganyika from 2002 through 2006, at two study sites, off Kigoma (north basin) and off Mpulungu (south basin). Using the CHEMTAX software for calculating contributions of the main algal groups to chlorophyll a, variations of phytoplankton composition and biomass were determined. We also investigated selected samples according to standard taxonomic techniques for elucidating the dominant species composition. Most of the phytoplankton biomass was located in the 0-40 m layer, with maxima at 0 or 20 m, and more rarely at 40 m. Deep chlorophyll maxima (DCM) and surface 'blooms' were occasionally observed. The phytoplankton assemblage was essentially dominated by chlorophytes and cyanobacteria, with diatoms developing mainly in the dry season. The dominant cyanobacteria were very small unicells (mostly Synechococcus), which were much more abundant in the southern basin, whereas green algae dominated on average at the northern site. A canonical correspondence analysis (CCA) including the main limnological variables, dissolved nutrients and zooplankton abundance was run to explore environment-phytoplankton relations. The CCA points to physical factors, site and season as key determinants of the phytoplankton assemblage, but also indicates a significant role, depending on the studied site, of calanoid copepods and of nauplii stages. Our data suggest that the factors allowing coexistence of several phytoplankton taxa in the pelagic zone of Lake Tanganyika are likely differential vertical distribution in the water column, which allows spatial partitioning of light and nutrients, and temporal variability (occurring at time scales preventing long-term dominance by a single taxon), along with effects of predation by grazers.

1. Abundance and bacterial production (BP) of heterotrophic bacteria (HBact) were measured in the north and south basins of Lake Tanganyika, East Africa, during seasonal sampling series between 2002 and 2007. The major objective of the study was to assess whether BP can supplement phytoplankton particulate primary production (particulate PP) in the pelagic waters, and whether BP and particulate PP are related in this large lake. HBact were enumerated in the 0-100 m surface layer by epifluorescence microscopy and flow cytometry; BP was quantified using 3 H-thymidine incorporation, usually in three mixolimnion layers (0-40, 40-60 and 60-100 m). 2. Flow cytometry allowed three subpopulations to be distinguished: low nucleic acid content bacteria (LNA), high nucleic acid content bacteria (HNA) and Synechococcus-like picocyanobacteria (PCya). The proportion of HNA was on average 67% of total bacterial abundance, and tended to increase with depth. HBact abundance was between 1.2 · 10 5 and 4.8 · 10 6 cells mL )1 , and was maximal in the 0-40 m layer (i.e. roughly, the euphotic layer). Using a single conversion factor of 15 fg C cell )1 , estimated from biovolume measurements, average HBact biomass (integrated over a 100-m water column depth) was 1.89 ± 1.05 g C m )2 . 3. Significant differences in BP appeared between seasons, especially in the south basin. The range of BP integrated over the 0-100 m layer was 93-735 mg C m )2 day )1 , and overlapped with the range of particulate PP (150-1687 mg C m )2 day )1 ) measured in the same period of time at the same sites. 4. Depth-integrated BP was significantly correlated to particulate PP and chlorophyll-a, and BP in the euphotic layer was on average 25% of PP. 5. These results suggest that HBact contribute substantially to the particulate organic carbon available to consumers in Lake Tanganyika, and that BP may be sustained by phytoplankton-derived organic carbon in the pelagic waters.

Herbivory and bacterivory by phagotrophic protists were estimated in the southern basin of the oligotrophic Lake Tanganyika at different seasons (in the rainy season in February-March 2007 and in the dry season in July-August 2006 and September 2007), using two independent methods: the selective inhibitor technique for assessing community grazing on picocyanobacteria (PCya) and fluorescently labelled bacteria (FLB) and Synechococcus (FLA) to estimate bacterivory and herbivory by phagotrophic nanoflagellates (NF) and ciliates. Protistan grazing impact on both heterotrophic bacteria and PCya was mainly due to NF, which contributed up to 96% of the microbial grazing. There was a clear selection of FLA by protists. PCya represented the main carbon source for both flagellates and ciliates in the mixolimnion, accounting for an average of 83% of the total carbon obtained from the ingestion of picoplanktonic organisms. Protists were the main consumers of particulate primary production (46-74% depending on season). Significant seasonal variation of grazing rates (0.011-0.041 h(-1)) was found, chiefly following variation of PCya production and biomass. Assuming a growth efficiency of 0.4, total protozoan production varied seasonally (189-313 g C m(-2) day(-1)) and was roughly half of particulate phytoplankton production. This study provides evidence that NF and PCya were tightly coupled in Lake Tanganyika and that herbivory by protists may be one of the reasons why this great lake has high productivity. Our results bring support to the idea that microbial herbivory is a major process in oligotrophic freshwater systems.