Phosphorus inputs to Lake Naivasha,Kenya, from its catchment and the trophic state of the lake

The main river supplying Lake Naivasha, Kenya, the Malewa, drains a catchment given over to largely subsistence cultivation and animal husbandry. The lake itself is the focus for an intensive horticultural industry based upon irrigation from the lake. The Malewa, however, is relatively independent of the impact of industry, and so its contribution to eutrophication of the lake was evaluated. Two periods of study, a very wet-dry and a `normal' wet-dry season showed that the river contribution of phosphorus led to a total phosphorus loading of 1.4 g m<sup>–2</sup> lake surface ann<sup>–1</sup> in the very wet period compared to 0.2 in the `normal'. Chlorophyll `a' in the open water of the lake was significantly related to soluble reactive phosphorus. The lake is now eutrophic by normal limnological criteria.     

Organochlorine and organophosphorus pesticide concentrations in water,sediment, and selected organisms in Lake Naivasha (Kenya)

Water, sediment, red swamp crayfish (Procambarus clarkii) and black bass (Micropterus salmoides) from Lake Naivasha were analyzed for selected organochlorine and organophosphorus pesticide residues. The mean p,p'-DDT, o,p'-DDT and p,p'-DDE residue levels recorded in black bass (28.3 (± 30.0), 34.2 (±54.0) and 16.1 (±16.1) g kg^–1, respectively) and crayfish (4.6 (±5.1), 3.2 (±2.8), and 1.4 (±1.1) g kg^–1, respectively), were higher than previously recorded. This indicated recent usage of technical DDT in the lake's catchment. Levels of p,p'-DDT, higher than those of p,p'-DDE further emphasized this. Mean lindane, dieldrin, -endosulfan and aldrin concentrations in black bass were 100.5, 34.6, 21.6 and 16.7 g kg^–1, respectively. The same residues were detected at lower concentrations in crayfish at 2.0, 2.0, 2.0 and 1.9 g kg^–1, respectively. The higher fat content (3.7 ± 2.7% SD) in black bass (compared to 0.6 ± 0.3% in crayfish) accounted for the significantly higher residue concentrations in black bass. Organophosphate pesticides were the most commonly used pesticides in the lake's catchment, but none was detected in any of the samples. The results indicate that there is need for further work to identify sources and fate of pesticide contaminants, as well as to improve monitoring of pesticide use throughout the catchment.     

Phytoplankton community structure and succession in the water column of Lake Naivasha,Kenya: a shallow tropical lake

Lake Naivasha is a shallow freshwater lake in the Rift Valley of Kenya. Since the 1980s, when the lake showed a seasonal shift between diatom and cyanobacterial dominance it has become moderately eutrophic. Its algal assemblage is now dominated by a persistent Aulacoseira italica population both numerically and in terms of contribution to overall primary production. Algal and cyanobacterial counts were used to derive Simpson's diversity, succession rate and total community succession, focusing on the 10 most numerically abundant taxa. 170 species were identified, 43 of which were in common with the 143 found in 1979–80, before the increase in trophic state. Most diatoms are indicators of moderate to high nutrient conditions. There is little horizontal or vertical variation in successional processes throughout the lake and although the absolute abundance of cells varies widely, proportional composition is relatively stable. In Crescent Island lagoon, the only regularly stratified site, hypolimnetic succession rates are lower than those in the epilimnion. Overall, community composition is controlled by mixing (and hence light regime) and nutrient availability. With `endless summer' conditions and full mixing, there is a successional pattern of `muted seasonality' adapted to physical instability and environmental stability.     

The status and future of the Lake Naivasha fishery,Kenya

I describe a laboratory system for investigating the role of light as a proximate cue for diel changes in locomotor activity and vertical location on the substrate of stream macro-invertebrates. The system consisted of computer-controlled halogen lamps positioned over a laboratory stream in which video-recordings were made of Stenonema modestum mayfly nymphs located on the undersides of unglazed tile substrates. Locomotor activity of study organisms in response to light changes were quantified during computer-programmed and reproducible light/dark (LD) cycles. The system provided the flexibility to simulate a variety of light environments so that the separate influences of light intensity and light change on diel activities of individuals and populations could be examined, which is difficult under natural light conditions. As a group, nymphs responded similarly to simulated twilight (light decrease from 7.9 × 10² to 6.9 × 10^–2 W cm^–2 at a constant –1.9 × 10^–3 s^–1 rate of relative light change) and to natural twilight, suggesting that proposed mechanisms of light control of diel activities in nature can be adequately tested in the simulated environment. However, locomotor activity and vertical movements among individual mayflies were highly variable under controlled conditions, suggesting that physiological differences influence their responses to environmental conditions.     

The dynamics and ecology of exotic tropical species in floating plant mats: Lake Naivasha,Kenya

The floating water fern Salvinia molesta has occurred in Lake Naivasha since the early 1960s and during this period has obstructed fishing activities and navigation. In recent years the extent of Salvinia has declined markedly. Since 1988, a second floating weed, Eichhornia crassipes (water hyacinth), has colonised the lake. Salvinia formed large mats around the edges, bays and lagoons and had very few plant or invertebrate species associated with it. Mats of E. crassipes, however, support a greater abundance and variety of animal and plant species, which shows a relationship with mat size. This seems to have facilitated plant succession but without a zonation typical of the classic hydroseral sequence for the lake.     

Towards an understanding of human impact upon the hydrology of Lake Naivasha,Kenya

The growth of a strain of Moina macrocopa (Straus 1820) isolated from an experimental stabilization pond in Marrakesh, was examined at seven concentrations of algae (6.25–6.25 × 10⁵ cells ml^–1 and at 5 different temperatures (15–30 °C)). Feeding conditions influenced the growth rate as well as the maximum size that reached 1.8 mm at 25 °C and at the highest algal concentration (6.25 × 10⁵ cell ml^–1). The life span and number of moltings reached a maximum (17.4 days and 13 moltings) at average nutrient concentrations (6.25 × 10⁵ cell ml^–1). Juvenile stages varied from 1 to 3 and adult ones from 6 to 8. In the temperature interval tested, growth rate increased with temperature while longevity decreased. Temperature had less effect on maximal size than nutrient availability. Population density (but not crowding) influenced longevity and survival but had no effect on growth.     

The aquatic plant communities of the Lake Naivasha wetland,Kenya: pattern,dynamics and conservation

The spatial and temporal patterns of the wetland plant communities at Lake Naivasha over the past decade, and the past five years in particular, are discussed in relation to the major controlling factors. The four communities are:- the emergent swamp, dominated by Cyperus species; the floating raft, dominated by the aliens Salvinia molesta and Eichhornia crassipes; the floating-leaved plants, represented only by Nymphaea caerulea; and the submerged angiosperms, consisting of three species of Potamogetons — P. pectinatus, P. schweinfurthii, P. octandrus — together with Najas pectinata. The major factors affecting their dynamics are:- water level changes which influence agricultural clearance, introduced crayfish Procambarus clarkii, and interactions between communities such as the physical effects of mobile floating rafts. The value of the different communities to the ecological and economic value of the lake is highlighted, and the possible damage from future activities discussed.     

Breeding biology of the Louisiana red swamp crayfish Procambarus clarkii Girard in Lake Naivasha,Kenya

This paper describes the breeding biology of the Louisiana red swamp crayfish (Procambarus clarkii Girard) that has been introduced in a tropical lake, Naivasha. Notable differences in crayfish biology are found between the place of origin in Louisiana and lake Naivasha.In Naivasha sexual maturity in females is reached at a larger size (42 mm carapace length (CL), 76 mm full length (FL)) than in males (40 mm CL, 71 mm FL). The corresponding figures for Chalmette area of Louisiana are 31–32 mm CL for both sexes and at Ben Hur Research Centre, Louisiana, 33 mm CL. The females weigh between 15 and 20 g at the onset of egg deposition. The corresponding Louisiana values are from 5 to 10 g. Crayfish in Naivasha as in Louisiana breed both inside burrows which are 10–70 cm deep, and on the sediment in shallow water at a depth of between 0.5 and 4 m. These observations suggest that reproduction is very much a lake edge activity. The number of breeding individuals in Naivasha varies with the trend of the water surface level. Sexually active females (form I) as judged from secondary sexual characters (appearance of basal hooks at the base of 3rd and 4th walking legs) persist throughout the year at no less than 60% of the adult male population. This is confirmed by the appearance of recently hatched juveniles (3.4 to 6.5 mm CL) in the water throughout the year. Similarly berried females in Naivasha are found throughout the year, though with peaks at irregular intervals. In Louisiana breeding takes place only from August through November with a single peak in October. The fecundity of crayfish in Naivasha is 433 (SE ± 21, N = 133) and there is a positive correlation between body length and egg number.     

Nitrogenase activity in the papyrus swamps of Lake Naivasha,Kenya

Nitrogenase activity (acetylene reduction activity) was found to occur universally in the Cyperus papyrus swamp in Lake Naivasha. Low rates of acetylene reduction activity (0.9–104.9 nmol C₂H₄ g d.wt. roots^-1 h^-1) were associated with excised roots of C. papyrus but higher rates of activity (89.0–280.4 nmol C₂H₄ g d.wt. roots^-1 h^-1) were associated with intact root systems of the plant. It was estimated that nitrogen fixation associated with young roots alone could supply about 26% of the nitrogen requirements of growing papyrus plants. Acetylene reduction activity in the lake bottom sediments was generally low and associated with adjacent papyrus stands. Plate counts of putative aerobic and facultatively anaerobic N₂-fixing bacteria associated with papyrus roots showed the presence of high numbers of diazotrophs (5.4 × 10⁶ CFU g d.wt. roots^-1). Fewer numbers of N₂-fixing bacteria were detected in the sediments (1.9 × 10³-3.2 × 10⁴ CFU g d.wt. sediment^-1).     

Nutrient release from swamp water,Cyperus papyrus L. organs and swamp sediments

Nitrogen and phosphorus releases from swamp water,Cyperus papyrus plant organs and papyrus swamp sediments were investigated over five weeks. The release of nutrients by dissolution of decomposing organic matter was very important for the swamp water under the prevailing anaerobic conditions. Large amounts of nutrients were found in the incubation water with plant organs and sediments within the first 48 hours. Leaching of the nutrients from the plant organs and sediment caused high initial values. Subsequently decomposition of the organic material caused slight changes in the nutrient levels in the water. Probably denitrification played a role in the nitrogen dynamics, while phosphorus was mineralized.     

A Study of a population of Micronecta scutellaris Stal (Hemiptera : Corixidae) in Lake Naivasha,Kenya

Micronecta scutellaris is one of the most abundant macro-invertebrates found in Lake Naivasha, Kenya and forms an important component of the diet of juvenile large mouthed bass.Its distribution in the lake and breeding biology are described and considered in relation to the lake's food chains.     

Plant biomass and nutrient levels of a tropical macrophyte (Cyperus papyrus L.) receiving domestic wastewater

The aboveground biomass and nutrient content ofCyperus papyrus were determined in a small tropical swamp receiving domestic wastewater. It was found that the biomass (4,955 g.m^–2 dry weight) was the highest ever reported for papyrus. The levels of both nitrogen and phosphorus in the plant organs were very high. The nitrogen concentrations of the various plant organs were 4.8% roots, 8.4% rhizomes, 4.5% scales, 4.8% culms, and 6.2% umbels on dry weight basis. As to phosphorus the concentrations were 0.09% roots, 0.11% rhizomes, 0.09% scales, 0.10% culms, and 0.13% umbels. The high biomass and nutrient contents of the plants may have been caused by the high nutrient levels in the surrounding water. Comparison of the nitrogen to phosphorus ratios in the plants to those in the surrounding water showed that the plants stored very high amounts of nitrogen.     

Distribution and abundance of the Louisiana red swamp crayfish Procambarus clarkii Girard at Lake Naivasha,Kenya between 1987 and 1999

The abundance and habitat distribution of Procambarus clarkii, an exotic herbivore at Lake Naivasha, Kenya, was studied between 1987 and 1999. Its growth and morphometrics were similar to other locations in the world where the species have been studied. P. clarkii was found primarily amongst the floating, exotic vegetation where it achieved densities of over 500 m^–2 by the end of the study. It is suggested that these densities became possible because the habitat offers protection against predation and an adequate food supply of bacterial-enriched detritus exists in the absence of green plant material. Both adults and juveniles forage on the lake bed at night.     

Factors affecting abundance and distribution of submerged and floating macrophytes in Lake Naivasha, Kenya

Macrophytes have been shown to perform important ecological roles in Lake Naivasha. Consequently, various studies regarding the impact of biotic factors on the macrophytes have been advanced but related studies on environmental parameters have lagged behind. In an attempt to address this gap, sampling on floating species and submergents was carried out in eight sampling sites in 2003 to investigate how they were influenced by a set of environmental factors. Soil texture (sandy sediments; P  < 0.05, regression coefficient = −0.749) and wind were the most important environmental parameters influencing the distribution and abundance of floating macrophytes. Combination of soil texture and lake-bed slope explained the most (86.3%) variation encountered in the submergents. Continuous translocation of the floating dominant water hyacinth to the western parts by wind has led to displacement of the submergents from those areas. In view of these findings, the maintenance and preservation of the steep Crescent Lake basin whose substratum is dominated by sand thus hosting most submergents remain important, if the whole functional purpose of the macrophytes is to be sustained.     

Aspects of the biodiversity of the rivers in the Lake Naivasha catchment

Deep-water Oculina coral reefs, which are similar in structure and development to deep-water Lophelia reefs, stretch over 167 km (90 nmi) at depths of 70–100 m along the eastern Florida shelf of the United States. These consist of numerous pinnacles and ridges, 3–35 m in height. Coral growth rates average 16.1 mm yr^–1 and biodiversity is very rich. Extensive areas of Oculina rubble may be due to human impacts (e.g. fish trawling and dredging, anchoring, bottom longlines) and natural processes such as bioerosion and episodic die-off. Early in the 1970s, the reefs were teeming with fish. By the early 1990s, both commercial and recreational fisheries, including scallop, shrimp, grouper, snapper and amberjack, had taken a toll on the reefs and especially on populations of grouper and snapper. A 315 km² (92 nmi²) area was designated the Oculina Habitat of Particular Concern (HAPC) in 1984, prohibiting trawling, dredging, bottom longlines and anchoring, and legislation was enacted in 2000 for expansion of the Oculina HAPC to 1029 km² (300 nmi²). The United States Coast Guard has been charged with surveillance and enforcement of the ban on bottom fishing and trawling. The primary difficulties in protecting these reefs and other deep-water Marine Protected Areas are their remoteness and time required to engage an enforcement vessel. Education regarding the nature and importance of these rich resources is important for better self regulation and surveillance by the fishing community. Only by bringing deep-water reefs to the public, the fishing community, and enforcement agencies, through video, photos, and education will there be better understanding and acceptance for the need of protection for these unseen resources. This paper reviews the current knowledge on the deep-water Oculina reefs, including the biology, geology, human impacts, and history of conservation and management.     

The effect of tubificids and chironomids on particle redistribution of lake sediment

The effects of chironomid larvae,Chironomus plumosus, and tubificid worms,Limnodrilus spp., on particle redistribution in lake sediment were investigated experimentally using pots containing sediments obtained from Lake Suwa, Japan. The chironomids and tubificids increased the water content of surface sediment. The chironomid larvae had no effect on particle size distribution, while tubificids continuously accumulated small particles on the surface sediment through their selective feeding activity. Particles larger than 0.125 mm were buried at a sediment depth of 6 cm. In Lake Suwa, long diatom frustules, large plant debris and blue-green algal flocs were found to accumulate in the deeper layer of the lake sediment inhabited by tubificids at high density.     

Massive fish kills within the Nyanza Gulf of Lake Victoria,Kenya

The morphometry and hydrology of the Nyanza Gulf of Lake Victoria can be greatly affected by violent storms. This can result in a condition in which nutrient-rich bottom mud is mixed with the sediment-laden runoff water from neighbouring marshes and rivers. This situation can lead to massive fish kills. It is such an event in 1984 which will be described that caused the sudden death of over 400 000 fish weighing over 2400 tonnes.The unusually low precipitation in the catchment area during that year resulted in a drop in the lake level from 12.4 to 11.8 m. This low water level, combined with a severe storm created the conditions which caused this large fish kill. The species affected were Lates niloticus and Oreochromis niloticus. Mortality was attributed to several factors which included high levels of suspended material in the water column (detritus and algae) which clogged the gills of the fish, low dissolved oxygen, low pH, and high concentrations of algae.     

Effects of the burrowing mayfly,Hexagenia, on nitrogen and sulfur fractions in lake sediment microcosms

Effects of the burrowing mayfly, Hexagenia, on nitrogen and sulfur fractions of sediment, and overlying water were determined. Laboratory microcosms were used to reproduce the benthic environment. The activities of Hexagenia increased sediment Eh (1.98 ± 0.486 (22) mV · day ⁻¹), and decreased pH in sediment (−0.007 ± 0.001 (22) day ⁻¹) and overlying water(-0.024 ± 0.004 (10) day⁻¹). In the control, Eh decreased and pH did not change. The presence of Hexagenia also markedly increased ammonia in sediment (5.46 ± 0.14 (22) ppm N · day⁻¹) and overlying water (0.792 ± 0.154 (10) ppm N · day⁻¹), while the control did not change. In addition, the sulfate fraction of sediment (0.177 ± 0.006 (17)% dry mass) and water (50.0 ± 4.9 (5) mg · I⁻¹) in microcosms with Hexagenia was greater than that of the control (0.151 ± 0.005 (16)% dry mass; 14.7 ± 1.71 (3) mg · 1⁻¹) at the termination of the experiment. Hexagenia may also stimulate the mineralization of carbon-bonded sulfur. The general role of Hexagenia in altering sediment chemistry is discussed.     

Aquatic biodiversity and saline lakes: Lake Bogoria National Reserve,Kenya

Lake Bogoria, in the Rift Valley of Kenya is an extreme saline lake (conductivity 40–80 mS cm^–1, alkalinity 1500 m equ l^–1). It is hydrologically more stable than the other, endorheic lakes in Kenya, because it is deep – maximum depth at present just over 10 m in an area of 3000 ha – and so does not have periods when it is dry. It is ecologically simple, with only one species dominating the phytoplankton – the cyanobacterium `spirulina', Arthrospira fusiformis. Its biomass and productivity were very high – biomass between 38 and 365 g l<sup>–1</sup> chlorophyll `a' and 3.4–21 × 10³ coils ml^–1 and net production between 0.24 and 1 gm C m³ h, the latter in a narrow zone of less than a metre. There were no macro-zooplankton in the plankton and the only grazer of A. fusiformis was the lesser flamingo, Phoeniconaias minor,which occurred irregularly in very high concentrations (in excess of 1 × 10⁶). Detritivory in the benthos was effected by a single chironomid species, Paratendipes sp., at a maximum density of 4 × 10⁴ m^–2. The mean daily emergence of adult chironomids was estimated to be 1 × 10³ m^–2, the maximum 3. There was no littoral plant community within the lake but 44 dicotyledonous and 31 monocotyledonous plant species in the drawn-down zone and adjacent to it. A diverse draw-down terrestrial invertebrate fauna, only superficially described here, processed the flamingo feathers and carcasses, with other detritus such as chironomid pupal exuviae and decaying A. fusiformis scum. About 50 bird species depended upon the chironomids, either as they emerged through the water column as flying adults or later on the shoreline as floating pupal exuvia and dead adults. The lake has high conservation value because of three bird species in particular – lesser flamingo, Cape teal and black-necked grebe. The former provides real economic value in a region otherwise impoverished, because of the spectacle of tens of thousands of flamingos set against the landscape of hot springs and fumaroles at the lake edge, which draws 15000 visitors per annum. P. minor has experienced three periods during the past ten years when major mortalities have occurred, the last of which killed 700 birds day^–1. This could have involved as many as 200000 birds (about 1/5th of the maximum population at this lake) if mortality was at a constant rate for the nine months it was observed. Causes of mortality have been suggested as avian tuberculosis, poisoning from cyanobacterial toxins or from heavy metal contamination at Lake Nakuru, but it is still not yet clear what contribution each makes to the problem.