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.     

Lakes of south-western Uganda

The vegetation of Lake Bunyonyi, a high altitude lake in Kigezi, S.W. Uganda, is described. The lake had two distinct plant communities. In the shallow sheltered bays and inflows extensive floating swamps of Cyperus papyrus L. and Cladium janiaicen. se Cranz. existed. On the steep, exposed shores there was only a narrow band of vegetation, dominated by firmly rooted Phragmites australis (Cav.) Steud., usually with papyrus behind. Floating-leaved Potamogeton and Nymphaea sometimes occurred outside the emergent species, and submerged species, including Ceratophyllum demersum L. grew down to 8 m. The pattern of vegetation, and the contribution of floating islands to the distribution of plants in the lake is discussed. Susceptibility to fragmentation, and distribution by wind are thought to be major factors governing the pattern. Oxygen proflies measured at the fringe of swamps indicated that the swamps did not cause appreciable deoxygenation of the water.     

Floating mats: their occurrence and influence on shoreline distribution of emergent vegetation

1. A study was conducted on the northern shore of Lake Victoria (Uganda) to determine the factors controlling the occurrence of floating root mats and the influence of the floating mats on the distribution of emergent vegetation. 2. Environmental conditions within 78 bays in the study area were characterised using bay size, wave exposure, water depth, littoral slope, sediment characteristics and water level fluctuations. Emergent plants that form floating root mats occur along the shores of these bays. The way in which commonly occurring shoreline vegetation was distributed across a wave‐exposure gradient was compared with their distribution across a water level fluctuation gradient. 3. Results suggested that wind–wave action and water level fluctuations are important factors determining the occurrence of floating mats. Mat‐forming plants occur in the most sheltered locations along the shore and in waterbodies with modest water level fluctuations. 4. The ability to form mats facilitated the lakeward expansion of emergent plants. Plants forming floating root mats had a larger depth range than non‐mat forming plants. 5. The initiation mechanisms for the floating mats of emergent vegetation in Lake Victoria appear to be: (i) invasion of mats of free‐floating plants by emergent vegetation; and (ii) detachment of emergent plants from the lake bed following flooding. 6. The formation of floating mats comes with a cost and benefit to emergent plants. The cost is increased vulnerability to damage by water level fluctuations or wind–wave action, leading to reduced horizontal distribution. The benefit is that deep flooding is avoided, thus increasing vertical distribution. The net effect may be to lead to dominance of mat‐forming plants in low‐energy environments and non‐mat‐forming plants in high‐energy environments.     

Phylogenetic taxonomy of the marine mite genus Rhombognathides (Acari: Halacaridae: Rhombognathinae)

This study examined the impacts of the alien waterweed, water hyacinth, on the abundance and diversity of aquatic macroinvertebrates in the littoral areas of northern Lake Victoria in Uganda. The weed had undergone explosive growth on the lake causing serious disruption to people, the economy and the ecosystem. This study was confined to impact of the weed in the littoral zone, not to the large floating mats of vegetation which float across the lake and clog large areas of shoreline.The littoral area studied comprised of fringing mats of Eichhornia crassipes (Mart) Solms (water hyacinth) to the lakeward of Cyperus papyrus; water hyacinth mats undergoing colonisation by Vossia cuspidata (Roxb.) Griff.; and a typical Cyperus papyrus L shore with no outer floating mat of water hyacinth. Numerical abundance (Nos. m^–2) and diversity (No. of taxa) of macroinvertebrates recovered from pure Eichhornia crassipes and the Eichhornia-Vossia succession increased from the fringe of the Cyperus papyrus towards the open water. In the typical Cyperus papyrus fringe, in the absence of water hyacinth, abundance was highest at the papyrus/open water interface and dropped off sharply towards open water. The Shannon–Weaver diversity index (H) of macroinvertebrates decreased progressively from pure Eichhornia crassipes stands, to Vossia/Eichhornia beds and Cyperus papyrus stands (H=0.56, 0.54 and 0.34, respectively) but were not significantly different. Dissolved oxygen decreased from open water into vegetation where it approached anoxia. Water hyacinth appeared to enhance the abundance and diversity of aquatic macroinvertebrates at the interface with the open water. The impoverished abundance and diversity of the macroinvertebrates deeper into the vegetation mats suggested negative environmental impacts of the water hyacinth when the fringe is too wide. Further research is recommended to establish the optimum width of the fringe of stationery water hyacinth that promotes maximum abundance and diversity of aquatic macroinvertebrates and, possibly, of other aquatic life. Since this study in 1997, there has been a dramatic decrease in Eichhornia infestations and by June 2000 it appeared largely to exist only as fringing vegetation in bays and inlets.     

Divergence in aerobic scope and thermal tolerance is related to local thermal regime in two populations of introduced Nile perch (Lates niloticus)

We tested whether thermal tolerance and aerobic performance differed between two populations of Nile perch (Lates niloticus) originating from the same source population six decades after their introduction into two lakes in the Lake Victoria basin in East Africa. We used short-term acclimation of juvenile fish to a range of temperatures from ambient to +6°C, and performed critical thermal maximum (CT_max) and respirometry tests to measure upper thermal tolerance, resting and maximum metabolic rates, and aerobic scope (AS). Across acclimation temperatures, Nile perch from the cooler lake (Lake Nabugabo, Uganda) tended to have lower thermal tolerance (i.e., CT_max) and lower aerobic performance (i.e., AS) than Nile perch from the warmer waters of Lake Victoria (Bugonga region, Uganda). Effects of temperature acclimation were more pronounced in the Lake Victoria population, with the Lake Nabugabo fish showing less thermal plasticity in most metabolic traits. Our results suggest phenotypic divergence in thermal tolerance between these two introduced populations in a direction consistent with an adaptive response to local thermal regimes.     

Functioning and dynamics of wetland vegetation of Lake Victoria: an overview

The aquatic macrophytic vegetation constituting the wetlands situated along the coast of Lake Victoria provides valuable services to both local and regional communities as well as an important ecological function through the transition between terrestrial and aquatic ecosystems. The wetland vegetation is typically rooted in the substrate on the landward side of the lake, but forms a floating mat towards the middle of the wetland and at the wetland/lake interface. Cyperus papyrus and Miscanthidium violaceum vegetation typically dominate the permanently inundated wetland areas along most of the shores of Lake Victoria. Due to the prevailing climatic and hydrological catchment conditions, these macrophytic plants (papyrus in particular) tend to exhibit high net productivity and nutrient uptake which strongly influences both wetland status and lake water quality. In addition, these wetlands provide important economic livelihoods for the local populations. The integrity and physical structure of these wetlands strongly influences their associated mass transport mechanisms (water, nutrients and carbon) and ecosystem processes. Wetland degradation in Africa is an increasing problem, as these ecosystems are relied upon to attenuate industrial, urban and agricultural pollution and supply numerous services and resources. In an integrated project focused on the wetlands of Lake Victoria, the ecological and economic aspects of littoral wetlands were examined and new instruments developed for their sustainable management.     

Development of macrophytic vegetation in the Agmon Wetland of Israel by spontaneous colonization and reintroduction

The draining of the Lake Hula and swamps, northern Israel, during the late 1950s resulted in the loss of a very diverse and rare ecosystem. Oxidation of the peat soil resulted in ground surface subsidence, while heavy autumn winds have eroded the dry peat. Moreover, agriculture on the peat soils is restricted, because of a nitrate surplus. Predictions that the sinking would continue and that more areas would go out of agricultural production led authorities to re-flood a portion of the Hula Valley in 1994. The aim of the present study was to monitor the spontaneous establishment of vegetation in the re-flooded area, the Agmon wetlands, and to reestablish some of the major plant species lost from the valley when Lake Hula was drained. Within the first two years, 74 plant species colonized the wetland spontaneously. Five out of 11 species designated for reintroduction were successfully established. Cyperus papyrus and Cynodon dactylon demonstrated sustainable potential for lake-shore stabilization. Cyperus papyrus was reintroduced from seedlings and rapidly became the dominant riparian species, while Cynodon dactylon established spontaneously. Re-introduced Nymphaea alba clones were established only in enclosures protected from grazing by the semi-aquatic mammal Myocastor coypu. Nuphar lutea and Iris pseudacorus showed better resistance to grazing. These results demonstrate a high potential for successful re-establishment of much of the original Hula swamp macrophytic vegetation by either spontaneous colonization by extant species from the surrounding areas or by introduction of locally extinct species. As such, there is a good chance that the associated faunal components of the former Lake Hula and swamps that have returned to the region since the Hula rehabilitation project commenced will continue to flourish.     

Habitat associations of papyrus specialist birds at three papyrus swamps in western Kenya

Birds of papyrus swamps have not been adequately studied in Kenya, and little is known about their ecology and habitat associations. Using fixed‐radius point counts and playbacks, we counted papyrus specialist birds and evaluated papyrus physical characteristics and levels of disturbance at a series of sample stations at three papyrus swamps of Dunga, Koguta and Kusa in the Kenyan sector of Lake Victoria. Papyrus height and density were significantly correlated across all sites but negatively correlated with levels of disturbance. Standardized point counts of swamp birds showed the papyrus gonolek Laniarius mufumbiri and Carruthers's cisticola Cisticola carruthersi to be the most abundant papyrus specialists across sites. Only Carruthers's cisticola numbers differed between sites. Overall, papyrus cover was the best predictor of the presence and abundance of all papyrus specialist birds, and significantly predicted the numbers of papyrus gonolek and white‐winged warbler Bradypterus carpalis.     

Physiological refugia: swamps,hypoxia tolerance and maintenance of fish diversity in the Lake Victoria region

In Lake Nabugabo, Uganda, a satellite of Lake Victoria, approximately 50% of the indigenous fishes disappeared from the open waters subsequent to the establishment of the introduced predatory Nile perch, Lates niloticus. This pattern is similar to the faunal loss experienced in the much larger Lake Victoria. Several of these species persisted in wetland refugia (e.g. ecotonal wetlands, swamp lagoons); however, deep swamp refugia (habitats lying well within the dense interior of fringing wetlands), are available only to a subset of the basin fauna with extreme tolerance to hypoxia. Although air-breathers are common in deep swamp refugia; we also documented a surprisingly high richness and abundance of non-air-breathing fishes. We describe several mechanisms that may facilitate survival in deep swamp refugia including high hemoglobin concentration, high hematocrit, large gill surface area and a low critical oxygen tension (P(c)). In addition, swamp-dwelling fishes showed lower PO(2) thresholds for onset of aquatic surface respiration than the lake-dwelling fishes. This suggests higher tolerance to hypoxia in the swamp fishes because they are able to withstand a lower oxygen tension before approaching the surface. We suggest that physiological refugia may be important in modulating the impact of Nile perch and indigenous fishes in the Lake Nabugabo region; this highlights the need to evaluate relative tolerance of introduced predators and indigenous prey to environmental stressors.     

Macrophyte species composition,distribution and diversity in relation to some physicochemical factors in the littoral zone of Lake Ziway,Ethiopia

The species composition, distribution and diversity of macrophytes in Lake Ziway were determined at the end of dry and wet seasons along nine littoral sites during 2010 using a belt transect method. Some physicochemical parameters were also measured. Fourteen macrophyte species were identified with low species diversity for the lake (H' value of 1.805). The macrophyte species composition of the lake has undergone many changes during the last few decades. Arundo donax, which was never reported earlier, attained the highest relative density (30.7%) and frequency (30.5%) in this study, followed by Echinochloa colona, Potamogeton schweinfurthii, Cyperus articulatus, Typha latifolia and Cyperus papyrus. The average densities of A. donax, C. articulatus, E. colona and Pistia stratiotes were positively correlated with nutrients, whereas the density of the other species had negative correlation with physicochemical variables (CCA). Generally, the present macrophyte composition and abundance indicates healthy ecological condition of the lake. But if nutrient levels continue to increase and water levels continue to decline, we expect further changes in macrophyte composition and especially a shift towards invasive floating species.     

Impacts of the C4 sedge Cyperus papyrus L. on carbon and water fluxes in an African wetland

Fluxes of CO₂ and H₂O vapour were measured by eddy covariance from a stand of the C₄ emergent sedge Cyperus papyrus (papyrus), which formed a fringing swamp on the north-west shore of Lake Naivasha, Kenya. The fluxes of CO₂ and H₂O vapour between the papyrus swamp and the atmosphere were large but variable, depending on the hydrology of the wetland system and the condition of the vegetation. These measurements, combined with simulation modelling of annual fluxes of CO₂, show that papyrus swamps have the potential to sequester large amounts of the carbon (1.6 kg C m^–2 y^–1) when detritus accumulates under water in anaerobic conditions, but they are a net source of carbon release to the atmosphere (1.0 kg C m^–2 y^–1) when water levels fall to expose detritus and rhizomes to aerobic conditions. Evapotranspiration from papyrus swamps (E) was frequently lower than evaporation from open water surfaces (E _o) and plant factors have a strong influence on the flux of water to the atmosphere. For the period of measurement E/E_o was 0.36.     

The role of the littoral zone in the functioning of a shallow tropical lake ecosystem

Lake Chilwa (Malawi) is a shallow tropical lake surrounded by an extensive zone of littoral swamp mainly composed of Typha domingensis. The data produced show that the physical and chemical conditions are different and more varied in the littoral region when compared with the open lake. Primary production in the littoral is confined almost entirely to Typha domingensis, which also acts as a nutrient pump by absorbing nutrients from the soils and eventually releasing them into the lake waters. Long term and short term exchanges in organic and inorganic materials between the littoral swamps and the open lake are described. These, as well as available evidence on the distribution, reproductive habits and diet of certain faunal groups, indicate that the littoral region of Lake Chilwa has an important regulating influence on the functioning of the whole lake.     

Primary productivity of papyrus (Cyperus papyrus) in a tropical swamp; Lake Naivasha,Kenya

Papyrus (Cyperus papyrus) standing biomass and the primary productivity of undisturbed and previously harvested areas of papyrus was measured in Lake Naivasha swamp, Kenya. Papyrus culm density in undisturbed swamp was estimated to be 13·1±1·9 culms m⁻² and aerial biomass was 3602 g m⁻². In undisturbed swamp the aerial productivity was 14·1 g m⁻² day⁻¹ while the previously harvested swamp reached a peak of 21·0 g m⁻² after 6 months. The annual aerial production rate of papyrus in Lake Naivasha was estimated to be 5150 g m⁻² year⁻¹. To sustain yields of regularly harvested papyrus swamps, the harvest intervals should exceed 1 year.     

The vegetation of Nyumba ya Mungu reservoir, Tanzania

A study was made of the aquatic macrophyte ecology of Nyumba ya Mungu; a nine year old man-made lake in northern Tanzania. The vegetation is described with the aid of percentage cover histograms and standing crop estimates. Concurrent physical, chemical and biological studies helped in ecological interpretations. Distinct vegetation communities were found in the deltaic swamps of the two major inflow rivers and it is concluded that this was due to water chemical differences. The water of the more alkaline River Kikuletwa appeared to influence the greater part of the perimeter swamps as the species composition largely coincided with that reported for higher pH, higher conductivity African lakes. Typha domingensis swamp was predominant and extensive, whilst Cyperus alopecuroides and Paspalidium geminatum were also common emergent species. Floating and submerged plants were relatively unimportant in the lake with respect to cover. The major zonations of species appeared to be a function of water depth, while chemistry seemed the main factor controlling vegetation pattern. In conclusion the significance of interrelationships between the macrophytes and other aquatic organisms is discussed.     

Permanent swamp vegetation of the Upper Nile

This paper describes vegetation in the nothern Sudd area of the Sudan. A visit by boat was made in April 1978 along the White Nile from Malakal to Lake No; to the inflow regions of the Bahr el Jebel, the Bahr el Zeraf and to the Bahr el Ghazal. Only the vegetation along the fringe of the rivers was observed as plants were tall and dense and it was not possible to penetrate into the inner swamps.The outer edge of the vegetation was fringed by extensive mats of free-floating Eichhornia crassipes which was easily dislodged and formed floating islands to be carried downstream. The main rivers, excluding the Bahr el Ghazal, were too deep and turbid to support beds of submerged species but plants such as Ceratophyllum demersum could be found amongst floating plants. Vossia cuspidata usually produced a broad zone to the landward of Eichhornia. It dominated the shallow water with a floating raft of stems growing out from the shore over the water surface. Massive stands of Cyperus papyrus and/or Phragmites karka, over 5 metres tall, dominated the vegetation behind and sometimes excluded the Vossia zone. The papyrus could form dense, floating rafts of rhizomes at the water's edge, but Phragmites always remained firmly rooted to the bottom. Floating papyrus and Vossia would become detached from the main beds and drifted downstream with Eichhornia.Creepers were common amongst papyrus but less common amongst Phragmites where the vegetation was generally more dense. Large stands of Typa domingensis were observed behind the fringe vegetation.The vegetation of the Bahr el Ghazal, in marked contrast to the rest, was stunted, and reasons for this are suggested. Species diversity was greater and the less turbid waters allowed considerable development of floating-leaved and submerged species.Estimates of fresh weights and heights of dominant vegetation types were attained but methods were very laborious and alternative ways have been suggested. The fresh weight of Eichhornia from a one metre square quadrat in the White Nile was around 70 kg. The shoots of Phragmites weighed 18.5 kg and Cyperus papyrus, 11.0 kg m^–2.     

Nutrient status and retention in pristine and disturbed wetlands in Uganda: management implications

Wetlands in Uganda experience different forms of human pressure ranging from drainage for agriculture and industrial development to over harvesting of wetland products. In order to develop sustainable management tools for wetland ecosystems in Uganda and the Lake Victoria Region, water quality analyses were carried out in a rural undisturbed (pristine) wetland (Nabugabo wetland in Masaka) and two urban wetlands that are experiencing human and urban development pressure (the Nakivubo wetland in Kampala and Kirinya wetland in Jinja). The former wetland forms the main inflow into Lake Nabugabo while the other two border the northern shore of Lake Victoria, Uganda. Nabugabo wetland buffers Lake Nabugabo against surface runoff from the catchment, while Nakivubo and Kirinya wetlands provides a water treatment function for wastewater from Kampala City and Jinja town respectively, in addition to buffering Lake Victoria against surface runoff. Water quality was assessed in all the wetland sites, and in addition nutrient content and storage was investigated in the main plant species (papyrus, Phragmites, Miscanthidium and cocoyam) in Nakivubo and Kirinya wetlands. A pilot experiment was also carried out to assess the wastewater treatment potential of both the papyrus vegetation and an important agricultural crop Colocasia esculenta (cocoyam). Low electrical conductivity, ammonium–nitrogen and ortho-phosphate concentrations were recorded at the inflow into Nabugabo wetland (41.5 μS/cm; 0.91 mg/l and 0.42 mg/l respectively) compared to the Nakivubo and Kirinya wetlands (335 μS/cm; 31.68 mg/l and 2.83 mg/l and 502 μS/cm; 10 mg/l and 1.87 mg/l respectively). The papyrus vegetation had higher biomass in Nakivubo and Kirinya wetlands (6.7 kg DW m⁻²; 7.2 kg DW m⁻² respectively), followed by Phragmites (6.5, 6.7), cocoyams (6.4, 6.6) and Miscanthidium (4.0, 4.2). The papyrus vegetation also exhibited a higher wastewater treatment potential than the agricultural crop (cocoyam) during the pilot experiment (maximum removal degree of ammonium–nitrogen being 95% and 67% for papyrus and yams). It was concluded that urbanisation pressure reduces natural wetland functioning either through the discharge of wastewater effluent or the degradation of natural wetland vegetation. It is recommended that wetland vegetation be restored to enhance wetland ecosystem functioning and for wetlands that are not yet under agricultural pressure, efforts should be made to halt any future encroachment.     

Biological diversity of the Yala Swamp lakes, with special emphasis on fish species composition, in relation to changes in the Lake Victoria Basin (Kenya): threats and conservation measures

During the second half of the last century, the Lake Victoria ecosystem has undergone drastic ecological changes. Most notable has been the decline in the populations of many endemic cichlid fishes. The lake has lost nearly 200 haplochromines and one tilapiine, Oreochromis esculentus. The above changes have been attributed to effects of species stocking and, in particular, from predation pressure by the introduced Nile perch, Lates niloticus. Other factors that have led to the decline of the endemic species include intensive non-selective fishing, extreme changes in the drainage basin, increased eutrophication, and the invasion of the lake by the water hyacinth, Eichhornia crassipes. However, the remnants of some species that had disappeared from Lake Victoria occur abundantly in the Yala Swamp lakes (Kanyaboli, Sare and Namboyo). This paper discusses the biodiversity of the swamp and the three lakes and gives suggestions for their conservation.     

Species richness in boreal swamp forests of SE Norway: The role of surface microtopography

Question: What is the relative importance of environmental gradients and surface microtopography (variation in vertical level within sampling units) for fine‐scale plant species richness in Picea abies swamp forests? Location: 11 swamp forests in SE Norway. Methods: We recorded species richness (number of species of vascular plants, mosses, Sphagnum and hepatics), depth to water table, soil base status and vertical range (microtopographic relief) in 2400 microplots, (each 1/16 m²), in 150 1‐m² plots in the 11 swamp forests. Relationships between species richness and environmental predictors were modelled by GLMM. Results: Moss and hepatic species richness increased with increasing microtopographic relief, most strongly for wet acid sites, in which similar trends were also found for Sphagnum. Relief was a poor predictor of vascular plant species richness. Conclusions: Microtopographic relief is a good predictor of local species richness in Picea abies swamp forests, partly because larger vertical variability means higher within‐plot habitat diversity with respect to the wet‐dry gradient, and partly because qualitatively new microhabitats associated with steep slopes are added in drier sites. The relationship between species richness and microtopographic relief is context dependent, differing in complex ways among species groups and among sites with different environmental conditions.     

The macrophytic vegetation of the draw-down area of Lake Kainji, Nigeria

Recent publications on the vegetation of Lake Kainji, Nigeria, give the impression that Echinochloa pyramidalis is the dominant macrophyte species of the lake. However, our experience of working on the lake since 1979 suggests a different floristic composition of the macrophyte vegetation of the lake. The most frequent macrophyte in Lake Kainji is Echinochloa stagnina. At all sites investigated, emergent macrophytes are more important than floating macrophytes. There is a possible successional shift from an Echinochloa pyramidalis sere to an Echinochloa stagnina stage. This argument does not, however, preclude the possibility of a mis-identification by early workers.     

Wetland Restoration at a Former Nike Missile Base in the Great Lakes Basin

Restoration of an abandoned 16-ha Nike missile base site located on a former wetland in the lower Detroit River (Michigan, U.S.A.) was investigated with an emphasis on wetland restoration. The site included a 2.7-ha abandoned missile base, a 1.3-ha lake, 10.4 ha of emergent and submersed wetlands, and 1.8 ha of uplands. Aquatic beds in the shallow bay connected to the river supported floating leaved and submersed aquatics including Nymphaea tiiberosa, Vallisneria antericana, Elodea canadensis, and Heteranthera dubia, with mats of green, filamentous algae. A 10-ha diked wetland adjacent to the site was dominated by Typha spp., Salix spp., Nymphaea tuberosa, Myriophyllum spicatum, Elodea canadensis, Chara sp., and Juncus effusus. Restoration objectives included a nature preserve, an outdoor recreation area, an experimental wetland complex, and an environmental education center. Ten alternative designs were suggested, including four with wetlands open to the bay, three with diked wetlands, and the rest involving island construction, a reconstructed upland site, or a wetland research facility. Alternatives were evaluated for their contribution to local ecology, research, education and wildlife, ease of maintenance, and probability of success. Construction of a wetland resembling conditions before construction of the base was recommended for its low maintenance and opportunity for research on non diked wetland design and construction in protected bays in the Laurentian Great Lakes. Recreational and educational opportunities were also recommended as part of the site restoration.