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.     

Interdemic variation in the foraging ecology of the African cyprinid, Barbus neumayeri

In the African cyprinid, Barbus neumayeri, populations from hypoxic waters have larger gills than populations from well-oxygenated streams. Differences in trophic morphology and feeding performance between these populations suggest a reduction in feeding efficiency in large-gilled fish that may reflect spatial constraints of the gills. However, this variation may also reflect interdemic variation in diets. In this study, we describe patterns of variation in diet, gut morphology, and prey availability for populations of B. neumayeri from swamp (low-oxygen) and stream (high-oxygen) sites in Kibale National Park, Uganda. Our results indicate that B. neumayeri are omnivorous, feeding primarily on benthic prey items; however, diets differed among swamp and stream sites for certain prey types. The observed dietary differences do not provide direct support for predictions based on variation in trophic musculature; hard-bodied prey were more common in low-oxygen sites. Prey availability also differed among sites; in particular, insect abundance and richness was generally lower in the swamp sites. Gut length was longer in one of the four populations, but did not conform to expectations based on diet differences. Condition and growth rates did not differ between populations from hypoxic and well-oxygenated sites, despite observed differences in prey availability and diet, suggesting that B. neumayeri may be distributed in a way that equalizes fitness among populations in different habitats.     

Papyrus swamps, hypoxia, and faunal diversification: variation among populations of Barbus neumayeri

To test whether patches of papyrus swamp contribute to diversification of populations of non-air-breathing fishes, the gill morphology of Barbus neumayeri was compared between a papyrus swamp and several tributaries which differed in oxygen regime. Total gill filament length differed among sites and was negatively related to dissolved oxygen availability, supporting strong selection pressure for low-oxygen tolerance in the swamp interior. Among recaptures of marked B. neumayeri over a 4·5-year period among the focal swamp and connected stream and river sites, 93% of fish were recovered at the site of capture. Some of the individuals that moved crossed physicochemical gradients and traversed long distances within the swamp/stream system. This movement rate would theoretically be sufficient to homogenize gene frequencies among populations. However, randomly amplified polymorphic DNA (RAPD) markers indicated significant genetic differentiation among sites and no relationship between genetic differences and geographical distances among sites suggesting habitat-specific selection pressures on dispersers, rather than insufficient dispersal.     

Effects of a cyperus papyrus L. swamp on domestic waste water

The effectiveness of a papyrus swamp for the removal of nutrients from domestic waste water was investigated in a small tropical swamp in Kenya. The study was conducted in a man-made impoundment transformed into a swamp which receives sewage effluents discharged into a stream.Spatial and temporal distribution of nutrients were determined in the swamp from October 1980 to August 1981. They included the measurement of nitrogen and phosphorus, together with temperature, pH, conductivity and dissolved oxygen.A comparison between the water quality characteristics of the input versus the output from the swamp indicated significant decreases in the mean temperature and conductivity by 20 and 23%, respectively. Dissolved oxygen was reduced by 85%, ammonium by 77% and orthophosphate by 80%.The results obtained indicate that papyrus swamps are efficient in nutrient removal for the purpose of domestic waste water renovation.     

Seasonal Influences on the Ecology of Testate Amoebae (Protozoa) in a Small <Emphasis Type="Italic">Sphagnum</Emphasis> peatland in Southern Ontario,Canada

Testate amoebae (Protozoa) were studied in spring, summer, and fall from the same microhabitats in a small Sphagnum-dominated peatland in southern Ontario, Canada. A total of 32 sampling stations were established in two wetland plant communities, 19 in an open Ericaceae low-shrub community and 13 in a closed Picea mariana and Larix laricina swamp community. Sphagnum was collected in each station for analysis of testate amoebae and measurement of soil water content parameters and water table depth in May, August, and October 2001. pH and dissolved oxygen of the groundwater under the Sphagnum were measured also. A total of 52 taxa including the rotifer, Habrotrocha angusticollis, were identified. Soil water content and water table variables emerged as the primary factors separating testate amoebae between the open bog/fen community and swamp community. Testate amoebae in the open bog/fen community showed a clear separation between the May sampling period and the August and October sampling periods. Sampling stations in May had much higher water table and were wetter than those in August and October. Conversely, testate amoebae in the swamp community did not show a clear difference between sampling periods. Soil moisture and water tables appear to be more constant in the swamp communities. Biological factors or other microscale environmental factors may need to be considered to explain seasonal changes in testate amoebae. A greater understanding of relationships between testate amoebae and microenvironmental factors is necessary to track seasonality in testate amoebae distributions.     

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.     

Papyrus wetlands a lunar-modulated refuge for aquatic fauna

A field study was undertaken of the use of papyrus (Cyperus papyrus) wetlands fringing Rubondo Island, Lake Victoria, Tanzania, as a refuge from predators by freshwater shrimps (Caridina nilotica) and tilapia (Oreochromis niloticus) larvae. During the full moon nights, fish larvae took refuge more than 5 m in the papyrus and did not venture out into the lake, while they ventured into fringing lake waters during moonless nights. The freshwater shrimps and lake flies (Chaoborus sp.) did not migrate with the moon. This fauna took refuge in the papyrus wetlands even though the dissolved oxygen concentration was as low as 0.4 ppm (<5% of saturation).     

Fish communities in the eastern,seasonal-floodplain of the Sudd,Southern Sudan

Annual flooding of the permanent Sudd swamps innundates large areas of Oryza longistaminata grasslands. Fish migration into a southeast section of the seasonal floodplain was led by Clarias gariepinus, Polypterus senegalus and several small species which were intent on spawning. Channa obscura and juveniles of Oreochromis niloticus were prominent among later entrants. From a total of 23 species caught in seasonal habitats, 7 contributed over 97% of total estimated numerical and biomass densities. The latter ranged from 435–580 kg ha^–1. In the 1982–83 flood-season entry to the floodplain was essentially restricted to species tolerant of oxygen-deficient waters and thereby able to penetrate the papyrus and Typha swamps of the permanent wetland. Migrants became stranded in temporary pools and subject to heavy cropping by birds.     

Water circulation and fish larvae recruitment in papyrus wetlands,Rubondo Island,Lake Victoria

A field study was undertaken of the hydrodynamics, water quality and adult and larval fish abundance in papyrus wetlands and surrounding coastal waters at Rubondo Island, Lake Victoria. Because they were exposed to the prevailing wind, the two bays facing east had small (0.01 km²) wetlands and were well flushed, with minimal accumulation of organic detritus and little oxygen depletion. Because it faced west, the third bay, Mlaga Bay, was sheltered and poorly flushed; organic matter accumulated in a large (1.41 km²) papyrus wetland. The rises and falls of the lake level at decadal time scales, may, by drowning or drying out wetlands, contribute significantly to storing organic matter as detritus. Vegetation decay in the wetlands, as well as primary production, resulted in large diurnal fluctuations of pH and dissolved oxygen concentration. Shading in daytime and cooling at night, cooled the wetlands water sufficiently to generate a baroclinic circulation whereby cold, wetland water sank and moved offshore in the lake while warm, lake water intruded in the wetlands near the surface. This flushing prevented the occurrence of anoxic conditions in the wetlands which were used by adult and larval fish, mainly tilapia, and freshwater shrimps (Caridina nilotica). No fish larvae were found at more than 150 m from the shore in the three bays. Both larval fish and Caridina nilotica appeared to move in and out of the wetlands at night, possibly as a result of low dissolved oxygen as well as sheltering from predation. Their abundance varied with the lunar phase, with maximum near new moon and first quarter. Being the only protected wetlands in Lake Victoria, Rubondo Island may increasingly become an important source of replenishment for fisheries in the lake which elsewhere appears overfished.     

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.     

Temporal dynamics of dissolved oxygen in a floating–leaved macrophyte bed

1. Oxygen concentrations in shallow vegetated areas of aquatic systems can be extremely dynamic. In these waters, characterizing the average oxygen content or frequency of low oxygen events (hypoxia) may require high frequency measurements that span seasons and even years. In this study, moored sondes were used to collect 15‐min interval dissolved oxygen (DO) readings in an embayment of the tidal Hudson River with dense coverage by an invasive floating leaved plant (Trapa natans) and in adjacent open waters. Measurements were made from late spring to summer over a 2‐year period (2005, 2006). 2. Oxygen concentrations were far more dynamic in the vegetated embayment than in the adjacent open waters and while hypoxic conditions never occurred in the open waters, they occurred frequently in the vegetated site. Overall the vegetated site was hypoxic (DO < 2.5 mg L^?1) 30% of the time and had an average oxygen concentration of 5.1 mg L^?1. Oxygen concentration was significantly (P < 0.0001, anova ) related to season, year and tide. Low tide periods during summer of 2006 had the lowest average oxygen concentration and the highest frequency of hypoxia. 3. The greater hypoxia in summer than spring is related to changes in plant morphology. In the spring and early summer when plants are submersed hypoxia occurs at lower frequency and duration than in the summer when dense floating vegetation covers the water. The tidal pattern in oxygen is related to hydrologic exchange with the non‐vegetated open waters. Year‐to‐year variation may be related to relatively small changes in plant biomass between years. 4. Oxygen concentrations in aquatic systems can be critical to habitat quality and can have cascading impacts on redox sensitive nutrient and metal cycling. For some systems with dynamic oxygen patterns neither weekly spot sampling nor short‐duration, high‐frequency measurements may be sufficient to characterize oxygen conditions of the system.     

Recent papyrus swamp habitat loss and conservation implications in western Kenya

Papyrus Cyperus papyrus swamps are characteristic of many wetlands of tropical Africa. Like most wetland habitats worldwide, they are under human pressure due to harvesting and reclamation for agriculture. Changes in papyrus cover were assessed using aerial photographs at three Important Bird Areas (IBAs) in the Kenyan sector of Lake Victoria. Papyrus area losses of 50% (Dunga), 47% (Koguta) and 34% (Kusa) occurred between 1969 and 2000. The habitat loss and degradation at the sites appeared purposeful, driven by demand for agricultural land and other papyrus products. Cleared papyrus and cultivated areas around all sites increased over the same period. If papyrus habitat loss continues at current rates, papyrus swamps at Dunga and Koguta will disappear by 2020, and Kusa will be reduced to only 19% of its 1969 area. Human population growth around the sites, with concomitant increase in land use activities and papyrus harvesting are the major factors that account for papyrus area reductions. Loss of total habitat is accompanied by deterioration in habitat quality within remaining areas. Papyrus physical structure (height and density) inversely correlate to human disturbances that include footpaths, cutting, burning, grazing and farming. The continued papyrus habitat loss and degradation represents a significant threat to biodiversity conservation particularly for papyrus-specialist birds and other papyrus-reliant species in western Kenya. The observed pattern of papyrus extents and land use changes at all sites provide the site-scale information necessary for papyrus conservation planning. In particular, conservation action is needed most urgently at Dunga and Koguta as they face severe land use pressures.     

A LIMNOLOGICAL INVESTIGATION OF LAKE LIAMBEZI,CAPRIVI

SUMMARY

A general investigation was carried out on Lake Liambezi to characterize the limnological features of the lake. The lake, 101 km² in area and situated within the tropics, was found to be shallow and bordered by reed swamps, particularly on the south-western shore. It was polymictic and consequently oxygen levels were high at all depths. Transparency was such that photosynthesis occurred to the bottom. Conductivity ranged from 17,3 to 41,2 mSm^?1 and the waters were moderately alkaline. Transfer of organic detritus from the reed swamp into the open water has resulted in the presence of high levels of organic nitrogen and phosphorus whilst ambient levels of inorganic forms were low. Planktonic chlorophyll a concentrations ranged between 1 and 104 μg l^?1 and Microcystis was the dominant phytoplankter. The dominant zooplankter was Bosmina. The important species of phytoplankton and zooplankton are listed and include two new species of blue-green algae which are described. The lake was found to be generally similar to most shallow tropical African lakes with a wide diversity of food sources for fish. Progressive encroachment by Salvinia appears to hamper access to fishing areas and the fishery potential is threatened.     

In situ and laboratory studies on the behaviour and survival of Pacific salmon (genus Oncorhynchus)

Juvenile Pacific salmon display a marked surface water orientation during downstream migration, estuarine and nearshore coastal rearing phases. Many estuaries in British Columbia are vertically stratified with a shallow, well-defined halocline which can restrict the dispersion of wastes discharged into less saline surface waters and impose constraints upon aquatic organisms. In situ experiments in an estuary receiving a surface discharge of treated pulp mill wastes, revealed conditions which were lethal to underyearling salmon at, and below the halocline (4.0–6.5 m depth). Behavioural bioassays determined that juvenile chinook salmon were biased towards the water surface and avoided waters at depth. Dissolved oxygen was the variable which affected this distribution most significantly. Surface waters receiving effluent from another pulp mill were lethal to juvenile salmon within 350 m, and a significant vertical avoidance response occurred within 350–950 m of the outfalls. The behavioural response was significantly correlated with in situ temperature, pH and colour (effluent).As a complement to field experiments we developed a 4500 l water column simulator (WCS) to examine salmon behaviour in the laboratory. We investigated the surface water orientation behaviour of juvenile salmon in relation to variations in salinity and dissolved oxygen. Under simulated vertically stratified estuarine conditions, the fish moved freely between overlying fresh water and salt water. Induction of hypoxic conditions in fresh water elicited a downward distribution shift towards the halocline and oxygenated, but more saline, waters. Avoidance reactions (50% level) occurred consistently up to 7–8 mg · l^–1 dissolved oxygen. Salmon continued to examine the hypoxic freshwater zone despite sub-optimal conditions.     

Ecology of a diplozoon parasite on the gills of the African cyprinid Barbus neumayeri

In oxygen‐deficient waters, the difficulties of oxygen uptake in gill parasites and their fish hosts may influence host and parasite densities, site selection by the parasite, and effects of the parasite on host condition. This study quantified the prevalence and intensity of the gill monogenean Neodiplozoon polycotyleus in the African cyprinid fish Barbus neumayeri from an intermittent forest stream in western Uganda. Oxygen levels were low in the stream over the 12‐month study, averaging only 2.5 mg litre^?1 (monthly range = 1.2–4.3 mg litre^?1). However, parasite prevalence was high (47.2%), suggesting high tolerance to low oxygen in N. polycotyleus. The prevalence of parasites varied with host body size, with the highest frequency of occurrence in the middle size classes. Prevalence also varied over the year; seasonal peaks of rainfall coincided with a lower frequency of N. polycotyleus. The significantly nonrandom frequency distribution of parasites among hosts suggests regulation of parasite numbers. Of the hosts infected, 37.1% harboured one N. polycotyleus parasite, and 62.9% harboured two parasites. No fish were infected with more than two diplozoons. There was evidence for strong site specificity by N. polycotyleus within hosts; 77.7% of the parasites were located on the filaments of the second gill arch, which may relate to increased oxygen availability. In addition, only one of the 178 infected fish had more than one parasite on one side of the branchial basket. Although N. polycotyleus is undoubtedly parasitic, we found no evidence of a negative parasitic effect on the condition or reproductive status of B. neumayeri.     

Major ion chemistry in a tropical African lake basin

SUMMARY. The amount of water lost by seepage from Lake Naivasha, calculated as the residual in the water budget, was 5 (1973), 11 (1974) and 20% (1975) of the total water loss. Direct measurements of seepage in nearshore shallows indicated that water entered the lake via ground-water seepage in the northern portion and left the lake in the southern portion. Naivasha lies in a topographic closed basin but is hydrologically a seepage lake; it is distinctive among the lakes lying in the endorheic rift valleys of Ethiopia, Kenya and Tanzania because the water is fresh. The mean chemical composition (mg l^?1) of bulk precipitation collected on two transects across the rift valley near L. Naivasha and at three stations near Nairobi was: Na, 0.54, K, 0.31; Ca, 0.19; Mg, 0.23; SO₄, 0.72 and Cl, 0.41. The major inflow, the Malewa River, was largely a solution of bicarbonate (1.15 m-equiv, l^?1), sodium (9 mg l^?1) and calcium (8 mg l^?1) and carried a total dissolved solid load (kg ha^?1) of 62 (1973) and 120 (1974). Total dissolved solids, sodium, calcium and bicarbonate concentrations were inversely related to discharge. Among the standing waters in the Naivasha basin, sodium and bicarbonate are predominant and the mean total dissolved solute content (mg l^?1) increased from a low under the northern papyrus swamp (217) to intermediate values in L. Naivasha (329), the nearshore lagoons (367) and Crescent Island basin (394) to a high in Oloidien Lake (831). Ion ratios and stability field diagrams indicated that calcite is formed in the Crescent Island basin and Oloidien Lake. Calcite was detected in the sediments of the two basins. Ion ratios also indicated that sulphate was retained in the sediments of the northern papyrus swamp. Several factors combine to keep L. Naivasha's water fresh. A large fraction of the water supplied to the lake comes from dilute rivers and rain. The lake does not lie in a closed basin, but loses water and solutes via seepage. Exchanges with sediments both in the pelagic and littoral regions of the lake are the major routes for solute movement and biochemical sedimentation is a major factor in removal of silica. Both geochemical and biochemical sedimentation account for much of the potassium, calcium, magnesium and bicarbonate removal. Only low levels of the conservative ions, sodium and chloride, are taken up by the sediments, while very low levels of sulphate and fluoride are actually released.     

The influence of fish size size on the avoidance of hypoxia and oxygen selection by largemouth bass

When an oxygen gradient ranging from c . 10 to 95% air saturation was formed in a 5 m chamber, largemouth bass Micropterus salmoides avoided water in which dissolved oxygen values were <27% air saturation. There was a significant ( P <0·05) correlation between fish mass and the level of dissolved aquatic oxygen that was selected. Small fish (23–500 g) utilized waters of lower oxygen levels than did the larger fish (1000–3000 g). The results of this study suggest that largemouth bass are able to sense and avoid hypoxic water, and select aquatic oxygen tensions that maintain their metabolic scope for growth and activity.     

The impact of fire regimes on populations of an endangered lizard in montane south‐eastern Australia

The Blue Mountains water skink (Eulamprus leuraensis; Scincidae) is restricted to less than 40 fragmented swamp sites, all within the Blue Mountains and Newnes Plateau areas of New South Wales, Australia. Climate change is expected to increase fire frequency in the area, potentially degrading habitat quality for this endangered reptile. We quantified lizard abundances in 12 swamps using standardized surveys, and constructed a Global Information System (GIS) database to determine fire‐histories for each swamp since 1967. The abundance of Blue Mountains water skinks was negatively correlated with fire frequency, but not with time since fire. Indirect impacts of fire (mediated via shifts in vegetation density, moisture levels, prey availability and post‐fire predation) may be more important than direct effects in these cool, moist habitats. Although lizards were less common in swamps close to urban areas, and less common in frequently burnt areas, viable populations of this endangered reptile still persist even in anthropogenically disturbed swamps and in swamps that have experienced up to four fires in 20 years. Future research could usefully extend these analyses to other swamps in the locality, and explore the broader impacts of fire regimes on the distinctive flora and fauna of this threatened ecological community.     

The benthic macrofauna community of Kenyan waters of Lake Victoria

The benthic macroinvertebrates in the Kenya waters of Lake Victoria (ca. 1400 km²) were surveyed during four 10 day sampling periods in February, April, August and November 1984. Fourty three taxonomic groups were recorded. Oligochaetes, molluscs and dipteran larvae were the most abundant and widely distributed groups. The dominant oligochaetes were Branchiura sowerbyii Beddard and the swamp worm Alma emini Michaelsen, which were abundant in all silty and soft mud areas in both littoral and open water zones. Melanoides tuberculata Müller, Bellamya unicolor Olivier and Caelatura spp. were the most abundant molluscs. Different patterns were observed between near-littoral stations (<8 m deep) and deep water stations (8–40 m deep). After 10 years (1994), the macroinvertebrates of Lake Victoria were examined again. No evidence was found of seasonal changes in the composition or benthos density, although local changes occurred in some species which were attributed to swarming behaviour or larval settlement patterns. The spatial distribution of the fauna may be influenced primarily by oxygen availability and industrial effluent discharged from paper mill and agro-based industries in the catchment of the lake. There has been a shift in abundance, from an Oligochaeta and Insecta dominated community in 1984, to the present community dominated by Mollusca and Oligochaeta. The ecological role of the benthic community and its recent changes to fisheries production in the lake is discussed.     

Responses of a fringing Cyperus papyrus L. swamp to changes in water level

Over a 9-year period (1993–2001), the land-water width of a papyrus fringe on the southern shore of Lake Naivasha, Kenya, varied between 40 and 60 m. Increases in width via rhizome spreading into open water followed the 1997/1998 El Niño flood when water depths rose by about 2 m. Germination of papyrus seeds also responded to water depth with a mean ± S.E. rate in experiments of 23 ± 6% after 21 days when water level was 5 cm below the sediment surface. No germination occurred when sediment was flooded or allowed to desiccate. Rhizome spreading from floating mats appeared to be favoured by deep water with seedling spread favoured on newly inundated, low-gradient slopes in shallow water. Although natural regenerative capacity was influenced by water depth, the height, density, biomass and chemical content of papyrus were not. Total average biomass along a land-water transect was 6950 ± 860 g m⁻² which was large in relation to nutrient and mineral contents. Culms contained 0.47 ± 0.14% N and 0.06 ± 0.05% P and rhizomes 0.71 ± 0.21% and 0.10 ± 0.06%. Sediment underlying the swamp was aerobic and there were small land-water gradients in the BOD of swamp water and sediment. However, chemical gradients were weak compared with wider papyrus swamps elsewhere. Lake and swamp water mixed in the narrow fringe studied and residence times for organic matter may not have been long enough for organic material to mineralise before entering lake water.