Two hundred years of a diverse Daphnia community in Lake Naivasha (Kenya): effects of natural and human-induced environmental changes

1. We used fossil diapausing eggs extracted from ²¹⁰Pb‐dated sediment cores to reconstruct historical changes in the Daphnia community of Lake Naivasha, a climate‐sensitive lake in Kenya which over the past 200 years has experienced a series of well‐documented natural and anthropogenic environmental changes. 2. Contiguous sampling and analysis of four cores yielded ephippial capsules of eight Daphnia species. Only two of these had been recorded previously in live collections from Lake Naivasha, and one species is new to science. The four more common species (Daphnia barbata, D. laevis, D. magna, and D. pulex) show striking differences in abundance patterns and population dynamics through time. Four other species (D. lumholtzi, D. curvirostris, D. longispina s.l., and Daphnia sp. nov. type Limuru.) appear to have been present only occasionally. Nevertheless, between 1895 and 1915 seven species of Daphnia inhabited Lake Naivasha simultaneously. 3. Despite considerable natural environmental change associated with climate‐driven lake‐level fluctuations, the Daphnia community of Lake Naivasha has been severely affected by human activities over the past century, especially the introduction of exotic fishes and water‐quality changes because of agricultural soil erosion. The recent reappearance of large‐bodied Daphnia species (D. magna, D. barbata, D. lumholtzi, Daphnia sp. nov. type Limuru) after 20–110 years of absence can be explained by their release from fish predation, following a dramatic increase in turbidity caused by excess clastic sediment input from eroded catchment soils. The small‐bodied species D. laevis has fared less well recently, presumably because the benefit of lowered predation pressure is counteracted by more pronounced negative effects of increased turbidity on this species and loss of submerged macrophyte beds which formerly served as predation refuge. 4. Our results suggest that, despite considerable environmental instability and the absence of specialised zooplanktivores, top‐down control of fish on large zooplankton is important in Lake Naivasha. Predation pressure from fish has led to clear‐cut shifts in local Daphnia species composition, but failed to drive the larger taxa to extinction.     

Behavioural responses to co-occurring threats of predation and ultraviolet radiation in Daphnia

1. Organisms in the wild are faced with multiple threats and a common response is a change in behaviour. To disentangle responses to several threats, we exposed two differently sized species of the freshwater invertebrate Daphnia to solar ultraviolet radiation (UVR) and predation from either moving pelagic or benthic ambush predators.
2. Using an advanced nanotechnology-based method, we tracked the three-dimensional movements of those mm-sized animals at the individual level. Each behavioural trial was performed both under conditions resembling night (no UVR) and day (UVR) and we examined patterns of the depth distribution and swimming speed by Daphnia across three treatments: no predator (control); bottom-dwelling damselfly (Calopteryx sp.); and fish (stickleback, Pungitius pungitius) predators. We also quantified the actual predation rate by the two predators on the two Daphnia species, Daphnia manga and Daphnia pulex.
3. We show that individual Daphnia are able to identify predators with different feeding habitats, rank multiple and simultaneously occurring risks and respond in accordance with the actual threat; complex responses that are generally associated with larger animals.
4. In a broader context, our results highlight and quantify how a cocktail of everyday threats is perceived and handled by invertebrates, which advances our understanding of species distribution in space and time, and thereby of population dynamics and ecosystem function in natural ecosystems.

     

Changes of phytoplankton communities in Lakes Naivasha and Oloidien, examples of degradation and salinization of lakes in the Kenyan Rift Valley

Increasing degradation of the water quality, caused by overuse and salinization, leads to considerable changes of the phytoplankton composition in Kenyan Rift Valley lakes. Exemplarily, the phytoplankton communities and biomasses of deteriorating freshwater Lake Naivasha and salinizing Lake Oloidien were studied between 2001 and 2005, accompanied by physico-chemical measurements (pH, total phosphorus and nitrogen, alkalinity, conductivity). Over the last three decades, the ecology of these two water basins has been subjected to dramatic changes, caused by excessive use of water and catchment area by man. In L. Naivasha a shift in the dominance of coccoid cyanobacteria towards dominance of Chlorophyceae (Botryococcus terribilis) was observed. Lake Oloidien exhibited a shift in the dominance of coccoid Chlorophyceae towards dominance of cyanobacteria (Arthrospira fusiformis, Anabaenopsis elenkinii). Phytoplankton findings and chemical data demonstrate that L. Naivasha has developed towards a eutrophic freshwater lake while L. Oloidien has progressed towards a hypereutrophic alkaline-saline lake. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Padisak     

Analysis of trophic relationships in two shallow equatorial lakes Lake Naivasha (Kenya) and Lake Ihema (Rwanda) using a multispecifies trophic model

A multispecifies trophic model called ECOPATH II, which can be used to describe the trophic relationships in aquatic ecosystems on a quantitative basis, is briefly presented. When properly used, it can help to explain the trophic relationships in ecosystems and possible evolution of fishstocks after modifications of the environment (e.g. eutrophication, introduction of a new population and/or a significant increase of the fishing effort), and to compare the trophic structure of several ecosystems. Examples are provided on two shallow lakes: Lake Ihema and Lake Naivasha. They are compared with Lake George which was previously documented.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

Larval chaoborid mandibles in surface sediments of small shallow lakes in Finland: implications for palaeolimnology

The aim of this study was to assess the value of phantom midge Chaoborus (Diptera: Chaoboridae) larvae as a palaeoenvironmental proxy in northern Europe. The presence of Chaoborus was examined in 80 small shallow lakes across Finland (60–70°N), based on their subfossil mandibles preserved in lake sediments. Chaoborus flavicans was present in 33 lakes, but was the most abundant midge taxon in only one of the lakes. Chaoborus crystallinus/obscurripes mandibles were present in two lakes. Statistical analyses showed that the distribution of Chaoborus was significantly correlated with lake depth and mean July air temperature. Chaoborus was absent in the northern study lakes, whereas the results indicated that C. flavicans was frequently found in small shallow lakes in southern Finland. Chaoborus flavicans was particularly abundant in fish-free, macrophyte-rich, dystrophic lakes, but absent in oligotrophic clear-water lakes having fish. The results of this study indicate that the subfossil mandibles of Chaoborus can provide valuable information in multiproxy palaeolimnological studies, especially when investigating past changes in water level and temperature in small shallow lakes in northern Europe. Guest editors: K. Buczkó, J. Korponai, J. Padisák & S. W. Starratt Palaeolimnological Proxies as Tools of Environmental Reconstruction in Fresh Water     

Physical-chemical influences on vernal zooplankton community structure in small lakes and wetlands of Wisconsin,U.S.A.

We sampled zooplankton communities from 54 small water bodies distributed throughout Wisconsin to evaluate whether a snap-shot of zooplankton community structure during early spring could be used for the purpose of differentiating lakes from wetlands. We collected a single set of zooplankton and water chemistry data during a one-month time window (synchronized from south to north across the state) from an open water site in each basin as a means to minimize and standardize sampling effort and to minimize cascading effects arising from predator–prey interactions with resident and immigrant aquatic insect communities. We identified 53 taxa of zooplankton from 54 sites sampled across Wisconsin. There was an average of 6.83 taxa per site. The zooplankton species were distributed with a great deal of independence. We did not detect significant correlations between number of taxa and geographic region or waterbody size. There was a significant inverse correlation between number of taxa and the concentration of calcium ion, alkalinity and conductivity. One pair of taxa, Lynceus brachyurus and Chaoborus americanus, showed a significant difference in average duration of sites of their respective occurrence. All other pairs of taxa had no significant difference in average latitude, waterbody surface area, total phosphorus, total Kjeldahl nitrogen, alkalinity, conductivity, calcium ion, sulfate, nitrate, silicate or chloride. Taxa were distributed at random among the sites – there were no statistically significant pairs of taxa occurring together or avoiding each other. Multivariate analysis of zooplankton associations showed no evidence of distinct associations that could be used to distinguish lakes from wetlands. Zooplankton community structure appears to be a poor tool for distinguishing between lakes and wetlands, especially at the relatively large scale of Wisconsin (dimension of about 500 km). The data suggest that a small body of water in Wisconsin could be classified as a wetland if it persists in the spring and summer for only about 4 months, and if it is inhabited by Lynceus brachyurus, Eubranchipus bundyi, and if Chaoborus americanus and Chydorus brevilabris are absent.     

Drought and aquatic community resilience: the role of eggs and seeds in sediments of temporary wetlands

1. A long‐lived bank of propagules consisting of eggs, seeds and spores is one mechanism that allows aquatic communities to survive drought. A drying (drought) event is, for aquatic organisms in a temporary wetland, a phase from which communities must recover. Such a dry phase is often considered a disturbance but should not be considered adverse or catastrophic for the organisms that have evolved to live in temporarily wet habitats. 2. This paper explores the parallels between the egg bank of zooplankton and the seed bank of aquatic plants as means of survival in temporary wetlands. The resilience of communities in temporary wetland ecosystems is assessed by examining dormancy, hatching, germination, establishment and reproduction of animals and plants from the egg and seed banks of wetlands with a range of wetting and drying regimes. 3. Both the zooplankton and aquatic plants of the temporary wetlands studied rely on their egg and seed banks as a means for surviving drying. These communities recover after the disturbance of drying by means of specific patterns of dormancy, dormancy breakage, hatching, germination, establishment and reproduction. Spatial and temporal patterns of species richness allow resilience through dormancy, as not all species are present at all sites and not all species hatch and germinate at the same time. Multiple generations in the egg and seed bank and complexity of environmental cues for dormancy breakage also contribute to the ecosystem's ability to recover after a drying event. A persistent egg and seed bank allows species‐rich communities to hatch, germinate and develop rapidly once dormancy is broken. Rapid establishment of species‐rich communities that reproduce rapidly and leave many propagules in the egg and seed bank also facilitates community recovery on flooding of a temporary wetland after a drying event. 4. To maintain the diversity of temporary wetland communities through droughts and floods we need to manage the dry and wet phases of wetlands. To conserve a wide range of wetland types, we need to maintain a variety of hydrological patterns across the landscape.     

Influence of predation by fish and water turbidity on a Daphnia gessneri population in an Amazonian floodplain lake,Brazil

The population behavior of Daphnia gessneri Herbst, 1967 in a floodplain lake (Lago Grande) of the lower Rio Solimões was investigated between April 1979 and March 1980 with regard to 1) predation by the fish called tambaqui (Colossoma macropomum, Characidae), 2) water level fluctuation and 3) water transparency. Zooplankton density samples were collected at two sites near mid-lake, where water depth and Secchi disc transparency were measured. In addition, qualitative samples of zooplankton and fish collections were taken at several sites in the adjacent floodplain areas. The author concludes that fluctuations in Daphnia gessneri populations correlate most with intense predation by fish and water turbidity.     

Sub-fossils of cladocerans in the surface sediment of 135 lakes as proxies for community structure of zooplankton,fish abundance and lake temperature

To elucidate the possibilities of using zooplankton remains in the surface sediment to describe present-days community structure and population dynamics of zooplankton, fish abundance and temperature, we compared contemporary data sampled in the pelagial during summer with the sediment record from the upper 1 cm of the sediment in 135 lakes covering a latitude gradient from Greenland in the north to New Zealand in the south. The abundance of three genera Bosmina, Daphnia and Ceriodaphnia of the total pool of ephippia was significantly related to the total abundance of the same taxa in the pelagic zone. However, in most lakes the abundance of Ceriodaphnia was higher in the sediment than in the water, which may be attributed to the overall preference by this genus for the littoral habitat. Using contemporary data from 27 Danish lakes sampled fortnightly during summer for 10 years, we found substantial inter-annual variations in the abundance of Daphnia spp., Ceriodaphnia spp., B. longirostris and B. coregoni. Yet, the sediment record mimicked the medium level well for most of the lakes, which suggests that the sediment record provides an integrated picture of the pelagic cladoceran community, which otherwise can be obtained only by long-term frequent contemporary sampling for several years. The contribution of Daphnia to the sum of Daphnia and Bosmina ephippia was negatively correlated with the abundance of fish expressed as catch per night in multi-mesh sized gill nets (CPUE). Yet, region-specific differences occurred, which partly could be eliminated by including nutrient state expressed as total phosphorus (TP) in a multiple regression. The average ratio of ephippia to the sum of ephippia and carapaces of Bosmina varied 40-fold between the sampling regions and was significantly negatively related to summer mean air temperature, and for Danish lakes also, albeit weakly, to fish CPUE but not to chlorophyll a. Apparently, temperature is the most important factor determining the ratio of parthenogenetic to ephippia producing specimens of Bosmina. We conclude that the sediment record of cladocerans is a useful indicator of community structure of pelagic cladocerans and the abundance of fish and temperature.     

Combined stable isotope and gut contents analysis of food webs in plant-dominated, shallow lakes

Summary 1. To determine feeding links between primary producers, invertebrates and fish, stable isotope analyses and gut content analyses of fish were conducted on the components of four shallow, eutrophic to hypertrophic, plant-dominated lakes.
2. Although separation of basal resources was possible, the diets of both fish and invertebrates were broad, comprising food from different compartments (planktonic, epiphytic/benthic), as well as from different trophic levels.
3. Mixing models were used to determine the extent to which periphyton production supported higher trophic levels. Only one species of invertebrate relied upon periphyton production exclusively.
4. Fish density affected the diets of invertebrates. The response was different for planktonic and epiphytic/benthic invertebrates. The proportion of periphyton production in the diets of zooplankton appeared to increase with fish density, whilst it decreased for other invertebrates.
5. As all zooplankton samples were collected in the open water at dusk, these results are further evidence for the diurnal horizontal migration of zooplankton. Although not conclusive, they are consistent with a behavioural response by invertebrates and zooplankton in the presence of fish.     

<Emphasis Type="Italic">Daphnia</Emphasis>species diversity in Kenya,and a key to the identification of their ephippia

The distribution of Daphniaspecies in tropical Africa is poorly known and understood. Daphniaare assumed rare in tropical regions, but systematic studies covering large areas are sparse. We sampled the active community (live zooplankton) and/or the dormant community (diapausing egg banks in the sediment) of 41 standing water bodies in Kenya in search for Daphnia.Overall the dormant communities yielded 11 species of Daphnia, a species richness more than twice the species richness found in the active communities. Dormant community species diversity better reflects the spatial, and particularly the temporal (multi-annual) variation in environmental conditions available to Daphniain these tropical standing waters. Hence, we suggest that the dormant community be taken into account when assessing local zooplankton diversity, especially in fluctuating tropical lake ecosystems, where the presence of each local Daphniaspecies in the active community may be strongly seasonal or erratic. Geographic distribution data from this study are supplemented with previous records of Daphniain East Africa to provide an overview of the known distribution of Daphniain Kenya and neighbouring countries. We also present a detailed key for morphological identification of the ephippia of the 11 Daphniaspecies encountered, complemented with photographs and drawings of diagnostic characters.     

小型浅水湖泊沉积物磷的赋存形态及其相关性分析

以孔目湖沉积物为研究对象, 应用七步连续提取法测定其中的不同形态磷, 探讨了该湖泊沉积物中各赋存形态磷的分布特征, 并对其进行了相关性分析。结果表明: 该湖泊遭受的磷内源负荷比较大, 磷污染严重; 沉积物中TP含量在2338.63-2954.98 mg•kg-1, 平均为2671.37 mg•kg-1; 沉积物中各形态磷含量从高到低依次为: Fe-P>De-P>Ca-P> OP>Al-P>Oc-P>Ex-P, 分别占TP的53.9%、28.7%、8.8%、6.2%、1.2%、0.9%、0.3%; 生物有效磷含量为1583.59 mg•kg-1, 占TP的59.28%; 沉积物中TP与Fe-P极显著相关, Oc-P与Ca-P和OP与Ca-P均相关, 而TP与Ex-P、Al-P、Ca-P和OP相关性较差, 说明沉积物中TP含量主要来自于Fe-P。这一研究结果为揭示小型湖泊富营养化发生机制提供了数据及理论支撑。     

Restoring lakes by using artificial plant beds: habitat selection of zooplankton in a clear and a turbid shallow lake

1. Return of large‐bodied zooplankton populations is of key importance for creating a shift from a turbid to a clear‐water state in shallow lakes after a nutrient loading reduction. In temperate lakes, recovery is promoted by submerged macrophytes which function as a daytime refuge for large zooplankton. However, recovery of macrophytes is often delayed and use of artificial plant beds (APB) has been suggested as a tool to enhance zooplankton refuges, thereby reinforcing the shift to a clear‐water state and, eventually, colonisation of natural plants. 2. To further evaluate the potential of APB in lake restoration, we followed the day–night habitat choices of zooplankton throughout summer in a clear and a turbid lake. Observations were made in the pelagic and littoral zones and in APB in the littoral representing three different plant densities (coverage 0%, 40% and 80%). 3. In the clear lake, the zooplankton (primarily Daphnia) were mainly found in the pelagic area in spring, but from mid‐May they were particularly abundant in the APB and almost exclusively so in mid‐June and July, where they appeared in extremely high densities during day (up to 2600 ind. L⁻¹). During night Daphnia densities were overall more equally distributed between the five habitats. Ceriodaphnia was proportionally more abundant in the APB during most of the season. Cyclopoids were more abundant in the high APB during day but were equally distributed between the five habitats during night. 4. In the turbid lake, however, no clear aggregation was observed in the APB for either of the pelagic genera (Daphnia and Bosmina). This may reflect a higher refuge effect in the open water due to the higher turbidity, reduced ability to orient to plant beds and a significantly higher fish density (mainly of roach, Rutilus rutilus, and perch, Perca fluviatilis) in the plant beds than in the clear lake. Chydorus was found in much higher proportions among the plants, while cyclopoids, particularly the pelagic Cyclops vicinus, dominated in the pelagic during day and in the pelagic and high density plants during night. 5. Our results suggest that water clarity is decisive for the habitat choice of large‐bodied zooplankton and that introduction of APB as a restoration measure to enhance zooplankton survival is only a useful tool when water clarity increases following loading reduction. Our results indicate that dense APB will be the most efficient.     

Metagenetic community analysis of microbial eukaryotes illuminates biogeographic patterns in deep-sea and shallow water sediments

Microbial eukaryotes (nematodes, protists, fungi, etc., loosely referred to as meiofauna) are ubiquitous in marine sediments and probably play pivotal roles in maintaining ecosystem function. Although the deep-sea benthos represents one of the world's largest habitats, we lack a firm understanding of the biodiversity and community interactions amongst meiobenthic organisms in this ecosystem. Within this vast environment, key questions concerning the historical genetic structure of species remain a mystery, yet have profound implications for our understanding of global biodiversity and how we perceive and mitigate the impact of environmental change and anthropogenic disturbance. Using a metagenetic approach, we present an assessment of microbial eukaryote communities across depth (shallow water to abyssal) and ocean basins (deep-sea Pacific and Atlantic). Within the 12 sites examined, our results suggest that some taxa can maintain eurybathic ranges and cosmopolitan deep-sea distributions, but the majority of species appear to be regionally restricted. For Operationally Clustered Taxonomic Units (OCTUs) reporting wide distributions, there appears to be a taxonomic bias towards a small subset of taxa in most phyla; such bias may be driven by specific life history traits amongst these organisms. In addition, low genetic divergence between geographically disparate deep-sea sites suggests either a shorter coalescence time between deep-sea regions or slower rates of evolution across this vast oceanic ecosystem. While high-throughput studies allow for broad assessment of genetic patterns across microbial eukaryote communities, intragenomic variation in rRNA gene copies and the patchy coverage of reference databases currently present substantial challenges for robust taxonomic interpretations of eukaryotic data sets.     

Vertical distribution of bacterial community structure in the sediments of two eutrophic lakes revealed by denaturing gradient gel electrophoresis (DGGE) and multivariate analysis techniques

Vertical distribution of bacterial community structure was investigated in the sediments of two eutrophic lakes of China, Lake Taihu and Lake Xuanwu. Profiles of bacterial communities were generated using a molecular fingerprinting technique, denaturing gradient gel electrophoresis (DGGE) followed by DNA sequence analysis, and the results were interpreted with multivariate statistical analysis. To assess changes in the genetic diversity of bacterial communities with changing depth, DGGE banding patterns were analysed by cluster analysis. Distinct clusters were recognized in different sampling stations of Lake Taihu. Canonical correspondence analysis (CCA) was carried out to infer the relationship between environmental variables and bacterial community structure. DGGE samples collected at the same sampling site clustered together in both lakes. Total phosphorus, organic matter and pH were considered to be the key factors driving the changes in bacterial community composition.