Ratio-dependent response of a temperate Australian estuarine system to sustained nitrogen loading

Classical resource- and the less studied ratio-dependent models of predator–prey relationships provide divergent predictions as to the sustained ecological effects of bottom-up forcing. While resource-dependent models, which consider only instantaneous prey density in modelling predator responses, predict community responses that are dependent on the number of trophic levels in a system, ratio-dependent models, which consider the number of prey per consumer, predict proportional increase in each level irrespective of chain length. The two models are only subtly different for systems with two or three trophic levels but in the case of four trophic levels, predict opposite effects of enrichment on primary producers. Despite the poor discriminatory power of tests of the models in systems with two or three trophic levels, field tests in estuarine and marine systems with four trophic levels have been notably absent. Sampling of phytoplankton, macroinvertebrates, invertebrate-feeding fishes, piscivorous fishes in Kooloonbung Creek, Hastings River estuary, eastern Australia, subject to over 20 years of sewage discharge, revealed increased abundances in all four trophic levels at the disturbed location relative to control sites. Increased abundance of phytoplankton at the disturbed site was counter to the predictions of resource-dependent models, which posit a reduction in the first trophic level in response to enrichment. By contrast, the increase in abundance of this first trophic level and the proportionality of increases in abundances of each of the four trophic groups to nitrogen loading provided strong support for ratio dependency. This first evidence of ratio dependence in an estuarine system with four trophic levels not only demonstrates the applicability of ecological theory which seeks to simplify the complexity of systems, but has implications for management. Although large nutrient inputs frequently induce mortality of invertebrates and fish, we have shown that smaller inputs may in fact enhance biomass of all trophic levels.     

Top-down control of phytoplankton in a shallow hypertrophic lake: Little Mere (England)

Top-down control of phytoplankton by zooplankton is possible through reductions in density of zooplanktivorous fish. Little Mere is a shallow lake where the effects of sewage effluent caused such a reduction. This allowed the large-bodied cladoceran, Daphnia magna Straus, to develop huge populations, preventing potentially large algal crops from developing.Subsequent diversion of the effluent is anticipated to lead to recovery of the fish community, reduced numbers of large-bodied grazers, and increased phytoplankton biomass. Whether the aquatic plant community, present in Little Mere, is resilient to such changes may depend upon whether cyanophytes are favoured, or not.     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

Predator personality structures prey communities and trophic cascades

Intraspecific variation is central to our understanding of evolution and population ecology, yet its consequences for community ecology are poorly understood. Animal personality – consistent individual differences in suites of behaviours – may be particularly important for trophic dynamics, where predator personality can determine activity rates and patterns of attack. We used mesocosms with aquatic food webs in which the top predator (dragonfly nymphs) varied in activity and subsequent attack rates on zooplankton, and tested the effects of predator personality. We found support for four hypotheses: (1) active predators disproportionately reduce the abundance of prey, (2) active predators select for predator‐resistant prey species, (3) active predators strengthen trophic cascades (increase phytoplankton abundance) and (4) active predators are more likely to cannibalise one another, weakening all other trends when at high densities. These results suggest that intraspecific variation in predator personality is an important determinant of prey abundance, community composition and trophic cascades.     

Ecological and socio‐economic impacts of invasive water hyacinth (Eichhornia crassipes): a review

1. Water hyacinth (Eichhornia crassipes) is one of the world’s most invasive aquatic plants and is known to cause significant ecological and socio‐economic effects. 2. Water hyacinth can alter water clarity and decrease phytoplankton production, dissolved oxygen, nitrogen, phosphorous, heavy metals and concentrations of other contaminants. 3. The effects of water hyacinth on ecological communities appear to be largely nonlinear. Abundance and diversity of aquatic invertebrates generally increase in response to increased habitat heterogeneity and structural complexity provided by water hyacinth but decrease due to decreased phytoplankton (food) availability. 4. Effects of water hyacinth on fish are largely dependent on original community composition and food‐web structure. A more diverse and abundant epiphytic invertebrate community may increase fish abundance and diversity, but a decrease in phytoplankton may decrease dissolved oxygen concentrations and planktivorous fish abundance, subsequently affecting higher trophic levels. 5. Little is known about the effects of water hyacinth on waterbird communities; however, increases in macroinvertebrate and fish abundance and diversity suggest a potentially positive interaction with waterbirds when water hyacinth is at moderate density. 6. The socio‐economic effects of water hyacinth are dependent on the extent of the invasion, the uses of the impacted waterbody, control methods and the response to control efforts. Ecosystem‐level research programmes that simultaneously monitor the effects of water hyacinth on multiple trophic‐levels are needed to further our understanding of invasive species.     

太湖西北湖区2003-2012年间氮磷浓度及浮游植物主要类群变化趋势分析

基于2003-2012年太湖竺山湖和西部沿岸区水体理化指标与浮游植物丰度的逐月监测数据,分析了两个湖区氮磷营养盐状态和浮游植物丰度以及浮游植物主要类群的年际变化趋势及季节变化特征,探讨了浮游植物群落变化与水温及营养盐指标间的关系。结果表明：10年间两个湖区氮磷营养盐浓度总体呈下降趋势,以竺山湖TN、NH₃-N浓度和西部沿岸区NO₃-N浓度下降最为显著;浮游植物丰度总体呈上升趋势,蓝藻在群落结构中日益占据绝对优势;季节变化上,氮营养盐浓度表现为春冬季节高于夏秋季节,TP浓度和浮游植物丰度呈相反的变化趋势。Pearson相关分析显示,水温、NH₃-N浓度和TN/TP是影响蓝藻丰度及其在浮游植物群落中所占比例的主要因素。在温度和营养盐结构的共同作用下,10年间两个湖区蓝藻水华暴发时间逐渐提前,而消退时间逐渐滞后,水华持续时间逐年上升。在全球变暖背景下,太湖水华治理需执行更加严格的氮磷限制阈值,且在重污染的西北湖区控磷依然是关键。     

Size-selective predation on pelagic microorganisms in Arctic freshwaters

Herbivorous zooplankton may have a pronounced influence on pelagicmicroorganisms in Arctic freshwaters. We quantified experimentallythe size-selective feeding of several zooplankton groups onpelagic microorganisms in high Arctic tundra systems. Our experimentsand field study focused on dominant herbivores in Arctic freshwaters,including the cladoceran Daphnia, the copepod Diaptomus andthe anostracan Branchinecta, and their effects on prey rangingin size from bacteria to large phytoplankton. Grazing experimentsshowed that Daphnia were effective predators on all types ofprey, whereas Diaptomus grazed preferentially on larger phytoplanktonwith low clearance rates for bacterial cells. Further analysisby flow cytometry indicated that Diaptomus grazed selectivelyon the largest bacteria. In contrast to the results obtainedin the controlled experiments, Arctic lakes and ponds with azooplankton community dominated by Daphnia had a higher bacterialproduction and abundance than systems not dominated by thisgrazer. This may indicate that the stimulatory effect of grazerson bacterial growth is more pronounced in natural systems, orthat factors other than zooplankton grazing are more importantin regulating bacterial abundance and production in naturalsystems. Although Arctic waters differ considerably from temperatesystems with respect to temperature and light regime, herbivore–preydynamics as well as the bacterial response to temperature appearto be similar between the climatic regions.     

Resource-niche complementarity and autotrophic compensation determines ecosystem-level responses to increased cladoceran species richness

This study examines the relationship between cladoceran species richness and ecosystem functioning. I conducted an experiment in which four cladocerans, Daphnia. magna, D. longispina, D. pulex and Chydorus sphaericus, were cultured in microcosms using different species combinations and levels of species richness. The results demonstrate that even within this closely related group of organisms the effects on ecosystem-level variables, such as total algae and zooplankton biomass, per capita productivity, and nutrient concentrations, as well as phytoplankton community structure, were highly variable between different combinations of these species. Since only four species where involved in this study, species-specific effects dominated the general relationship between species richness and ecosystem functioning. Particular combinations of species resulted in effects that indicated more efficient grazing. These effects, which were most pronounced in combinations including both D. magna and C. sphaericus, were manifested as an indirect effect as the prey community shifted towards grazing-resistant species. As a result, the productivity of the prey community decreased, because phytoplankton species with lower per capita productivity became more dominant. I suggest that the primary mechanism that caused this significant effect was complementarity in prey-size use of D. magna and C. sphaericus. In terms of prey-size range, D. pulex and D. longispina were redundant when D. magna was present and were quickly out-competed by the latter despite higher per capita filtering efficiency. The results show that different mechanisms are important for different combinations of species. Furthermore, the ability of the prey community to respond to changes of consumer species composition is an important factor in experiments in which consumer species richness is experimentally manipulated. Received: 2 November 1998 / Accepted: 8 September 1999     

Vertical distribution of Chlamydomonas changes in response to grazer and predator kairomones

Individuals in aquatic communities frequently assess their biotic environment through infochemicals. In particular, kairomones are commonly involved in interactions between predator and prey. However, the relationship between individuals and chemicals produced by other organisms that are not direct predators, but may indicate the presence of a predator, is not well characterized. We used experimental microcosms to test whether the unicellular green alga Chlamydomonas reinhardtii alters vertical migration patterns in response to kairomones produced by zooplankton ( Daphnia ) and planktivores (fish). Our results suggested that phototaxis in C. reinhardtii was strongly affected by the type of kairomone present, the concentration of the kairomone, and the duration of exposure to the kairomone. Kairomones generally increased phototaxis in C. reinhardtii . The adaptive significance of such behavioral changes in natural settings would depend largely on local community composition. The similarity in phototactic responses of C. reinhardtii to Daphnia and fish kairomone suggest that, in at least this species of phytoplankton, the underlying genetic elements responsible for kairomone detection may be responsive to a broad range of chemical stimuli, allowing this species to adjust its phototaxis in response to not only the presence of its grazers, but also to predators of its grazers.     

Impact of submerged macrophytes on fish-zooplanl phytoplankton interactions: large-scale enclosure experiments in a shallow eutrophic lake

1. The impact of changes in submerged macrophyte abundance on fish-zooplankton-phytoplankton interactions was studied in eighteen large-scale (100 m²) enclosures in a shallow eutrophic take. The submerged macrophytes comprised Potamategon pectinatus L., P. pusillus L. and Callitriche hermaphroditica L. while the fish fry stock comprised three-spined sticklebacks, Gasterosteus acuteatus L., and roach, Rutilus rutilus L. 2. In the absence of macrophytes zooplankton biomass was low and dominated by cyclopoid copepods regardless of fish density, while the phytoplankton biovolume was high (up to 38 mm³1) and dominated by small pennate diatoms and chlorococcales. When the lake volume infested by submerged macrophytes (PVI) exceeded 15–20% and the fish density was below a catch per unit effort (CPUE) of 10 (approx. 2 fry m^?2), planktonic cladoceran biomass was high and dominated by relatively large-sized specimens, while the phytoplankton biovolume was low and dominated by small fast-growing flagellates. At higher fish densities, zooplankton biomass and average biomass of cladocerans decreased and a shift to cyclopoids occurred, while phytoplankton biovolume increased markedly and became dominated by cyanophytes and dinoflagellates. 3. Stepwise multiple linear regressions on log-transformed data revealed that the biomass of Daphnia, Bosmina, Ceriodaphmia and Chydorus were all significantly positively related to PVI and negatively to the abundance of fish or PVI x fish. The average individual biomass of cladocerans was negatively related to fish, but unrelated to PVI. Calculated zooplankton grazing pressure on phytoplankton was positively related to PVI and negatively to PVI x fish. Accordingly the phytoplankton biovolume was negatively related to PVI and to PVI x zooplankton biomass. Cyanophytes and chryptophytes (% of biomass) were positively and Chlorococcales and diatoms negatively related to PVI, while cyanophytes and Chlorococcales were negatively related to PVI x zooplankton biomass. In contrast diatoms and cryptophytes were positively related to the zooplankton biomass or PVI x zooplankton. 4. The results suggest that fish predation has less impact on the zooplankton community in the more structured environment of macrophyte beds, particularly when the PVI exceeds 15–20%. They further suggest that the refuge capacity of macrophytes decreases markedly with increasing fish density (in our study above approximately 10 CPUE). Provided that the density of planktivorous fish is not high, even small improvements in submerged macrophyte abundance may have a substantial positive impact on the zooplankton, leading to a lower phytoplankton biovolume and higher water transparency. However, at high fish densities the refuge effect seems low and no major zooplankton mediated effects of enhanced growth of macrophytes are to be expected.     

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Top-down and bottom-up community regulation in marine rocky intertidal habitats

Strong top-down control by consumers has been demonstrated in rocky intertidal communities around the world. In contrast, the role of bottom-up effects (nutrients and productivity), known to have important influences in terrestrial and particularly freshwater ecosystems, is poorly known in marine hard-bottom communities. Recent studies in South Africa, New England, Oregon and New Zealand suggest that bottom-up processes can have important effects on rocky intertidal community structure. A significant aspect of all of these studies was the incorporation of processes varying on larger spatial scales than previously considered (10’s to 1000’s of km). In all four regions, variation in oceanographic factors (currents, upwelling, nutrients, rates of particle flux) was associated with different magnitudes of algal and/or phytoplankton abundance, availability of particulate food, and rates of recruitment. These processes led to differences in prey abundance and growth, secondary production, consumer growth, and consumer impact on prey resources. Oceanographic conditions therefore may vary on scales that generate ecologically significant variability in populations at the bottom of the food chain, and through upward-flowing food chain effects, lead to variation in top-down trophic effects. I conclude that top-down and bottom-up processes can be important joint determinants of community structure in rocky intertidal habitats, and predict that such effects will occur generally wherever oceanographic ‘discontinuities’ lie adjacent to rocky coastlines. I further argue that increased attention by researchers and of funding agencies to such benthic–pelagic coupling would dramatically enhance our understanding of the dynamics of marine ecosystems.     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

Effects of an invasive consumer on zooplankton communities are unaltered by nutrient inputs

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Dietary changes in predators and scavengers in a nocturnally illuminated riparian ecosystem

Aquatic and terrestrial ecosystems are linked by fluxes of carbon and nutrients in riparian areas. Processes that alter these fluxes may therefore change the diet and composition of consumer communities. We used stable carbon isotope (δ¹³C) analyses to test whether the increased abundance of aquatic prey observed in a previous study led to a dietary shift in riparian consumers in areas illuminated by artificial light at night (ALAN). We measured the contribution of aquatic‐derived carbon to diets in riparian arthropods in experimentally lit and unlit sites along an agricultural drainage ditch in northern Germany. The δ¹³C signature of the spider Pachygnatha clercki (Tetragnathidae) was 0.7‰ lower in the ALAN‐illuminated site in summer, indicating a greater assimilation of aquatic prey. Bayesian mixing models also supported higher intake of aquatic prey under ALAN in spring (34% versus 21%). In contrast, isotopic signatures for P. clercki (0.3‰) and Pardosa prativaga (0.7‰) indicated a preference for terrestrial prey in the illuminated site in spring. Terrestrial prey intake increased in spring for P. clercki under ALAN (from 70% to 74%) and in spring and autumn for P. prativaga (from 68% to 77% and from 67% to 72%) and Opiliones (from 68% to 72%; 68% to 75%). This was despite most of the available prey (up to 80%) being aquatic in origin. We conclude that ALAN changed the diet of riparian secondary consumers by increasing the density of both aquatic and terrestrial prey. Dietary changes were species‐ and season‐specific, indicating that the effects of ALAN may interact with phenology and feeding strategy. Because streetlights can occur in high density near freshwaters, ALAN may have widespread effects on aquatic–terrestrial ecosystem linkages.     

Positive indirect effect of aquatic insects on terrestrial prey is not offset by increased predator density

1. Changes in one prey species' density can indirectly affect the abundance of another prey species if a shared predator eats both species. Sometimes, indirect effects occur when prey straddle habitats, including when riparian predator populations grow in response to emergent aquatic insects and increase predation on terrestrial prey. However, predators may largely switch to aquatic insects or become satiated, reducing predation on terrestrial prey. 2. To determine the net indirect effect of aquatic insects on terrestrial arthropods via generalist spider predators, a field experiment was conducted mimicking midge influx and a wolf spider numerical response inside enclosures near an Icelandic lake. Lab mesocosms were also used to assess per capita rates of spider predation u nder differing levels of midge abundance. 3. Midges always decreased sentinel prey predation, but this effect increased with predator density. When midges were absent, predation increased 30% at a high spider density, but predation was equal between spider treatments when midges were present. In situ arthropods showed no effect of midge or spider treatments, although non‐significant abundance patterns were observed congruent with sentinel prey results. 4. In lab mesocosms, prey survivorship increased ≥50% where midges were present and rapidly saturated; the addition of 5, 20, 50, and 100 midges equivalently reduced spider predation, supporting predator distraction rather than satiation as the root cause. 5. The present results demonstrate a strong positive indirect effect of midges and broadly support the concept that predator responses to alternative prey are a major influence on the magnitude and direction of predator‐mediated indirect effects.     

上海滴水湖两种枝角类牧食对浮游植物群落影响的模拟研究

于2012年9月通过浮游动物添加实验,研究了大型溞(Daphnia magna)和隆腺溞(Daphnia carinata)对上海滴水湖浮游植物群落结构的影响.结果表明,实验组氨氮和可溶性活性磷浓度与空白组相比显著增加,大型溞组浮游植物密度较空白组降低了70.3％,隆腺溞组浮游植物密度较空白组降低了80.0％,叶绿素a浓度分别下降了80.4％和75.2％,叶绿素a与氨氮、可溶性活性磷呈显著相关性.浮游植物的群落结构较空白组也发生较大变化,硅藻、蓝藻和绿藻密度比例明显降低,隐藻、裸藻和甲藻在实验结束时没有检出,说明大型溞和隆线溞能有效控制水体中浮游植物密度,并能够对浮游植物群落结构产生影响.同时,附着藻类密度较空白组也有明显减少,说明大型溞和隆线溞对附着藻类也有一定的牧食作用.     

Effects of artificial ultraviolet-B radiation on experimental aquatic microcosms

1. The effects of prolonged ultraviolet-B (UVB) radiation on freshwater communities were studied in indoor microcosms (600 L) with artificial light sources, simulating a clear, shallow, mesotrophic aquatic ecosystem. A range of six intensities (in duplicate) of UVB radiation, ranging from 0 (control) to 9.56 kJ m⁻² day⁻¹ at the water surface, was applied for 8 weeks. The UVB radiation levels, attenuation, shading and scattering were comparable to those in Dutch shallow freshwater systems. Physical, chemical and biological variables were monitored weekly.
2. The UVB treatment did not affect the abundance, species composition or biovolume of the phytoplankton or zooplankton communities, nor did it affect the periphyton or the macroinvertebrate community. A few species showed a significant response on some of the sampling dates, but there was no negative UVB effect at the community level. Overall, the ecosystems in the microcosms were not affected by the UVB treatment.
3. In a bio-assay, a laboratory clone of Daphnia pulex , not subjected to UVB radiation, was fed with seston from the microcosms. Daphnia pulex feeding on seston from the control microcosms grew faster, had better survival and better reproduction than D. pulex feeding on seston from the UVB treated microcosms. The phytoplankton–zooplankton interaction may have been influenced by the UVB treatment.
4. The dissolved oxygen content (DOC) concentrations in the microcosms were around 5 mg L^−1. The DOC levels in Dutch systems rarely fall below 10 mg L^−1. This might provide sufficient protection against the detrimental effects of increased UVB radiation.     

Predation by an exotic lizard,Anolis sagrei,alters the ant community structure in betelnut palm plantations in southern Taiwan

Abstract 1. Predators can affect prey directly by reducing prey abundance and indirectly by altering behavioural patterns of prey. From previous studies, there is little evidence that ant community structure is affected by vertebrate predation. 2. Researchers tend to consider the interactions between vertebrate predators and ants to be weak. The present study examined the impact of the exotic invasive lizard, Anolis sagrei, on the ant community structure by manipulating the density of lizards within enclosures. The natural density of A. sagrei in the field was surveyed and used as the stocking density rate in the lizard‐present sub‐enclosures. 3. Before the lizard density was manipulated, there was no difference in the ant diversity between sub‐enclosures. After the lizard density manipulation, the ant diversity in sub‐enclosures with A. sagrei present was significantly different from that of enclosures where the lizards were absent, although the overall ant abundance did not differ significantly. 4. The ant diversity difference was generated by a significant reduction of the ant species Pheidole fervens in sub‐enclosures with A. sagrei present. Such an abundance change might be the result of direct predation by the lizards, or it might be generated by a foraging site shift by this ant. 5. The results of this study thus demonstrated that the invasion of an exotic vertebrate can significantly alter the community structure of ants, perhaps through the combined direct and indirect effects of lizards on ants.     

淮河中游浮游甲壳动物群落结构的季节动态

2007年1-12月对淮河中游浮游甲壳动物群落结构的季节动态进行研究。共记录浮游甲壳动物24种, 其中枝角类8属13种、桡足类9属11种。枝角类在4月和9月形成两个峰值, 即(28.2±21.6) ind/L和(40.8±10.1) ind/L, 其优势种分别为僧帽溞 Daphnia cucullata和脆弱象鼻溞 Bosmina fatalis。捕食性桡足类-近邻剑水蚤Cyclops vicinus vicinus、广布中剑水蚤Mesocyclops leuckarti和台湾温剑水蚤Thermocyclops taihokuensis分别在4月、5月和6月形成较大的密度。汤匙华哲水蚤Sinocalanus dorrii和中华窄腹剑水蚤Limnoithona sinensis分别在5月和8月占优势。小型浮游植物(≤20 μm)生物量在4月达到最大值, 之后快速下降, 而较大型浮游植物(>20 μm)生物量从4月上升, 到7月达到最大值。典型冗余分析(RDA)显示, 溞属Daphnia的仲春下降与捕食性桡足类(尤其是近邻剑水蚤)的摄食压力、浮游植物生物量的季节变化密切相关。