Relative impacts of Daphnia grazing and direct stimulation by fish on phytoplankton abundance in mesocosm communities

• 1 Planktivorous fish were hypothesised to influence the abundance of algal biomass in lakes by changing zooplankton grazing, affecting zooplankton nutrient recycling and by direct recycling of nutrients to phytoplankton. The relative roles of direct fish effects vs. zooplankton grazing were tested in mesocosm experiments by adding to natural communities large grazing zooplankton (Daphnia carinata) and small planktivorous fish (mosquitofish or juveniles of Australian golden perch).
• 2 The addition of Daphnia to natural communities reduced the numbers of all phytoplankton less than 30 µm in size, but did not affect total biomass of phytoplankton as large Volvox colonies predominated.
• 3 The addition of Daphnia also reduced the abundance of some small (Moina, Bosmina, Keratella) and large (adult Boeckella) zooplankton, suggesting competitive interactions within zooplankton.
• 4 The addition of mosquitofish to communities containing Daphnia further reduced the abundance of some small zooplankton (Moina, Keratella), but increased the numbers of Daphnia and adult Boeckella. In spite of the likely increase in grazing due to Daphnia, the abundance of total phytoplankton and dominant alga Volvox did not decline in the presence of mosquitofish but was maintained at a significantly higher level than in control.
• 5 The addition of juveniles of golden perch to communities containing Daphnia reduced the abundance of small zooplankton (Moina), increased the abundance of large zooplankton (adult Boeckella) but had no significant effect on Daphnia and total phytoplankton abundance.
• 6 The results of the present study suggest that some planktivorous fish can promote the growth of phytoplankton in a direct way, probably by recycling nutrients, and even in the presence of large grazers. However, the manifestation of the direct effect of fish can vary with fish species.

     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

Vertical and seasonal distributions of micro-organisms,zooplankton and phytoplankton in a eutrophic lake

The vertical distributions of bacteria, picoalgae,protozoan and metazoan zooplankton, and phytoplanktonin the highly eutrophic Lake Köyliönjärvi(SW Finland) were studied monthly during the period ofice-cover in January-April 1996. For comparison, wealso provide some data on the distributions of theplankton during the summer. The whole watercolumn remained oxic during the ice-covered period,although the near-bottom oxygen concentrations werealways very low. The heterotrophic nanoflagellateswere more abundant in winter than in summer, butciliates, picoalgae and bacteria were more numerous insummer. In general both zooplankton and phytoplanktonhad low biomass during the ice-covered period.However, the biomass of the diatom Aulacoseiraislandica ssp. islandica was high under the icein April. The calanoid copepod Eudiaptomusgraciloides was the dominant zooplankton species fromJanuary to March, but had almost disappeared by thebeginning of April and did not increase again until inJune. The dominant rotifer species in winterwere Keratella cochlearis, Filinia terminalis,and Filinia longiseta in the surface water andRotaria neptunia near the bottom.     

Effects of zooplankton on sedimentation in pelagic ecosystems: Theory and test in two lakes of the Canadian shield

A simple heuristic theory based on conservation of matter and describing the fate of autotrophic production in pelagic ecosystems was developed to assist in conceptualizing how zooplankton affect sedimentation processes in lakes and oceans. The theory predicted that effects of zooplankton on the fraction of autotrophic incorporation that sediments (the export ratio) will be a function of factors related both to zooplankton digestion and egestion and to tendencies of particulate matter to sediment directly prior to mineralization. As a result, effects of zooplankton grazing on the export ratio were predicted to be site-dependent, a function of physical conditions and zooplankton communities characterizing an ecosystem. The theory was tested by monitoring autotrophic production of C, N, and P, sedimentation of C, N, and P, and zooplankton biomass in two lakes of the Canadian shield characterized by contrasting morphometry and food-web structure. In Lake 110, a small elongate lake protected from wind exposure, export ratios of C, N, and P declined strongly with zooplankton biomass. In contrast, in L240, a larger lake with considerably greater wind exposure, export ratios increased with zooplankton biomass. These results were consistent with predictions of our theory that effects of zooplankton on sedimentation processes will depend on the tendency of particulate matter to directly sediment relative to the tendency of egested materials to sediment. However, no significant differences in relationships between export ratios for C, N, and P and zooplankton biomass were found.     

A bioassay study of effects of zooplankton,iron and NTA on the phytoplankton productivity of a marl lake

Additions of iron and NTA had minor stimulatory effects on the phytoplankton productivity of samples of water from two calcareous Michigan lakes in autumn and winter contrary to effects at other seasons. Added Daphnia pulex significantly reduced phytoplankton productivity by grazing at this time of year, in all tests at all levels of addition from 4–32 animals/l, greatly in excess of any possible beneficial effects of increased nutrient availability.     

Relations between planktivorous fish abundance,zooplankton and phytoplankton in three lakes of differing productivity

Hydrobiologia - Water chemistry, phytoplankton, zooplankton and fish populations werestudied over several years in three shallow, non-stratified lakes withdiffering nutrient loadings and fish...     

The dynamics and role of limnetic zooplankton in Loosdrecht lakes (The Netherlands)

The paper summarizes the results of a ten-year (1981–1991) zooplankton research on the Lake Loosdrecht, a highly eutrophic lake. The main cause of the lake's eutrophication and deteriorating water quality was supply up to mid 1984 of water from the River Vecht. This supply was replaced by dephosphorized water from the Amsterdam-Rhine Canal in 1984. The effects of this and other restoration measures on the lake's ecosystem were studied. Despite a reduction in the external P-load from ca. 1.0 g P m^–2 y^–1 to ca. 0.35 g m^–2 y^–1 now, the filamentous prokaryotes, including cyanobacteria and Prochlorothrix, continue to dominate the phytoplankton.Among the crustacean plankton Bosmina spp, Chydorus sp. and three species of cyclopoid copepods and their nauplii are quite common. Though there was no major change in the composition of abundant species, Daphnia cucullata, which is the only daphnid in these lakes, became virtually extinct since 1989. Among about 20 genera and 40 species of rotifers the important ones are: Anuraeopsis fissa, Keratella cochlearis, Filinia longiseta and Polyarthra. The rotifers usually peak in mid-summer following the crustacean peak in spring. The mean annual densities of crustaceans decreased during 1988–1991. Whereas seston (< 150 µm) mean mass in the lake increased since 1983 by 20–60%, zooplankton (> 150 µm) mass decreased by 15–35%.The grazing by crustacean community, which was attributable mainly to Bosmina, had mean rates between 10 and 25% d^–1. Between 42 and 47% of the food ingested was assimilated. In spring and early summer when both rotifers and crustaceans have their maximal densities the clearance rates of the rotifers were much higher. Based on C/P ratios, the zooplankton (> 150 µm) mass contained 2.5 times more phosphorus than seston (< 150 µm) mass so that the zooplankton comprised 12.5 % of the total-P in total particulate matter in the open water, compared with only 4.5% of the total particulate C. The mean excretion rates of P by zooplankton varied narrowly between 1.5 and 1.8 µg P 1^– d^–1, which equalled between 14 and 28% d^–1 of the P needed for phytoplankton production.The lack of response to restoration measures cannot be ascribed to one single factor. Apparently, the external P-loading is still not low enough and internal P-loading, though low, may be still high enough to sustain high seston levels. Intensive predation by bream is perhaps more important than food quality (high concentrations of filamentous cyanobacteria) in depressing the development of large-bodied zooplankton grazers, e.g. Daphnia. This may also contribute to resistance of the lake's ecosystem to respond to rehabilitation measures.     

The relation between bacterial carbon and dissolved humic compounds in oligotrophic lakes

Abstract The mean annual biomass of planktonic bacteria showed large variations both within and between lakes. The lowest bacterial biomass was found in acidified lakes (7.8–12.1 μg C · 1⁻¹), and tended to increase with increasing water colour (up to 44.1 μg C · 1⁻¹). The highest recorded bacterial biomass was 138 μg C · 1⁻¹. The mean annual bacterial biomass equalled 23–45% of the algal biomass. Zooplankton biomass was high, compared to algal biomass (40–50%). Multiple regression analysis of 10 variables showed a strong positive correlation between bacterial biomass and humic content ( r = 0.74, P < 0.001), while other parameters, except pH, showed no correlation. The observation thus strongly supports the role of humic compounds in aquatic secondary production.     

Structure and dynamics of zooplankton communities in the littoral zone of some North Carolina lakes

Species composition and seasonal, dynamics of zooplankton in the littoral zone were studied in three piedmont North Carolina lakes for one year. Nygaard's compound index indicated oligotrophic conditions in Belews Lake and eutrophic conditions in Salem Lake and Lake 150. A total of 32 species of zooplankton were found in the samples. Significantly fewer species of truely littoral zooplankton were present in the oligotrophic lake regardless of season (P < 0.05). Eutrophication appears to favor increased diversity in zooplankton of the littoral zone. Habitat heterogeneity, provided by a well developed littoral zone containing aquatic macrophytes, may be the major factor contributing to the maintenance of more taxa of zooplankton in eutrophic systems.Distinct taxa of Cladocera and Copepoda are present in the littoral zone, exhibiting seasonal dynamics that are largely different from the limnetic fauna. Such dynamics may result from predation by characteristically limnetic species of cyclopoid copepods which seek prey in the littoral zone. The numerical and seasonal abundance of predators and prey are inversely related and the true littoral taxa, represented in the present study by the genera Alona, Chydorus, Pleuroxus, Sida, Simocephalus, and Eucyclops, contained no predaceous species. The seasonal dynamics of these genera are very similar even in widely separated geographical regions, indicating that the factor(s) responsible operate independently of climatic and chemical variables. Intrazooplankton predation appears to be a more plausable explanation than segregation along depth which, although consistent with data from studies of limnetic waters, cannot be related to populations of zooplankton in shallow littoral areas.     

Influence on phytoplankton biomass in lakes of different trophy by phosphorus in lake water and its regeneration by zooplankton

In 49 unpolluted lakes of north-eastern Poland the biomass of algae in summer is significantly related to the concentration of total phosphorus and to the rate of phosphorus regeneration by zooplankton. Using a model with equations describing these relationships, the biomass of blue-green algae and other phytoplankton groups was predicted for 14 polluted lakes. A good approximation of actual values was obtained only for the biomass of blue-green algae calculated from the estimated rate of P regeneration by zooplankton in these lakes. It is hypothesized that more-or-less edible algae of other classes did not show dependence on the rate of input of regenerated P because their biomass was heavily reduced by grazing of zooplankton.     

The outcome of the invasion of Florida lakes by Daphnia lumholtzi

1. We examined 7–12 years of monthly to quarterly historical data from 15 lakes in FL, U.S.A. to determine the extent and outcome of invasion by the alien cladoceran Daphnia lumholtzi Sars. 2. The alien species was found in 10 of the 15 lakes, including Florida’s three largest lakes: Okeechobee, George and Apopka. All the surveyed lakes had resident populations of the smaller native species Daphnia ambigua Scourfield. 3. In most of the lakes, D. ambigua occurred seven to ninefold more often in plankton samples than D. lumholtzi, and at 10‐ to 100‐fold higher maximal densities. One exception was a small lake in central Florida (Lake Jesup), where D. lumholtzi attained high densities on several occasions in the 10 years of sampling. 4. In Lake Okeechobee, where data were of sufficient quality and quantity to perform statistical analyses, the results of canonical correlation analysis indicated that high densities of D. lumholtzi were correlated with lower concentrations of suspended solids, high algal biomass and higher temperature, whereas the opposite conditions were correlated with high densities of D. ambigua. 5. Based on the majority of data, D. lumholtzi has successfully invaded many lakes in Florida, yet it has not become a substantive component of the zooplankton. Additional research is needed to determine whether resources, fish predation or some other factor is responsible for this outcome of invasion.     

Midsummer zooplankton assemblages in four types of wetlands in the Upper Midwest,USA

The objective of this study was to characterize the zooplankton and phytoplankton assemblages of four different types of wetlands and to evaluate their use as environmental indicators. Total abundances, community composition, and species diversity were evaluated for zooplankton and phytoplankton assemblages from 24 wetlands and related to water quality variables. During August 1995, six representative sites were sampled from four types of wetlands designated as constructed, impacted, non-impacted, or temporary. The plankton assemblages of all wetlands were dominated by cosmopolitan crustacean, rotifer, and phytoplankton taxa typical of lake plankton communities. Species diversity, richness, and evenness of zooplankton and phytoplankton assemblages did not differ significantly among the wetland types. Total zooplankton abundance was significantly (p < 0.01) related to chlorophyll a and total phosphorus concentrations over the range of trophic conditions. Mean zooplankton densities and phytoplankton biovolumes were similar among the wetlands, however, the relative abundances of major zooplankton groups differed among the wetland types. Cyanophytes, primarily Oscillatoria spp., were a major component of the phytoplankton across all four wetland types, and were significantly more abundant within the constructed and temporary sites. On average, rotifers accounted for 79% of total zooplankton abundance within the constructed wetlands and were much less dominant in the non-impacted and temporary wetlands. Cladoceran, copepodite, and adult copepod concentrations were low in the constructed and impacted wetlands and increased in the non-impacted and temporary wetlands in conjunction with increased chlorophytes and cryptophytes. Our preliminary survey suggests that abiotic factors which are known to directly affect phytoplankton may indirectly affect zooplankton composition in such a way as to use zooplankton assemblages as indicators of water quality. However, further study incorporating seasonal dynamics and the influence of predators on zooplankton assemblages is needed to fully assess the use of zooplankton community composition as an environmental indicator for wetland systems. This revised version was published online in July 2006 with corrections to the Cover Date.     

Calcium content of crustacean zooplankton and its potential role in species distribution

SUMMARY 1. The occurrence and species composition of crustacean zooplankton in Norwegian lakes was related to ambient Ca concentrations, pH, total phosphorus and chlorophyll a , and to the presence/absence of other zooplankters and a planktivorous fish (roach: Rutilus rutilus ).
2. Ambient Ca concentrations appeared to influence the distribution of Daphnia species: Daphnia longispina often occurred in Ca-rich lakes with low fish-predation pressure while the smaller Daphnia cristata often occurred in opposite conditions.
3. Body Ca contents were measured in zooplankters from a wide range of localities, to examine Ca requirements and thus the potential for Ca-limitation of common species.
4. All Daphnia species had relatively high specific Ca contents [ranging from 0.8 to 4.4% Ca dry weight (DW)⁻¹] compared with other cladocerans and also copepods (ranging from 0.1 to 0.4% Ca DW⁻¹). Within the Daphnia genus, the specific Ca content increased with increasing body size of the species, and thus the large-bodied species had especially high Ca demands.
5. Because of their high Ca demands, species of Daphnia could be competitively disadvantaged in softwater lakes relative to less Ca-demanding species.     

Steemann Nielsen and the zooplankton

E. Steemann Nielsen is remembered by most biological oceanographers and limnologists as having introduced the ¹⁴C method for measuring photosynthesis in 1952. The present paper is to recall that he was interested in the phytoplankton as part of the plankton community and was much aware of the role of grazing in affecting, if not determining, the concentrations of phytoplankton and, thus, also its rate of production. His principal statements to this effect were made with the open, oligotrophic subtropical and tropical oceans in mind where phytoplankton concentrations exhibit little seasonal change. This paper shows that Steemann Nielsen's sentiment also applies to non-static situations, especially phytoplankton blooms. Of the blooms in Cushing's North Sea Calanus patches of 1949 and 1954 and the two low-latitude, open-sea iron fertilization experiments (IronEx I, II) of the 1990s, more than half or even most of the newly formed cells were lost daily. In these examples, the same water was revisited, mixing was considered, and sinking was an unimportant loss term, so that grazing was the principal cause of mortality. Because of the grazing losses and the subsequent regeneration the CO₂ draw down in the fertilized water was much lower than the ¹⁴C uptake. Moreover the examples show that over the course of the blooms, the rate and even the sign of temporal change of phytoplankton abundance had little relation to the rate of cell division, as already postulated by Riley's 1946 model of the seasonal cycle of phytoplankton on Georges Bank. Thus, in most situations in the open sea and, presumably, large lakes, the rates of cell division (instead of photosynthesis by itself) and of mortality (most often from grazing) are needed for understanding and predicting the temporal change of phytoplankton abundance, a principal goal of biological oceanography. The mechanism maintaining the actual abundance of phytoplankton in the quasi-steady state prevailing over most of the ocean much of the time is still unclear.     

Evidence that fish structure the zooplankton communities of turbid lakes and reservoirs

1. Visually foraging fish typically exclude large zooplankton from clear‐water lakes and reservoirs. Do fish have the same effect in turbid waters, or does turbidity provide a refuge from visual predation? 2. To test the hypothesis that fish exclude large zooplankton species from turbid sites, I searched for populations of medium or large Daphnia species in turbid, fish‐containing reservoirs of south‐central Oklahoma and north‐central Texas, U.S.A., and surveyed the literature for accounts of Daphnia species in turbid habitats worldwide. 3. Only small Daphnia species and the exuberantly spined Daphnia lumholtzi were detected in the turbid reservoirs. The Daphnia species in the reservoirs are smaller than other Daphnia species that occur in the area but were not detected. An extensive survey of the literature suggests that large Daphnia may be found in the lakes of extreme turbidity [Secchi disk depth (SD) < 0.2 m] but that only small and spiny Daphnia are likely to occur in more typical turbid locations (1.0 m > SD > 0.2 m) unless some additional factor reduces the influence of fish predation in such sites. 4. The field samples from Texas and Oklahoma together with the literature review suggest that the effect of visually foraging planktivorous fish on the size structure of turbid‐water zooplankton communities may often be as strong or even stronger than the effect of fish on clear‐water zooplankton communities.     

The seasonality of phytoplankton in African lakes

Although some study of the subject began in 1899, wide-ranging information from African water-bodies has only become available since 1950. Important developments included the establishment of long-term centres of research, the adoption of improved methods for quantitative algal sampling, the more intensive study of environmental conditions, the beginnings of experimental testing, and the improvement of taxonomic knowledge.At higher latitudes (> 20 °) examples of pronounced algal seasonality are long-established; they are accompanied and influenced by marked changes in radiant energy income and so water temperature, and often by effects of seasonal water input. Illustrations are given from lakes in Morocco and South Africa.More generally in Africa, including the tropical belt, annual patterns of phytoplankton seasonality are usually either dominated by hydrological features (water input-output) or by hydrographic ones (water-column structure and circulation). Examples of both types are discussed, together with instances (e.g. L. Volta) of combined hydrological and hydrographic regulation. In both the seasonal abundance of diatoms is often distinct and complementary to that of blue-green algae, with differing relationships to vertical mixing and water retention.Horizontal variability in the seasonal cycle is especially pronounced in the larger or morphometrically subdivided lakes. Some inshore-offshore differentiation is also known to affect phytoplankton quantity (e.g. L. George) and species composition (e.g. L. Victoria). Longitudinal differentiation is common in elongate basins especially when with a massive or seasonal inflow at one end (e.g. L. Turkana, L. Nubia, L. Volta); occasional terminal upwelling can also be influential (e.g. southern L. Tanganyika). Such examples grade into the longitudinally differentiated seasonality of flowing river-reservoir systems, as studied on the Blue and White Niles.The annual amplitude of population density, expressed in orders of magnitude (=log₁₀ units), is one measure of seasonal variability. It can exceed 3 orders both in systems subject to hydrological wash-out (e.g. Nile reservoirs) and in the more variable species components of lakes of long retention (e.g. L. Victoria). Low amplitudes can be characteristic of some components (e.g. green algae in L. Victoria) or of total algal biomass (e.g. L. George, L. Sibaya).Seasonal changes may be subordinated to inter-annual ones, especially in shallow and hydrologically unstable lakes (e.g. L. Nakuru).     

Evidence of coprophagy in freshwater zooplankton

1. Vertical transport of nutrients in sedimenting faecal material is greatly reduced by coprophageous organisms. Unfortunately, nearly all work on faecal production, sedimentation and coprophagy has dealt with copepods in marine ecosystems. Here, we report the first evidence of coprophagy in freshwater zooplankton from oligotrophic and eutrophic lakes. We used ¹⁴C‐labelled algae and faecal material to estimate the rates of algal clearance and coprophagy. 2. Measured feeding rates per individual on faecal material were similar (Daphnia pulex, D. rosea, Leptodiaptomus tyrelli) or even higher (D. lumholtzi) than filtering rates on phytoplankton. This finding does not necessarily implicate active selection of faeces over algae because: (i) we did not use the same food concentrations for faeces and algae, and (ii) grazers of slightly different sizes were used in each test. 3. Weight‐specific clearance rates of L. tyrelli and Holopedium gibberum on faecal matter (0.084–0.089 mL μg^?1 h^?1) were higher than in the daphniids (0.026 mL μg^?1 h^?1). 4. The data indicate that coprophagy in freshwater ecosystems is an important mechanism of nutrient recycling, and this process should be taken into account when studying nutrient fluxes within lakes and reservoirs.     

The effects of Daphnia on nutrient stoichiometry and filamentous cyanobacteria: a mesocosm experiment in a eutrophic lake

1. Stoichiometric theory predicts that the nitrogen : phosphorus (N : P) ratio of recycled nutrients should increase when P‐rich zooplankton such as Daphnia become dominant. We used an enclosure study to test the hypothesis that an increased biomass of Daphnia will increase the relative availability of N versus P sufficiently to decrease the abundance of filamentous cyanobacteria. The experiment was conducted in artificially enriched Lake 227 (L227) in the Experimental Lakes Area (ELA), north‐western Ontario, Canada. Previous studies in L227 have shown that the dominance of filamentous, N‐fixing cyanobacteria is strongly affected by changes in the relative loading rates of N and P. 2. We used a 2 × 2 factorial design with the addition or absence of D. pulicaria and high or low relative loading rates of N and P (+NH₄, –NH₄) in small enclosures as treatment variables. If Daphnia can strongly affect filamentous cyanobacteria by altering N and P availability, these impacts should be greatest with low external N : P loading rates. The phytoplankton community of L227 was predominantly composed of filamentous Aphanizomenon spp. at the start of the experiment. 3. Daphnia strongly reduced filamentous cyanobacterial density in all enclosures to which they were added. The addition of NH₄ had only a small impact on algal community composition. Hence, we conclude that Daphnia did not cause reductions in cyanobacteria by altering the N : P ratio of available nutrients. 4. Despite the lack of evidence that Daphnia affected filamentous cyanobacteria by altering the relative availability of N and P, we found changes in nutrient cycling consistent with other aspects of stoichiometric theory. In the presence of Daphnia, total P in the water column decreased because of an increase in P sedimentation. In contrast to P, a decrease in suspended particulate N was offset by an increase in dissolved N (especially NH₄). Hence, dissolved and total N : P ratios in the water column increased with Daphnia as a result of differences in the fate of suspended particulate N versus P. There was minimal accumulation and storage of P in Daphnia biomass in the enclosures. 5. Our experiment demonstrated that Daphnia can strongly limit filamentous cyanobacterial abundance and affect the biogeochemical cycling of nutrients. In our study, changes in nutrient cycling were apparently insufficient to cause the changes in phytoplankton community composition that we observed. Daphnia therefore limited filamentous cyanobacteria by other mechanisms.     

Predicted warming and browning affect timing and magnitude of plankton phenological events in lakes: a mesocosm study

1. Aquatic ecosystems in Northern Europe are expected to face increases in temperature and water colour (TB) in future. While effects of these factors have been studied separately, it is unknown whether and how a combination of them might affect phenological events and trophic interactions. 2. In a mesocosm study, we combined both factors to create conditions expected to arise during the coming century. We focused on quantifying effects on timing and magnitude of plankton spring phenological events and identifying possible mismatches between resources (phytoplankton) and consumers (zooplankton). 3. We found that the increases in TB had important effects on timing and abundance of different plankton groups. While increased temperature led to an earlier peak in phytoplankton and zooplankton and a change in the relative timing of different zooplankton groups, increased water colour reduced chlorophyll‐a concentrations. 4. Increased TB together benefitted cladocerans and calanoid copepods and led to stronger top‐down control of algae by zooplankton. There was no sign of a mismatch between primary producers and grazers as reported from other studies. 5. Our results point towards an earlier onset of plankton spring growth in shallow lakes in future with a stronger top‐down control of phytoplankton by zooplankton grazers.     

Variation in spatial and temporal gradients in zooplankton spring development: the effect of climatic factors

1. We examined the temporal and spatial heterogeneity of zooplankton in lake surface waters during the spring of 3 years in Lake Washington, U.S.A., a large lake with a high production of sockeye salmon fry. 2. We show large within‐season and among‐year variation in the horizontal distribution of temperature, chlorophyll a concentration, and zooplankton in the lake. The main pattern, a delay in zooplankton population increase from the north‐ to the south‐end of the lake, recurred in each year and was persistent within each spring. 3. The delay is primarily caused by the development of a temperature gradient during spring warming, as cold mountain water enters the south end of the lake, while warm water enters the north end via a river draining a nearby lake. Climate factors, such as air temperature and precipitation during winter and spring, appear to influence the extent of the delay of zooplankton increase. 4. If the climate continues to warm, the temporal disconnection in zooplankton development between lake areas immediately influenced by cold river inflow and areas that are influenced by spring warming may increase in magnitude. Thus, the different areas of the lake may not contribute equally to fish production.