Elemental composition of two coexisting Daphnia species during the seasonal course of population development in Lake Constance

Summary In 1986 and 1987 the seasonal course of population density, life history traits and biomass composition (CHN-elemental and ash) of two coexisting Daphnia species from L. Constance (Überlinger See) were studied. The two daphnids are morphologically similar, but demonstrate different seasonal variation of behaviour (i.e. diurnal vertical migration). With the exception of nitrogen content, the mean biomass composition of adults and that of eggs were significantly different: the dry biomass of eggs (adults) contained on average 55.6 (46.1) % C, 7.9 (6.5) % H, 9.7 (9.7) % N, and 10.0 (23.8) % ash. These values were similar to those of other daphnids; it may be concluded that the average daphnid composition is not species specific, but is determined mainly by the position in the life cycle and by the nutritional state of the animal. A distinct seasonal variation in the elemental composition of the biomass of adult daphnids was observed. It related significantly to the concentration of food, as indicated by the epilimnetic chlorophyll-a concentrations and by Secchi depth. From the amplitude of seasonal variation of elemental composition and from the shape of relationships between the elemental composition of daphnids and their food, it can be concluded that in summer the non-migrating D. galeata lives under stronger food limitation than the migrating H. hyalina. In spring, there were no differences in the biomass composition of either species. In summer, the adult fcmales of D. galeata contained significantly less carbon, and hydrogen than those of D. hyalina. In most parameters of biomass (i.e. biomass composition of adults) and of life history (i.e. clutch size, mean adult weight), D. galeata displayed higher coefficients of variation. A tendency in D. galeata to have peaks of population development carlier in the season and in D. hyalina to have them towards autumn was observed. These contrasting responses of the populations development, the seasonal variability of the biomass composition seem to support the view of Geller (1986) who called D. galeata and D. hyalina exploitative and conservative strategists respectively.This research was supported by the Deutsche Forschungsge-meinschaft within the Sonderforschungsbereich Cycling of Matter in Lake Constance (SFB 248), and by scholarship to R.B. from the Deutscher Akademischer Austauschdienst (DAAD)     

Seasonal Vertical Migration and Aestivation of <Emphasis Type="Italic">Rhyacodrilus hiemalis</Emphasis> (Tubificidae,Clitellata) in the Sediment of Lake Biwa,Japan

The vertical distribution of the tubificid worm Rhyacodrilus hiemalis Ohtaka, the numerically dominant species of oligochaete in the littoral of Lake Biwa, was studied with special reference to seasonal vertical migration in the lake sediment. Monthly collections of lake sediment cores were made using PVC tubes. Core sections of sample sediments ranged from 76 to 117 cm. The vertical distribution of the worms showed no diurnal variation; therefore diel vertical migration was not evident. Seasonal downward migration started in April, and upward migration started in October. From December to March, almost all worms remained in the near-surface sediment layer (surface to 30 cm deep), while from July to September almost all worms remained deeper than 30 cm. However, few individuals migrated deeper than 90 cm. No discontinuous layers were found in grain size composition, water content, loss on ignition, particulate carbon, nitrogen or phosphorus. In deep sediment there was no free oxygen, as evidenced by negative ORP values. For 4 months in summer, R. hiemalis aestivated, probably utilizing anaerobic respiration. It appeared that R. hiemalis moved deeper in the sediment in response to sediment temperature, because sediment temperatures in the deep layers seemed to converge at around 20–21 °C in the summer months. The life history traits of seasonal vertical migration and summer aestivation perhaps arose as an adaptation to the climatic conditions accompanying the geographical origin of R. hiemalis, and they also serve to minimize predation risk during summer when most invertebrate predatory fishes are active.     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.     

Migrations of haemoglobin-rich Daphnia longispina in a small,steeply stratified,humic lake with an anoxic hypolimnion

Migrations of Daphnia longispina were studied in a small humic lake with an exceptionally shallow oxic epilimnion. Horizontal distributions showed clear avoidance of the shoreline, which might be explained by the lower density of predators (Chaoborus sp. and Notonecta sp.) in the central parts of the lake. In early summer all size classes of D. longispina exhibited upward nocturnal vertical migration, descending to the upper hypolimnion in daytime. Later in summer, when the nocturnally migrating Chaoborus sp. had grown large enough to graze on small Daphnia, the latter seemed to shift towards twilight migration. However, large Daphnia individuals showed no synchronized migration; rather their bimodal vertical distributions suggested asynchronous vertical migration. Large individuals showed a particular tendency to concentrate near to the oxycline, close to the dense phytoplankton and bacteria populations in the upper part of the anoxic hypolimnion. According to vertical trap experiments, large D. longispina visited the anoxic hypolimnion and might harvest its abundant food resources. The high haemoglobin content of large individuals seems a specific adaptation to allow access to low oxygen water and hence to maximize grazing potential, in both epi- and hypolimnion, and minimize predation pressure. By staying predominantly in cooler water near the oxycline, Daphnia might also minimize its energy consumption to adjust to low food availability while sustaining a sufficiently high population density to exploit those unpredictable short periods with abundant food which are common in small headwater lakes. It is suggested that migrations of zooplankton are a complex behavioural adaptation which may not be explained by any single factor. In humic lakes with shallow stratification, vertical migrations seem to offer particularly high potential advantages, because of the short distances between dramatically different environments in the water column. In further studies more emphasis should be placed on migrations of individuals rather than populations, and migrations should be considered as a dynamic part of the structure and function of the whole planktonic ecosystem.     

Density and depth variations of Daphnia multilocus genotypes during a summer period in Lake Maarsseveen

The genotype composition of a Daphnia population complex during a summer period in Lake Maarsseveen (The Netherlands) was determined by allozyme analysis. The depth distribution, diel vertical migration and several parameters of the total population were measured. Young-of-the-year (0⁺) perch (Perca fluviatilis) were caught and species and allozyme types of Daphnia in the perch gut were also analysed. During May 1997, the densities of D. hyalina, D. galeata, the back-cross D. g × h — hyalina and the multilocus allozyme genotypes of the hybrid D. g × h decreased, except one multilocus genotype (MMMF). Total population size decreased and the ratio of females with eggs to those without eggs decreased as well. Food limitation during this clear-water phase in the lake is considered responsible. All genotypes, except MMMF, gradually descended in the water column. This drift is thought to be a reaction to the abundantly present 0⁺ perch or to the kairomones of this fish, although predation on the daphnids was still absent. In June, diel vertical migration started, except again part of the MMMF subpopulation. The other part migrated over a short distance compared with the other taxa and allozyme types. Within two weeks, the upper 5 m of the epilimnion was devoid of Daphnia, and guts of perch were predominantly filled with MMMF. The daphnids in the gut and the lake did not differ in allozyme type composition. By the end of July, population density had increased again. The size and composition of the Daphnia population complex continuously changed during the study period, as did the depth distribution of the components. Different genotypes within the population complex seem to have developed different strategies to cope with starvation and predation and the state at a particular moment can be understood only if past and present factors are considered.     

Competition in natural populations of Daphnia

I investigated the competitive relationships between two species of Daphnia, D. galeata and D. cucullata, and their interspecific hybrid. The term hemispecific competition was introduced to describe competition between parental species and hybrids. In eutrophic Tjeukemeer both parental species were found to compete with the hybrid, whereas competition between D. galeata and D. cucullata seemed limited. Although the effect of competition on life history traits of daphnids may be profound, the influence of the competitors on the seasonal dynamics of the Daphnia species seems limited.     

Competitive ability of Daphnia under dominance of non-toxic filamentous cyanobacteria

It is generally assumed that Daphnia is more susceptible to the inhibitory effects of filamentous cyanobacteria than small cladocerans since daphnids have a larger gape size and filtrate the filaments, whereas small cladocerans do not. This study addresses the question whether food limitation has the potential to modify this scenario of cladoceran response to dominance of non-toxic filamentous cyanobacteria. Daphnia galeatawas grown under limited (0.1 mg C l^–1) and unlimited concentrations (1.0 mg C l^–1) of high-quality food algae both in the absence/presence of non-toxic filamentous Aphanizomenon flexuosum. As the effects of these cyanobacteria on D. galeatawere positive under food limiting conditions and negative at the high food density, it was concluded that D. galeatawas mainly affected by nutritional quality due to its ability to ingest the filaments, while mechanical interference with food collection was not important. In competition experiments between D. galeataand Bosmina longirostris, D. galeatawas the dominant species at regular additions of food (1.0 mg C l^–1) in the absence of Aphanizomenon. In the presence of these cyanobacteria, D. galeatawas inhibited during the first days of the experimental period. However, the negative effect at the initially high food density was outweighed by nutrition at food limiting conditions and the outcome in competitive dominance was not changed. The results demonstrate that the ability of D. galeata to ingest large-sized non-toxic cyanobacteria can be considered as advantageous under food limiting conditions.     

Size-dependent habitat choice in Daphnia galeata Sars and size-structured interactions among zooplankton in a subarctic lake (lake Lombola, Norway)

Seasonal changes in vertical distribution of Daphnia galeata and other zooplankters were monitored in lake Lombola (69° 07&prime; N). Depth-habitat use, availability of edible algae and zooplankton densities were recorded to examine seasonal changes in intensity of competition between Daphnia and the other herbivores in the lake. Early in July, the exephippial generation of Daphnia aggregated near the surface, independently of body-size. In late July, when fish planktivory was expected to increase, the daphnids moved down during the day. In August, as intraspecific competition for food intensified, small and large Daphnia partitioned the water column, with larger individuals staying deeper. In September, Daphnia became dominated by large individuals, edible phytoplankton reached the seasonal minimum, and the vertical distribution of Daphnia gradually stretched out towards the surface. The observations on food availability and zooplankton densities suggest that interspecific competition intensified by the end of July. Species and stages that were most exposed to exploitative and interference competition by Daphnia were those staying deeper, because their vertical distribution overlapped more with the larger, competitively superior daphnids. These susceptible competitors included Keratella cochlearis and Synchaeta, among the rotifers, and nauplii and early copepodite stages of Cyclops scutifer. Depth-habitat use is discussed in relation to copepod development, zooplankton dynamics and predator-mediated coexistence.     

Multiple defence strategies of <Emphasis Type="Italic">Daphnia</Emphasis><Emphasis Type="Italic">galeata</Emphasis> against predation in a weakly stratified reservoir

In the presence of size-selective fish daphnids were shown to exhibit two alternative inducible defence strategies: They may either escape predation by active migration or adopt a life history strategy, e.g., reproduce earlier and at a smaller size. Depending on the type of habitat, migration may either be vertically (in deep stratified lakes) or horizontally (in shallow lakes with macrophytes) oriented. Concerning behavioural defence strategies, daphnids living in medium-deep, weakly stratified water bodies with a poorly developed littoral face a dilemma, since the littoral provides no shelter and the availability of a deep-water refuge is unpredictable. We studied the population dynamics, life history changes (size at maturity) and daytime vertical distribution of Daphnia galeata in a weakly stratified reservoir in relation to predation by juvenile fish during 6 years. While temperature gradients were usually small, oxygen concentrations suggest that a low-oxygen refuge for daphnids was available in every year to some extent. Our results indicate that, depending on predation intensity and stratification patterns, daphnids exhibit both, behavioural and life history defences. In years with a high biomass of young-of-the-year (YOY) perch Daphnia abundance declined rapidly at the end of the clear water stage while at the same time the vertical distribution at daytime shifted to deep strata providing a low-oxygen refuge and the size at maturity decreased. However, while the life history response in some years lasted throughout most of the summer period, a shift in daytime vertical distribution was exhibited for much shorter periods. Both traits were much less expressed in years with low YOY fish densities and no negative correlation between them could be verified. We suggest that under high predation pressure in this relatively shallow reservoir no strictly alternative (either behavioural or life history) strategies exist, but that daphnids make use of the full range of possible anti-predator defences available, at least during short periods when predation is most intense. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

The influence of fish kairomones on the induction and vertical distribution of sexual individuals of the Daphnia galeata species complex

To assess the potential production of hybrids and backcrosses in a semi-natural environment, we studied the combined effect of fish kairomone, and food level on the production of males and ephippial females in different clones of five Daphnia taxa from the D. galeata species complex. We also studied the diel vertical migration (DVM) of these sexual daphnids under the same varying conditions. This was done to test the hypothesis that males and ephippial females have different migrating strategies, which would increase their mating probability. The study was carried out in two large-scale indoor mesocosms, the so-called `plankton towers' in the Max-Planck Institute in Plön, Germany.Although all of the Daphnia taxa produced ephippial females in the course of the experiment, only D. galeata produced a significant number of males. Fish kairomones had a significant negative influence on the production of ephippial females. We found no DVM in the D. galeata males. They stayed at a depth between 5 and 6 m both day and night, 1 or 2 m above the thermocline. The ephippial females of D. cucullata x hyalina migrated, whereas ephippial females of the other taxa showed no DVM but came significantly closer to the surface in the presence of fish kairomones. We conclude that males and sexual females co-occur in this species complex both in time and space. Therefore, a regular production of hybrids and backcrosses in this species complex seems likely. Fish kairomones do not seem to significantly influence this process.     

No Evidence for Adaptive Micro-Evolution to a Decrease in Phosphorus-Loading of a Daphnia Population Inhabiting A Pre-Alpine Lake

Many polluted lakes in Europe are being restored and phosphorus concentrations have dropped dramatically in these lakes. We studied the genetic structure of Daphnia galeata over the past 30 years in Lake Greifensee, Switzerland, a period during which the phosphorus concentration in the lake reduced dramatically. Distinct genotypes of D. galeata were hatched from diapausing eggs extracted from six different time horizons in dated sediments. We compared juvenile growth, size and egg production of D. galeata reared on high-P and low-P algae to investigate whether Daphnia have evolved to grow better on phosphorus-limited algae. Our results indicate that life histories of D. galeata differed significantly between both food types. We also found significant clone effects for size and egg number. But we found no significant interaction between the depth from which the clones were selected and food quality. This means that we found no evidence for adaptive micro-evolution in response to P reduction in the lake. We discuss our results in relation to other studies that found evidence for adaptive micro-evolution in comparable time frames.     

Influence of spring warming on the predation rate of underyearling fish on Daphnia– a deterministic simulation approach

1. According to previous field studies in the biomanipulated Bautzen reservoir (Germany), a midsummer decline of the dominating zooplankter, Daphnia galeata, was suggested to be initiated by a simultaneous occurrence of low fecundity of the daphnids and a selective feeding of underyearling fish on mature daphnids. The timing of both processes was assumed to be triggered by spring water temperature. However, the field data were not appropriate for testing whether yearly differences in spring warming are strong enough to control the predation rate of underyearling fish on daphnids.
2. By combining field data on fish growth, feeding and population mortality, the daily uptake of Daphnia by a virtual population of underyearling perch (Perca fluviatilis) was simulated. In addition, the daily predatory mortality of mature daphnids was calculated independently. Scenarios with a warm and a cold spring were compared. Furthermore, the delayed warming of a pelagic zone of a lake versus a littoral one was simulated. Sensitivity of the simulation to changes in five parameters was tested.
3. In both the warm spring scenario and the littoral warming scenario, more daphnids in general and more mature daphnids in particular were eaten, compared with the cold spring and pelagic scenarios. The predatory mortality of mature daphnids was driven by the increasing gape size of growing fish such that in warmer years the fish reach earlier the size at which they can eat mature daphnids.
4. The simulation was most sensitive to changes in daily mortality rate of the fish and to the size at maturity of the daphnids. Since at least the fish mortality is also temperature‐dependent via the growth rates of fish, the predation rate of perch on D. galeata in Bautzen reservoir is substantially increased during a warm spring. This underlines the assumption that even a slight global warming may have a decisive influence on food web processes due to the fine‐scaled patterns of trophic interactions in lakes.     

Vertical distributions of zooplankton and population dynamics of Daphnia in a meromictic lake

Vertical distributions of zooplankton were studied in relation to profiles of temperature, oxygen and chlorophyll a in Roi Lake, a small meromictic lake in central Alberta. Zooplankton were distributed fairly evenly through the oxygenated part of the water column in early summer, but a gradual descent of several species became evident in June. The vertical distribution of chlorophyll was dominated by a huge peak at the 8- to 9-m-deep chemocline. the location of a plate of photosynthetic sulfur bacteria. Ambient concentration of chlorophyll was a poor predictor of the numbers of zooplankton and the fecundity of Daphnia pulicaria at different depths, and per capita birth rates of Daphnia were usually highest in the surface waters. The reproductively disadvantageous restriction of daphnids to deep water by late summer and their catastrophic decline in the face of high ambient concentrations of chlorophyll suggest that factors other than temperature and food supply are important in influencing the dynamics and distribution of zooplankton in this lake.     

Reduction of the filtration rate of Daphnia galeata by dissolved photosynthetic products of edible phytoplankton

The filtration rate of Daphnia galeata was determined in in situ experiments in Bautzen Reservoir and in laboratory experiments, where daphnids were exposed to filtrates that previously contained either natural phytoplankton or cultured eukaryotic algae (Scenedesmus obliquus or Asterionella formosa), respectively. Individual filtration rate (FR) was measured using fluorescent beads, taking into account ingested beads in the gut only. Compared to heated control treatments (100 °C), dissolved compounds released by the nutritious cultured algae during the preconditioning phase or by the natural phytoplankton assemblages from Bautzen Reservoir strongly reduced the filtration rate of D. galeata (down to 60%). Heating deactivated these dissolved compounds. A significant correlation was found between primary production measured in situ and the reduction of FR in the filtrate of reservoir water, indicating that extra-cellular products released during photosynthesis triggered the reduction of the filtration rate. The ratio of ingested to collected beads was used to quantify the proportion of food, which was not only collected but passed the mouth of D. galeata. The ratio of ingestion to collection was compared between filtered and unfiltered reservoir water both media identical with respect to the concentration of dissolved compounds, whereas other factors (e.g. food concentration, temperature, filtration rate) were different. The changes in this ratio between filtered and unfiltered reservoir water suggest that D. galeata is capable of a chemosensory control of the ingestion behaviour by detecting external metabolites.     

Trade-offs inDaphnia vertical migration strategies

Summary Planktonic animals performing diel vertical migration (DVM) experience a tradeoff between reduced mortality and reduced reproductive output due to lower food availability in their refuge. Models of DVM as an evolutionarily stable strategy predict that, under certain conditions, strategies of both migration and non-migration can coexist. Vertical profiles of animal abundances during day and night, however, do not allow any discrimination between the behaviour of individuals or subpopulations. We used length-body protein regressions as a measure of the nutritional state ofDaphnia to distinguish possible sub-populations differing in their migration strategy. An overwhelming part of the population migrated downwards during the day. However, the few daphnids in the epilimnion during the day had significantly higher protein content than the animals in the deep water, indicating that these daphnids did not migrate randomly but remained in the surface food-rich water all day. This shows that migrating animals gain no metabolic advantage over non-migrating ones.Supported by a F.P.U. grant (Spanish Goverment)     

Bacterial epibionts of Daphnia: a potential route for the transfer of dissolved organic carbon in freshwater food webs

The identification of interacting species and elucidation of their mode of interaction may be crucial to understand ecosystem-level processes. We analysed the activity and identity of bacterial epibionts in cultures of Daphnia galeata and of natural daphnid populations. Epibiotic bacteria incorporated considerable amounts of dissolved organic carbon (DOC), as estimated via uptake of tritiated leucine: three times more tracer was consumed by microbes on a single Daphnia than in 1 ml of lake water. However, there was virtually no incorporation if daphnids were anaesthetised, suggesting that their filtration activity was essential for this process. Microbial DOC uptake could predominantly be assigned to microbes that were located on the filter combs of daphnids, where the passage of water would ensure a continuously high DOC supply. Most of these bacteria were Betaproteobacteria from the genus Limnohabitans. Specifically, we identified a monophyletic cluster harbouring Limnohabitans planktonicus that encompassed sequence types from D. galeata cultures, from the gut of Daphnia magna and from daphnids of Lake Zurich. Our results suggest that the epibiotic growth of bacteria related to Limnohabitans on Daphnia spp. may be a widespread and rather common phenomenon. Moreover, most of the observed DOC flux to Daphnia in fact does not seem to be associated with the crustacean biomass itself but with its epibiotic microflora. The unexplored physical association of daphnids with heterotrophic bacteria may have considerable implications for our understanding of carbon transfer in freshwater food webs, that is, a trophic ‘shortcut'' between microbial DOC uptake and predation by fish.     

Density and depth variations of Daphnia multilocus genotypes during a summer period in Lake Maarsseveen

The genotype composition of a Daphnia population complex during a summer period in Lake Maarsseveen (The Netherlands) was determined by allozyme analysis. The depth distribution, diel vertical migration and several parameters of the total population were measured. Young-of-the-year (0⁺) perch (Perca fluviatilis) were caught and species and allozyme types of Daphnia in the perch gut were also analysed. During May 1997, the densities of D. hyalina, D. galeata, the back-cross D. g x h – hyalina and the multilocus allozyme genotypes of the hybrid D. g x h decreased, except one multilocus genotype (MMMF). Total population size decreased and the ratio of females with eggs to those without eggs decreased as well. Food limitation during this clear-water phase in the lake is considered responsible. All genotypes, except MMMF, gradually descended in the water column. This drift is thought to be a reaction to the abundantly present 0⁺ perch or to the kairomones of this fish, although predation on the daphnids was still absent. In June, diel vertical migration started, except again part of the MMMF subpopulation. The other part migrated over a short distance compared with the other taxa and allozyme types. Within two weeks, the upper 5 m of the epilimnion was devoid of Daphnia, and guts of perch were predominantly filled with MMMF. The daphnids in the gut and the lake did not differ in allozyme type composition. By the end of July, population density had increased again. The size and composition of the Daphnia population complex continuously changed during the study period, as did the depth distribution of the components. Different genotypes within the population complex seem to have developed different strategies to cope with starvation and predation and the state at a particular moment can be understood only if past and present factors are considered.     

Ingestion Rates of Some Daphnia Species in a Shallow Lake (Lake Balaton,Hungary)

Filtering rates of three species of Daphnia were studied in situ by the ¹⁴C technique during the summer in a lake with a water temperature of 20-25°C (Lake Balaton, Hungary). The renewal time of gut contents proved to be 3-5 min in each of the three species whether fed on natural or artificial food. From May to September, filtering rates of D. cucullata, D. hyalina and D. galeata averaged 7.8, 10.3 and 9.2 ml ind.⁻¹ day⁻¹, respectively. Identically long individuals of all three species had almost the same filtering rates in spring, whereas in summer animals of identical weights had similar filtering rates regardless of species because of the length variability of the helmets. The grazing rate of adult filter-feeding crustaceans is suggested to be approximately 7 % per day.     

Is the midsummer decline of Daphnia really induced by age-0 fish predation? Comparison of fish consumption and MDaphnia mortality and life history parameters in a biomanipulated reservoir

We analysed the temporal pattern of a Daphnia galeata populationand the development of the age-0 Ash community in a long-termbiomanipulated lake (Bautzen reservoir, Germany) during springand early summer of two successive years. In Bautzen reservoir,the age-0 Rsh-Daphnia interaction is a key process within thefood web due to the low abundance of adult zooplanktivorousfish. Daily consumption of daphnids by age-0 fish, as estimatedby a bioenergetics model, was compared to daily mortality ratesof daphnids. In addition, we estimated life history parametersof Daphnia that may indicate predation impact by fish. A midsummerdecline of daphnids occurred only in July of 1996, whereas in1995 the Daphnia biomass remained >4 mg wet weight 1^–1for the entire summer. The percentage of total Daphnia mortalitydue to fish predation before the onset of the midsummer declinewas –2% day^–1. Temporal patterns of individual size,clutch size and size at maturity also indicated that the 1996midsummer decline of daphnids was not the exclusive consequenceof age-0 fish predation. Instead, low reproductive capacityof daphnids also contributed significantly to the decline. Consequently,year-to-year variation of the Daphnia dynamics may be determinedby a fine-tuned ‘timing’ between the period of reducedfecundity and the time of the strongest predation impact byage-0 fish.     

Temperature impact on the midsummer decline of Daphnia galeata: an analysis of long-term data from the biomanipulated Bautzen Reservoir (Germany)

1. The influence of water temperature on occurrence and duration of a midsummer decline (MSD) of Daphnia galeata was studied in the biomanipulated Bautzen Reservoir in Germany. The proportion of piscivores in the fish community of the reservoir has been enhanced experimentally since 1981. As a consequence, Daphnia galeata has dominated the zooplankton. Over 18 years of study (1981–1998), a long‐lasting MSD (longer than 30 days) occurred in 7 years, whereas a short MSD (shorter than 30 days) was observed in 6 years. During the remaining 5 years, an MSD was not observed.
2. Two hypotheses were examined to explain the observed patterns. First, we postulated that high water temperature during winter and early spring (January–April) leads to an MSD after an early and high spring peak of daphnids. On the other hand, low temperature during winter and early spring should not cause an MSD owing to a slower increase of the population, resulting in a later peak of daphnids. Second, we hypothesized that the mean water temperature during early summer (May and June) influences the occurrence of an MSD (by controlling young‐of‐the‐year (YOY) fish predation on daphnids).
3. The water temperature during winter and early spring explains 83%, and the early summer water temperature 55%, of interannual variation in the occurrence of an MSD.
4. The interannual variation in duration of an MSD was neither explained by temperature during winter and early spring nor by early summer temperature alone, but in 14 of the 18 years (78%) by a combination of both.
5. We conclude that water temperature during winter and early spring had a strong impact on Daphnia mortality by influencing height and timing of the spring peak which, in turn, influenced the extent of overexploitation of their food resources. By contrast, the water temperature during early summer probably influenced the mortality of daphnids caused by predation of YOY fish. The relative timing of both sources of mortality, which depends on the temperature regime during the first 6 months of the year, is the key process in controlling the occurrence and duration of an MSD. A long‐lasting MSD, therefore, is likely in Bautzen Reservoir only if temperatures are high during winter and early spring, as well as during early summer.
6. As a consequence of climate warming, recent climate records reveal warming during winter, spring and early summer in middle Europe, rather than an increase in mean annual temperatures. If our findings and conclusions are related to this regional and temporal pattern of climate warming, an increasing frequency of years with a long‐lasting MSD and, consequently, a decreasing efficiency of biomanipulation can be predicted.