Indirect effect of planktivorous fish on the growth and reproduction of Daphnia galeata

Both the size of Daphnia galeata primiparae and the average egg volume were smaller in individuals reared from neonata in water inhabited for the preceding 24 hours by Y-O-Y roach when compared with control animals reared in the same water with the exclusion of fish. The smaller size of experimental primiparae resulted from smaller increments in some instars and from the earlier maturation of about two-thirds of those that matured in the 4th instar, while the rest of the experimental animals and all the controls matured in the 5th instar. There were larger clutches and smaller eggs in experimentals when compared with the similar sized controls, but the difference hasn't proved to be statistically significant.     

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.     

Life history response of <Emphasis Type="Italic">Daphnia galeata</Emphasis> to heterogeneous conditions within a reservoir as determined in a cross-designed laboratory experiment

Growth and reproduction were studied in the laboratory in a cross-designed experimental set-up in four Daphnia galeata subpopulations collected from different locations (with respect to water characteristics) in a reservoir (epilimnion, metalimnion and hypolimnion in the deepest part of the reservoir near the dam and epilimnion of the upstream part of the reservoir) and in a laboratory clone of the same species. The results of two-way ANOVA revealed significant effects of the two parameters manipulated – source of water used for cultures and Daphnia subpopulation – on the life history characteristics of growth and reproduction. The water from the upstream part of the reservoir was the most favourable culture medium for all characteristics of the Daphnia groups studied (the largest primiparae, clutches and eggs, the shortest postembryonic development time and filtering setae). The poorest performances were recorded in the downstream, epilimnetic and metalimnetic waters. The primiparae in the hypolimnetic water were smaller but had relatively larger clutches of smaller eggs and slightly longer postembryonic development times. The Daphnia subpopulation originating from the hypolimnion had the smallest primiparae, the largest clutches, the smallest eggs and the shortest postembryonic development, whereas the opposite was found in animals from the epilmnion. These differences in ecologically relevant traits were supported by analysis of the quasi-neutral genetic markers that indicated significant site-dependent differences in clonal structure between the subpopulations. There was no consistent trend to higher within-group variance in the life history traits in the genetically heterogeneous subpopulations from the reservoir compared to the laboratory clone.     

Estimation of phosphorus metabolic rates of Daphnia galeata using a three-compartment model

A compartment model to estimate the different phosphorus metabolicrates in Daphnia galeata is presented. The model has three compartments:gut, metabolic pool and structural pool. Existing two-compartmentmodels used for carbon and phosphorus turnover in Daphnia donot allow estimation of ingestion and egestion rates. We extendedexisting two-compartment models with one more compartment, thegut, which allowed us to estimate both the ingestion and theegestion rates. Parameters of the model are estimated from asingle experiment of feeding unlabelled Daphnia with ³²P-labelledScenedesmus obliquus. Separate experiments with juvenile andadult daphnids were carried out in order to compare their metabolisms.This model permits a reliable estimation of the different metabolicrates of Daphrna in a single experiment and discriminates clearlybetween animals of different sizes.     

Cyclomorphosis as a factor explaining success of a Daphnia hybrid in Tjeukemeer

Recent studies have shown that Daphnia hybrids are intermediate to their parental species for several life history traits, and that they combine advantageous traits of the parental species, thereby decreasing their risk of being preyed upon by planktivorous fish. In this study D. galeata, D. cucullata and their interspecific hybrid are compared with respect to cyclomorphosis and juvenile growth as a potential mechanism to avoid invertebrate predation. Helmets and tail spines are known to reduce predation risk, whereas fast juvenile growth reduces the period that Daphnia are vulnerable to predation. In laboratory cultures, I tested the hypothesis that, with regard to cyclomorphosis, juvenile hybrids are more comparable to the juveniles of D. cucullata, and with respect to growth rate, they are more comparable to D. galeata. The hybrid did not differ significantly from D. galeata in the time needed to reach 0.9 mm body length. In contrast, the relative spine and helmet length of juvenile hybrids were more comparable with those of D. cucullata, whereas in adult instars these characters were more similar to D. galeata. Thus during the first juvenile instars, the hybrids combine the relative long helmet and spine of D. cucullata, with the faster growth of D. galeata. This combination of characteristics might reduce hybrid vulnerability to invertebrate predation in field situations.     

What can we learn from Daphnia filtering screens?

The relative filtering setae length (RFSL the length of filteringsetae on the third pair of thoracic limbs expressed as a percent of carapace length) was determined in three developmentalcategories of a reservoir population of Daphnia galeata andits relationship to phytoplankton fluctuation. RFSL of adultfemales immediately after moulting reflects the current foodconditions in the reservoir (higher RFSL at lower food level,and vice versa), whereas RFSL of adult females close to moultingreflects the food conditions several days or weeks ago. RFSLof juvenile animals reacts the most quickly to changing foodconditions, provided the temperature allows frequent moulting.Higher RFSL can also indicate poor-quality food for daphnids,in situations when food quantitative parameters (such as chlorophylla) show high values.     

Facing multiple enemies: parasitised hosts respond to predator kairomones

During their lifetime most organisms are exposed to various enemies influencing their victims in multiple direct and indirect ways. Most studies concentrate on the effects of one enemy at a time, thereby not taking into account that in nature organisms are often simultaneously exposed to more than one enemy. We conducted a life-history experiment to investigate the simultaneous effects of predators (fish, Leuciscus idus) and parasites (microparasite, Caullerya mesnili) on their victim (Daphnia galeata). D. galeata were exposed to predator kairomones, parasites or both. D. galeata are able to sense the presence of fish predators via chemical cues (= kairomones). Both fish predator kairomones and microparasite infections influence the life history of Daphnia. Some of the effects of fish predator kairomones are directly opposed to microparasite effects; fecundity, for example, is increased in the presence of fish kairomones and decreased in Daphnia parasitised with C. mesnili. We investigated the influence of both threats on age at maturity, body size at different adult instars, fecundity and survival of one D. galeata clone. In the presence of fish kairomones, all D. galeata matured significantly earlier and increased the number of eggs in the second brood significantly. Parasitised D. galeata matured significantly earlier than non-parasitised ones in the absence and presence of fish kairomones. An infection with the microparasite C. mesnili led to significantly lower clutch sizes at the second adult instar, to significantly smaller body sizes from adult instar three onwards and to significantly reduced survival. No significant interaction effect between the responses to fish presence and to parasite infection was found for any of the investigated life-history traits. The lack of interaction effects between the exposure to predator kairomones and parasite infection was most likely due to the different timing of the effects. Fish kairomones affected D. galeata early in its life history whereas C. mesnili increased in its effects over time. Our results show that parasitised D. galeata are able to exhibit life-history responses to fish predator presence early in their lives. Thus, D. galeata parasitised with C. mesnili have a similar chance as non-parasitised D. galeata to escape from fish predation via life-history changes. Since older parasitised D. galeata are smaller, they may have an even better chance to escape visual predators under actual predation.     

Life history patterns of parental and hybrid Daphnia differ between lakes

1. We investigated whether Daphnia galeata × hyalina hybrids of Lake Constance and Lake Greifensee show the same pattern of life history parameters as previously reported for D. galeata × cucullata hybrids and whether such a pattern is consistent between Daphnia populations from those two lakes. 2. Hybrids in Lake Constance were intermediate in size compared with the parental species. Hybrids in Lake Greifensee were smaller than D. galeata. The intrinsic growth rate (r) of hybrids from Lake Constance was not significantly different from the faster growing parental taxon D. galeata. However, r of hybrids from Lake Greifensee was significantly lower than that of D. galeata. 3. The observed juvenile body length differences between the taxa varied with the clutch number. The first clutch juvenile lengths of the three taxa did not differ for Lake Constance. First clutch juveniles of Lake Greifensee D. galeata were smaller than hybrid first clutch juveniles. The third clutch juvenile length did not differ between taxa from Lake Greifensee, but D. galeata juveniles from Lake Constance were bigger than those of D. hyalina. 4. The life history pattern found in Lake Constance corresponds to previous findings from other studies. The hybrids in this lake combine the faster population growth of one parental species with a relatively small size. In the case of Lake Greifensee hybrids, the relatively large size of first clutch juveniles and the small size of the adults could be interpreted as dual adaptations to invertebrate and fish predation. We speculate that the lower population growth rate of the hybrids is a trade‐off for this twofold protection.     

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     

Predator-induced phenotypic plasticity in Daphnia pulex: uncoupling morphological defenses and life history shifts

Chemical cues from a predator Chaoborus sp. induce morphological defense (neck spine) and life history shifts (later reproduction, decreased fecundity but larger juvenile size) in the waterflea Daphnia pulex. These shifts have been interpreted either as costs of defense or as separate adaptation. In order to investigate if the life history shifts can be separated from the morphological defense, Daphnia pulex individuals were exposed to chemical cues from Chaoborus at different stages of life for variable periods. The daphnids that were exposed to Chaoborus started their reproduction later than the controls, although the differences were not statistically significant. Neck spine was induced only if daphnids were exposed to Chaoborus in an early stage of their life. Numbers of eggs produced were not affected by the different treatments, but egg mortality was higher in mothers exposed to Chaoborus. With these treatments it was possible to see neck spine induction without measurable life history changes or costs. On the other hand, irrespective of neck spine presence, the Chaoborus chemical(s) had an effect on Daphnia pulex mothers.Publication no 2159. Netherlands Institute of Ecology, Centre for LimnologyPublication no 2159. Netherlands Institute of Ecology, Centre for Limnology     

Life history shifts in response to different levels of fish kairomones in Daphnia

Life history parameters, such as age and size at maturity, neonatesize and number of eggs in the first clutch, of a clone of thehybrid Daphnia galeata x hyalina were determined at six levelsof fish kairomones. It was found that the age and size at maturitydecreased gradually with increasing kairomone level, while thenumber of eggs in the first clutch increased linearly. At thesame time, neonate size decreased linearly with increasing kairomonelevel. In a starvation experiment, newly born neonates of mothersnot cultivated in the presence of kairomones survived the longest.These results show that Daphnia is able to adjust its life historytraits to different levels of kairomones, and thus to differentfish densities.     

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)     

Food quality effects on instar‐specific life histories of a holometabolous insect

1. It is a long‐standing challenge to understand how changes in food resources impact consumer life history traits and, in turn, impact how organisms interact with their environment. To characterize food quality effects on life history, most studies follow organisms throughout their life cycle and quantify major life events, such as age at maturity or fecundity. From these studies, we know that food quality generally impacts body size, juvenile development, and life span. Importantly, throughout juvenile development, many organisms develop through several stages of growth that can have different interactions with their environment. For example, some parasitoids typically attack larger instars, whereas larval insect predators typically attack smaller instars. Interestingly, most studies lump all juvenile stages together, which ignores these ecological changes over juvenile development.
2. We combine a cross‐sectional experimental approach with a stage‐structured population model to estimate instar‐specific vital rates in the bean weevil, Callosobruchus maculatus across a food quality gradient. We characterize food quality effects on the bean weevil's life history traits throughout its juvenile ontogeny to test how food quality impacts instar‐specific vital rates.
3. Vital rates differed across food quality treatments within each instar; however, their effect differed with instar. Weevils consuming low‐quality food spent 38%, 37%, and 18% more time, and were 34%, 53%, and 63% smaller than weevils consuming high‐quality food in the second, third, and fourth instars, respectively. Overall, our results show that consuming poor food quality means slower growth, but that food quality effects on vital rates, growth and development are not equal across instars. Differences in life history traits over juvenile ontogeny in response to food quality may impact how organisms interact with their environment, including how susceptible they are to predation, parasitism, and their competitive ability.

     

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.     

Carbon as an indicator of Daphnia condition in an alpine lake

Carbon–length relationships can be used to indicate the condition of daphnids in natural situations. We examined the Daphnia galeata population of an alpine lake (Oberer, Arosasee, Switzerland), where most individuals display diel vertical migration behaviour (DVM). Normally, migrating daphnids face a trade-off between `predator safe areas' in the hypolimnion and `food rich areas' with high predation risk. However, in alpine lakes, with chlorophyll a and POC maxima typically in deeper layers and weak temperature gradients, migrating species are confronted less with this trade-off. We investigated the seasonal changes in carbon content of D. galeata in different depth strata in relation to environmental parameters of the lake. The carbon content of D. galeata was high in spring, but declined considerably in summer and increased slightly in autumn. The low values indicate that D. galeata are food limited for much of the year. The slopes of the regression lines between carbon content and body length varied seasonally, but were not significantly different among depths on a given date. In summer, D. galeata individuals residing in the deep layers during the day had a significantly higher carbon content than individuals in the surface layers. During the rest of the year, the carbon content of individuals was similar among all depth strata. We conclude that migrating D. galeata and individuals remaining in deep layer had better body conditions compared to non-migrating D. galeata in this alpine lake during summer, when migration amplitudes were highest.     

Influence of temperature on growth, reproduction and longevity of Moina salina Daday, 1888 (Cladocera, Moinidae)

This paper presents the results of a study of the effects oftemperature on the growth, reproduction and longevity of thecladoceran Moina salina, a species of potential use as livefood in marine aquaculture. The growth rate of M.salina increasedwith increasing temperature. Some parameters of development,such as length at death and the number of adult instars, werealso positively related to temperature. Other parameters (durationof juvenile and adult instars) decreased with increasing temperature,while the number of juvenile instars was unaffected. An increasein temperature resulted in a reduction in age at maturity anda decrease in the number of days between broods. The numberof young per female, the number of broods per female, the numberof youngper day of reproductive life, and the number of youngper brood, increased up to a temperature of 25°C. At 15and 20°C. substantial degeneration of eggs and/or embryosoccurred. Likewise, temperature affected the type of reproductioncarried out by sexual females. Temperature and longevity wereinversely correlated. It was concluded that temperature actsas a very important factor regulating the life cycle of M.salina.Temperature >30°C may correspond to sublethal levels,while a temperature of 15°C is considered to impose stress.The range 20–25°C is optimal for the development andreproduction of this species.     

Adult, not juvenile mortality as a major reason for the midsummer decline of a Daphnia population

We analysed dynamics and structure of a Daphnia galeata populationprior to and during a midsummer decline of this species in Bautzenreservoir (Saxony, Germany). Patterns of juvenile and adultmortality were determined by combining field data with laboratoryestimates of juvenile growth. After establishing high densities,fecundity and recruitment of D.galeata markedly decreased, whereassize at maturity was high. Immediately before the populationdecline, adult mortality increased and remained high even afterthe decline, whereas juvenile mortality was low during the wholeinvestigation period, and virtually absent after the decline.We conclude that the succession of events leading to a midsummerdecline of Daphnia is as follows. (i) A quick increase in Daphniaabundance leads to the formation of a strong ‘peak cohort’of about the same age. (ii) During the clear-water phase, foodconditions deteriorate, fecundity declines and hence, recruitmentis low. Juvenile mortality during this period is low, but present.(iii) Adult mortality increases when the ‘peak-cohort’reaches its mean life-span, which is reduced due to interactionsbetween age-specific and starvation-induced mortality. At thispoint, Daphnia population dynamics can no longer be explainedwithout the onset of size-selective predation. Hence, the timingbetween enhanced mortality due to senescence on the one handand predation on the other hand, both directed towards adultdaphnids, may be decisive for the initiation of a midsummerdecline of Daphnia.     

Daphnia galeata life history response to heterogeneous food conditions and dissolved chemicals in the Rímov Reservoir

Laboratory cultivation experiments were carried out in order to determine the response of D. galeata to food conditions and dissolved chemicals in the water collected in three contrasting sites of the Rímov Reservoir. There were two types of cultivation media: (1) lake water with natural seston, filtered through a 40 m mesh size sieve, (2) lake water filtered through a membrane filter 0.45 m (or 0.22 m), with Scenedesmus subspicatus culture added to a concentration of 1.0 mg POC l^–1. In the treatment with natural seston, the parameters investigated (body length, filtering setae length, clutch size, egg volume and postembryonic development time) were determined mainly by the food level, which was higher in the upper part of the reservoir than near the dam. In one series, significantly larger clutches and smaller eggs together with longer postembryonic development (PED) were detected in daphnids cultivated in hypolimnetic water, compared to those in epilimnetic water. As particulate carbon concentrations were similar, this indicates that in that case food quality played a role shaping life histories in Daphnia. In some of the experiments with the filtered water, there was a detectable effect of dissolved chemical(s), which induced shorter PED and smaller primiparae in the animals cultivated in water from the upper part of the reservoir.     

The effect of accelerations in light increase on the phototactic downward swimming of Daphnia and the relevance to diel vertical migration

The effect of relative increases in light intensity on photobehaviourwas studied in the hybrid Daphnia galeata x hyalina. We firstcarried out a series of experiments to study the influence offish kairomone on several response variables of light-inducedswimming. With fish kairomone present, an increase in the percentageof reacting daphnids to 100% was found at almost all ecologicallyoccurring relative light change rates that were above threshold.The relationship between the relative increase in light intensity(stimulus) and the time expiring between the onset of the stimulusand the start of the downward swimming response was not influencedby fish kairomone, nor did kairomone alter the functional relationshipbetween stimulus strength and downward displacement velocity,although velocity increased. During the previous experiments,various light change rates were applied, but per test run theserates were constant. The natural relative light increase inthe early morning consists of continuously increasing relativelight change rates, turning into decreasing rates after themaximum is reached     

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.