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.     

Sexual reproduction in Daphnia: interspecific differences in a hybrid species complex

To investigate whether reproductive variation in the Daphnia galeata-cucullata-hyalina hybrid species complex can explain the occurrence of hybrids and backcrosses, a total of 43 clones from three north German lakes were tested for allocation to sexual reproduction under equal stress conditions. Six replicates per clone were followed until the seventh adult instar. I used the following cues to promote sexual reproduction: short photoperiod, water from a crowded Daphnia culture, fish-conditioned water and low food concentration. For each animal, clutch size and clutch sex were recorded. Ephippia, which were empty since the animals were cultured individually, were only produced by D. cucullata and D. cucullata x hyalina (26% and 6% of the broods, respectively), whereas almost all taxa produced males (the range was 2–15%). Intraspecific variation for male and ephippial production was also found. The fact that the taxa tested show different responses to the stimuli of-fered may indicate that there is reproductive isolation between them. This is supported by field data from the Tjeukemeer (The Netherlands) which shows that D. galeata mostly produces ephippia in spring and D. cucullata in autumn. The presence of hybrids with ephippia in both seasons, however, shows that backcrossing is also possible.     

Amplified Fragment Length Polymorphism (AFLP) Reveals Species-Specific Markers in the Daphnia Galeata–HyalinaSpecies Complex

Daphnia often occur in species complexes that consist of two or more co-occurring species and their hybrids. Hybrid individuals are often capable of sexual reproduction and so backcrossing with introgression occurs. To better understand hybridization and backcrossing frequency, we sought to develop PCR-based, species-specific markers in the Daphnia galeata–hyalina species complex using amplified fragment length polymorphism (AFLP). This technique produces large numbers of reproducible markers for assessing diversity across the nuclear genome and provides several advantages over mtDNA and microsatellite approaches. We examined 28 clones of D. galeata, D. hyalina, and their hybrids isolated from Lake Constance on the Swiss-German border. Using a single AFLP primer combination we found five potential species-specific markers, defined as bands that occurred in >80% of one parental species and <20% of the other. Two bands appeared to be co-dominant and were present (homozygous) in D. galeata, absent in D. hyalina, and heterozygous in the hybrid. We conclude AFLP could provide enough PCR-based, species-specific markers to identify species, hybrids, and backcrosses from even small amounts of tissue (i.e. resting eggs).     

Seasonal variability in the relationship between body length and individual dry weight as related to food abundance and clutch size in two coexisting Daphnia species

Dry weights of individual Daphnia galeata and D. hyalina, whichcoexist in Lake Constance, were determined weekly during a twoyear investigation. From each sample and each species a length–weightrelationship (LWR) was calculated resulting in 54 (65) regressionequations based on N=2466 (3032) in D. galeata (D. hyalina).The constants (Ln(A)) and (B) of the LWRs showed a marked seasonalvariability, ranging from Ln(A)=1.0784–2.4740(1.0152–2.3860)and B=2.14–4.20 (2.11–4.15). The seasonal variabilityof the LWR was estimated with four different models. Condensingthe 54 (65) equations into one seasonal cycle resulted in amodel with an explained variance of r²=0.873 (0.869). Threemore models were established by calculating multiple regressionswith take stratification, average fecundity (E), and food concentrationas determining variates. Differences in LWRs of the two Daphniaspecies were significant (p <0.001). As the most useful approximationduring the period of stratification, LWRs are recommended forD. galeata: Ln(W)=(1.5674+0.0287^*(E))+Ln(L)^*(3.3611+0.0111^*(E)) as a generalized LWR for an epilimnetic daphnid, and for D.hyalina: Ln(W)=(1.5593+0.0613^*(E))+Ln(L)^*(3.2709+0.0017^*(E)) as LWR for a daphnid which migrates diurnally between epi andhypolimnion. The explained variance of this model is r²=0.843(0.826). The influence of food concentration in the epilimnionon LWRs was found to be significant on Ln(A) but not on (B). ^*This research project was supported by the Deutsche Forschungsgemeinschaft     

Taxon-specific reaction norms to predator cues in a hybrid Daphnia complex

1. Previous studies have shown that interspecific hybridisation is common among taxa from the Daphnia galeata/hyalina/cucullata species complex. We investigated the influence of predator kairomones on the morphology and life histories of nine clones belonging to three taxa (pure D. galeata, F₁ hybrids between D. galeata and D. hyalina, and backcrossed D. hyalina) of this species complex. Predators exerting positive (fish) and negative (Chaoborus larvae) size‐selective predation were tested. 2. The most responsive traits were size at maturity and size of neonates. Despite large between‐clone variation, discriminant analysis revealed that the three taxa were distinct from each other in key life‐history traits. F₁ hybrids did not react in an intermediate way compared to the other taxa: the multivariate distances between F₁ hybrids and either taxon were larger than between pure D. galeata and backcrossed D. hyalina. 3. The average plasticity (calculated across all traits) was similar for all three taxa. With regard to the size at maturity and neonate body size, the strength of the response was a function of the intrinsic values of these traits expressed in the control. For example, for size at maturity, smaller individuals showed a significantly stronger reaction to Chaoborus kairomones than larger ones. 4. Finally, we monitored seasonal changes in body size, egg number and population density of pure D. galeata and F₁ hybrids in Greifensee (Switzerland). The two taxa experienced similar seasonal changes in body size but, on some sampling dates, they differed in mean egg number. The observed seasonal changes in Daphnia body size were consistent with what would be expected if the predator assemblage shifted from fish to Chaoborus over the course of the summer. The fluctuations in the frequencies of Daphnia taxa, however, were not related to seasonal variation in Daphnia body size. 5. Experimental data suggest that temporally heterogeneous predation regimes might be an important condition stabilising the co‐occurrence of Daphnia hybrids with parental taxa. Predation regimes, however, cannot solely explain dynamic changes in taxon frequency in Greifensee.     

Daphnids respond to algae-associated odours

Using a Y-tube olfactometer, it was found that Daphnia galeatax hyalina moves to the arm with odour from either of two ediblealgal species (Scenedesmus acuminatus and Oscillatoria limnetica)rather than to the alternative arm with clean water. However,no differential response was observed when odours of the toxiccyanobacterium (Oscillatoria agardhu) were tested. We suggestthat odours associated with edible algae attract Daphnia whereasnon-edible algae do not elicit attraction of Daphnia.     

Daphnia populations in three interconnected lakes with roach as the principal planktivore

Data on two co-existing Daphnia species, D.cucullata (G.O.Sars)and D.hyalina (Leydig), from three neighboring and interconnectedlakes in northern Poland, revealed an extremely confined rangeof population density fluctuations throughout summer in eachof the two species in each of the three lakes, with D.cucullatabeing an order of magnitude more abundant than D.hyalina (meanof 11.5 ind. l^–1 for D.cucullata and 1.1 ind. l^–1for D.hyalina with 95% confidence limits of ±2.5 and±0.5 ind. l^–1, respectively), in spite of markedlychanging fecundity. There was no apparent phase of decline orincrease that could be related to the distinct phases of lowor high fecundity resulting from different food levels. Analysisof the gut contents of roach (Rutilus rutilus), the dominantplanktivore in the three lakes, revealed identical selectivityfor each of the two Daphnia species, suggesting that the lowerdensity of the D.hyalina population was compensated for by thegreater conspicuousness of individuals of this species. It isconcluded that the population density of these Daphnia speciesremains far below the carrying capacity of the habitat and doesnot depend on food levels. Food availability merely sets therate of population increase, while the actual population densityin the lakes studied reflects the species' vulnerability topredation by planktivorous fish.     

Experimental interspecific hybridization in Daphnia

Hybridization is a common phenomenon in Daphnia (Cladocera; Anomopoda); interspecific hybrids have been found between several species and hybrids are found in many European lakes. Although much information on the morphology, ecology and genetics of hybrids is available, little is known about the level of reproductive isolation among species or about the relative fitness of hybrids and parental species. In order to facilitate studies on differentiation and speciation processes and comparative experimental studies on hybrids and recombinant genotypes, we present the first successful laboratory crossing experiments of two different Daphnia species, D. galeata and D. cucullata. Males and sexual females from two D. galeata and two D. cucullata clones were reciprocally crossed, juveniles hatched from resting eggs and reared until maturity. Hatching and juvenile survival rates of hybrids were relatively low (12.1% and 24%, respectively). D. galeata and D. cucullata clones vary in their level of successful interspecific matings and in the number of subsequent offspring. In general, hybrid crosses between D. cucullata females and D. galeata males were more successful than reciprocal crosses.     

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.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

The energy budget of two sympatric Daphnia species in Lake Constance: productivity and energy residence times

Summary Two Daphnia species, D. hyalina and D. galeata, are living in Lake Constance. Both populations show logistic growth, with the phase of exponential increase in May and the phase of steady state during the summer. In spring, food is not limited and both daphnids reveal an exploitative strategy: females mature early, primiparae are small, egg weights are small, but clutch sizes are large. From June on, when food limiting conditions prevail, D. hyalina starts to migrate vertically with a diurnal rhythm whereas D. galeata keeps on living in the epilimnion over the whole season. The differing thermal environments of the populations lead to short generation times (age of first reproduction) of 11 days for D. galeata in mid-summer, and to 47 days (July) for D. hyalina. The annual production (P) of D. hyalina and D. galeata were 18 and 24 g DW/m², respectively. The average standing stocks (B) were 1.6 and 0.9 g DW/m². This corresponds to turnover rates of 11 x and 27 x per season (200 days), and to turnover times of 18 days and 7.5 days, respectively. From 1980 to 1982 the cumulative annual primary production (PPR) varied between 260 and 330 g C/m². The common productivity (P) of the functional component Daphnia (=sum of both Daphnia species) followed the PPR, but the Daphnia stading stock was constant at 2.5 g DW/m². The increase of secondary production (P) was a consequence of a shift in abundance between the two Daphnia species: at low PPR, D. hyalina was 4 x more abundant, but at high PPR, the two species were equally abundant. Such internal regulatory mechanism within a system component is in accordance with the hypothesis of Cheslak and Lamarra (1981): increase of energy residence time (B/R) with declining nutrients and increasing strength of competitive interrelationships within a given functional component of an ecosystem.This research was supported by the Deutsche Forschungsgemeinschaft within Sonderforschungsbereich Stoffhaushalt des Bodensees (SFB 248)     

Does trimethylamine induce life-history reactions in Daphnia?

Laboratory experiments were carried out to investigate life-history reactions of Daphnia to trimethylamine (TMA), a substance, which has recently been found to induce a similar phototactic reaction in Daphnia as fish kairomones. The effects of different treatments (control, fish kairomone and TMA) on the life-history traits of five clones of D. magna and one clone of D. galeata and D. hyalina, respectively, were studied. Only D. magna exhibited significant reactions to TMA and to fish kairomone. D. galeata showed no significant responses to either TMA or to fish kairomone. D. hyalina reacted to the fish treatment with two traits (size and number of eggs at maturity). The comparison of the reaction norms to TMA and fish kairomone demonstrated that the directions of some responses to both factors were similar but were different for others such as for size at maturity. TMA resulted in a larger size at maturity in D. magna, whereas fish kairomone had no significant influence on this trait. We, therefore, conclude that trimethylamine is not the primary causative chemical agent in fish induced life-history adaptations in Daphnia.     

The contribution of differential hatching success to the fitness of species and interspecific hybrids

Resting egg banks of microcrustaceans have been used to reconstruct the evolutionary and ecological history of species. However, recent studies provided evidence for a discrepancy between dormant propagules in the sediment and the planktonic population. This pattern raises two questions: First, what is the value of data on resting egg banks for population dynamics over time and second, which component of the reproductive cycle causes the observed inconsistency? In our study we focussed on the second question by comparing the taxon composition of a resting egg bank with the reproductive success of ex-ephippial hatchlings. Species and interspecific hybrid identification of dormant and hatched stages was achieved through the application of restriction fragment length polymorphism analysis of an internal transcribed spacer region. We found no significant deviation between the proportion of hatched Daphnia galeata, D. galeata × hyalina and D. hyalina individuals and the observed taxon composition of the resting egg bank. However, species and hybrids differed in their mode and relative success of reproduction. We conclude that the components of reproductive success in Daphnia contribute differentially to the fitness of species and interspecific hybrids. The discrepancy between resting egg banks and “active” planktonic populations results not from differential hatching of species but from the reproductive success of ex-ephippial females and the timing and frequency of sexual reproduction of the different taxa.     

Inter- and intraspecific differences in Daphnia life histories in response to two food sources: the green alga Scenedesmus and the filamentous cyanobacterium Oscillatoria

The effects of two food sources on life history traits of Daphniagaleata, Daphnia cucullata and their interspecific hybrid. D.cucullataxgaleata,were studied. For each taxon, two clones were reared on botha green alga (Scenedesmus obliquus) and a filamentous cyanobacterium(Oscillatoria limnetica). Reproduction on Oscillatoria was generallylower than on Scenedesmus, but a positive population growthrate was still achieved, except in one hybrid clone Life historytraits showed differences among the intraspecific clones. Daphniacucullata, the smallest of the three taxa, was the best of thethree in utilizing Oscillatoria as food, but D.galeata and thehybrid did not differ clearly. The clones within the taxa alsodiffered in the extent they could utilize Oscillatoria as food.The results are discussed in relation to the co-existence ofthese taxa in nature.     

The possible origin ofDaphnia cucullata procurva Poppe 1887 in the lakes of the Pomeranian Lakeland by hybridization in the past

Besides the genusBosmina, the genusDaphnia offers considerable taxonomic difficulties due to the existence of a very great number of different morphotypes and their intergrades. Especially the morphotypes clustering aroundDaphnia galeata Sars, 1864,D. hyalina Leydig, 1860, andD. cucullata Sars, 1862 offer a bewildering picture. In the last years an attempt has been made to understand this phenomenon mainly as the result of hybridization and colonization of new biotopes by populations with hybridogenetic polymorphism and genotypic diversity (founder effect). Now it will be suggested on a morphological base thatD. cucullata procurva Poppe, 1887, a unique and aberrant formtype restricted to few lakes along the river Brda in Poland, is probably an old and stabilized hybrid originating from interspecific hybridization betweenD. galeata andD. cucullata.     

Differences in genetic structure and ecological diversity between parental forms and hybrids in a Daphnia species complex

Hybridization is a common phenomenon in Daphnia species complexes. Hybrids often dominate in Daphnia populations; therefore it is worthwhile to look for principal differences between parental and hybrid populations with respect to their genetic structure and clonal differentiation. We studied natural populations of members of the Daphnia galeata/hyalina/cucullata complex in three lakes. In one of these lakes, one parental species (D. galeata) and one hybrid (D. galeata × cucullata) were investigated more intensively. The frequency of sexual reproduction was higher in parental populations, whereas clonal diversity was higher in hybrid populations. Ecological differentiation among clonal groups was more pronounced in the D. galeata × cucullata hybrid compared to D. galeata, whereas selection intensity was weaker. These results are discussed with respect to stability of clonal groups, multiple hybridizations and selective constraints.     

Molecular Genetic Diagnostics of Some Daphnia Species (Crustacea,Cladocera) from the Volga River

Population genetic structure of the widespread Daphnia species from Kuibyshev and Saratov reservoirs was examined using RAPD–PCR technique with four mitochondrial DNA (16S rRNA) specific primers. One of the Daphnia clones examined from the Volga region appeared to be conspecific to the North American Daphnia galeata, while another clone was most likely a hybrid between D. galeata and D. cucullata.     

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.     

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.     

A major shift in Daphnia genetic structure after the first ice‐free winter in a German reservoir

1. Climate warming may cause disruption of trophic linkages in aquatic ecosystems and lead to changes in abundance and genetic structure of zooplankton populations. We monitored the community of the Daphnia galeata‐hyalina hybrid complex in the Saidenbach Reservoir (Saxony, Germany) using allozyme electrophoresis for three consecutive years (2005–07), including one (2007) following an unusually warm winter that prevented the formation of ice cover for the first time in the history of the reservoir. 2. Genetic composition during the 2007 season differed substantially from the two preceding years that experienced the usual 3‐month ice period. Three abundance peaks in June, July and October 2007 were dominated by hybrids of Daphnia galeata x hyalina, whereas in the 2005 and 2006 seasons two peaks in June and September were dominated by Daphnia hyalina genotypes. 3. The genetic composition of the pool of diapausing eggs produced in autumn and the rate of change of genotype abundance during the following spring indicate recruitment of the D. hyalina subpopulation from ex‐ephippial animals during the spring population increase. 4. The differing potential to contribute to the overwintering animal pool or to the inoculum from diapausing eggs was confirmed by results from laboratory life‐table experiments. Daphnia galeata clones survived longer and produced parthenogenetic offspring under winter conditions, whereas D. hyalina clones showed a shorter lifespan and produced resting eggs. 5. Our results indicate a profound role of recruitment strategy in the observed shift in genetic composition. Increasing winter temperatures predicted in the context of climate change may thus favour overwintering animals, leading to an increase in the contribution of these genotypes to the population. Such microevolutionary processes may dampen possible seasonal mismatches between daphnid populations and their food or predator populations.