Genetic markers, genealogies and biogeographic patterns in the cladocera

Cladoceran crustaceans are an important component of zooplankton in a wide range of freshwater habitats. Although the ecological characteristics of several cladoceran species have been well studied, biogeographical studies have been hampered by problematic taxonomic affiliations. However, recently developed molecular techniques, provide a powerful tool to subject aquatic taxa to comparative analyses. Here we highlight recent molecular approaches in aquatic ecology by presenting a simple method of DNA preparation and PCR amplification of the mitochondrial DNA (16S rDNA) in species from nine different families within the cladocera. On a broad taxonomic scale, sequence analysis of this mtDNA fragment has been used to produce the first molecular based phylogeny of the cladocera. This analysis clustered the cladoceran families in a fashion similar to that suggested by previous systematic classifications. In a more detailed analysis of the family Daphniidae, nuclear randomly amplified polymorphic DNA (RAPD), mitochondrial restriction fragment length polymorphism (RFLP) and morphological analyses were combined to identify species and interspecific hybrids within the Daphnia galeata species complex across 50 lakes in 13 European countries and one lake in Africa. The study revealed interspecific hybridization and backcrossing between some taxa (D. cucullata and D. galeata) to be widespread, and species and hybrids to frequently occur in sympatry. Genetic, as well as morphological information, suggests the occurrence of D. hyalina outside the Holarctic.     

Predator-induced life-history changes and the coexistence of five taxa in a Daphnia species complex

Interspecific hybridization is common in water fleas of the Daphnia galeata species complex (e.g. D. galeata , D. cucullata , D. hyalina and their interspecific hybrids). We studied the effect of fish on the life histories of five taxa of this species complex originating from the Plußsee, northern Germany. Using four clones per taxon, we found that fish kairomones reduce size at birth and size at maturity significantly. For size at maturity larger taxa showed a significantly stronger reaction to fish kairomones than the smaller taxa. With respect to the intrinsic rate of increase, r , we compared two predation regimes (positive size selective and not selective). We found that under the fish predation regime most clones had a higher r when cultured with fish kairomones, leading to a higher r , and a stronger reaction for the smaller taxa. We conclude that fish predation might be an important factor influencing the co-occurrence of Daphnia parental taxa with their hybrids.     

How do migrating daphnids cope with fish predation risk in the epilimnion under anoxic conditions in the hypolimnion?

The vertical distribution of Daphnia in experimental tubes isinfluenced by fish kairomone concentration in the ‘epilimnion’and oxygen conditions in the ‘hypolimnion’. Daphnidstrade off reduced predation mortality in a refuge against disadvantagesdue to unfavourable oxygen conditions.     

The ecological forecast horizon,and examples of its uses and determinants

Forecasts of ecological dynamics in changing environments are increasingly important, and are available for a plethora of variables, such as species abundance and distribution, community structure and ecosystem processes. There is, however, a general absence of knowledge about how far into the future, or other dimensions (space, temperature, phylogenetic distance), useful ecological forecasts can be made, and about how features of ecological systems relate to these distances. The ecological forecast horizon is the dimensional distance for which useful forecasts can be made. Five case studies illustrate the influence of various sources of uncertainty (e.g. parameter uncertainty, environmental variation, demographic stochasticity and evolution), level of ecological organisation (e.g. population or community), and organismal properties (e.g. body size or number of trophic links) on temporal, spatial and phylogenetic forecast horizons. Insights from these case studies demonstrate that the ecological forecast horizon is a flexible and powerful tool for researching and communicating ecological predictability. It also has potential for motivating and guiding agenda setting for ecological forecasting research and development.     

Starving with a full gut? Effect of suspended particles on the fitness of <Emphasis Type="Italic">Daphnia hyalina</Emphasis>

Suspended particles are abiotic factors that can affect the abundance of cladocerans such as daphnids. Ultra-oligotrophic Lake Brienz, situated in the front ranges of the Swiss Alps, is dominated by two major inflows that annually transport over 300,000 tons of suspended glacial material into the lake. A laboratory flow-through experiment was performed to test whether these suspended particles have an influence on the fitness of Daphnia hyalina from Lake Brienz, measured as body size, fecundity and juvenile growth rate, especially when they are simultaneously exposed to low food concentrations. Our results show that the concentration of suspended particles present in Lake Brienz does not reduce the fitness of daphnids, even at very low food concentration. In fact, a low concentration of suspended particles increased the fitness. Reduction of fitness could only be observed at a suspended particle concentration of over 25 mg l⁻¹, a level that has never been recorded in Lake Brienz. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

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).     

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.     

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.     

Rapid evolutionary loss of metal resistance revealed by hatching decades‐old eggs

We investigated the evolutionary response of an ecologically important freshwater crustacean, Daphnia, to a rapidly changing toxin environment. From the 1920s until the 1960s, the use of leaded gasoline caused the aquatic concentration of Pb to increase at least fivefold, presumably exerting rapid selective pressure on organisms for resistance. We predicted that Daphnia from this time of intense pollution would display greater resistance than those hatched from times of lower pollution. This question was addressed directly using the resurrection ecology approach, whereby dormant propagules from focal time periods were hatched and compared. We hatched several Daphnia genotypes from each of two Swiss lakes, during times of higher (1960s /1980s) and lower (2000s) lead stress, and compared their life histories under different laboratory levels of this stressor. Modern Daphnia had significantly reduced fitness, measured as the population growth rate (λ), when exposed to lead, whereas those genotypes hatched from times of high lead pollution did not display this reduction. These phenotypic differences contrast with only slight differences measured at neutral loci. We infer that Daphnia in these lakes were able to rapidly adapt to increasing lead concentrations, and just as rapidly lost this adaptation when the stressor was removed.     

Parasites driving host diversity: Incidence of disease correlated with Daphnia clonal turnover*

According to the Red Queen hypothesis, clonal diversity in asexual populations could be maintained by negative frequency‐dependant selection by coevolving parasites. If common clones are selected against and rare clones gain a concomitant advantage, we expect that clonal turnover should be faster during parasite epidemics than between them. We tested this hypothesis exploring field data of the Daphnia–Caullerya host–parasite system. The clonal make‐up and turnover of the Daphnia host population was tracked with high temporal resolution from 1998 until 2013, using first allozyme and later microsatellite markers. Significant differences in the clonal composition between random and infected subsamples of Daphnia populations were detected on six of seven tested occasions, confirming genetic specificity of the host–parasite interaction in this system. We used time series analysis to compare the rates of host clonal turnover to the incidence of parasitism, and found that Caullerya prevalence was significantly associated with microsatellite‐based clonal turnover. As alternate hypotheses, we further tested whether turnover was related to a variety of biotic, abiotic, and host demographic parameters. Other significant correlates of turnover were cyanobacterial biomass and (weakly) temperature. Overall, parasitism seems to be a strong driver of host clonal turnover, in support of the Red Queen hypothesis.