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.     

Seasonal variations in the condition of two Daphnia species and their hybrid in a eutrophic lake: evidence for food limitation

As seston and chlorophyll concentrations in eutrophic lakesare usually high all year round, it was hypothesized in thepast that food limitation is of major importance for the seasonaldynamics of herbivorous zooplankton in such lakes. Since directmeasurements of food are hampered by the lack of knowledge onthe exact nature of the food in eutrophic conditions, indirectmeasurements are necessary to estimate the degree of food limitationin these circumstances. Hence, we used laboratory- and field-derivedrelationships between the body length and body carbon contentof different species of Daphnia, which were collected from thehighly eutrophic Tjeukemeer, the Netherlands. From a seasonalsurvey of the carbon content of the daphnids. we concluded thatin Tjeukemeer D.galeata, D.galeata x cucullata and D.cucullataare food limited during the largest part of the year. Sincethe condition of the hybrids was relatively high as comparedwith the parental species when the food concentration was high,D.galeata x cucutlaia is expected to be the more successfultaxon during periods of high food availability.     

Daphnia invasion: population dynamics of Daphnia assemblages in two eutrophic lakes with particular reference to the introduced alien Daphnia ambigua

The population dynamics of Daphnia species (D.cucullata, D.ambigua,D.galeata) in two small, productive lakes were studied and experimentswith field enclosures were carried out to elucidate the ecologicalniches of the introduced, alien species D.ambigua. Daphnia ambiguocoexisted temporally with the similarly sized D.cucullata, butlived in deeper water. Although both species had the same clutchsizes, similar proportions of ovigerous females, and death rates.D.cucullata was nurnerically dominant over D.ambigua. This suggeststhat the observed difference in abundance is simply a resultof the slower population growth rates of D.ambigua at lowertemperatures in greater depths. The experiments with field enclosuresshowed that when D.culcullata and D.ambigua were forced to livein the same layer of water, D.ambigua was numerically dominantover D.culullata both in the enclosures with and without fish.As the difference between the abundances of the two speciesdecreased in the enclosure with fish, D.cucullata seems to bebetter protected against fish predation than D.ambigua. In theenclosures without fish, D.ambigua and D.cucullata were numericallysurpassed by the larger species D.galeara. The present resultsimply that in some productive lakes with a high fish stock,D.mbigua invaded the niche of D.galeara and that the speciescan become an important component of the native zooplankton.The results also support the opinion that co-existing Daphniacan have very similar dynamics and niches. ¹ Present address: Zoologisches Institut, Christian-Albrechts-UniversitätKiel, Olshausenstrasse 40, D-24098 Kiel, FRG     

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.     

Spatial and temporal patterns of sexual reproduction in a hybrid Daphnia species complex

Evidence for extensive interspecific hybridization among speciesof the genus Daphnia has been accumulating on a global scale.Although there is evidence for limited gene flow between taxavia hybridization, many species still maintain discrete morphologicaland molecular characteristics. We studied temporal and spatialpatterns of sexual reproduction within the Daphnia galeata–hyalina–cucullatahybrid species complex in a lake (Plußsee), located innorthern Germany. Allozyme electrophoresis allowed us to trackseasonal changes in taxon composition as well as the quantificationof back-crosses. Sexually-reproducing animals (ephippial femalesand males) were mainly found in autumn. The simultaneous presenceof sexual morphs of D. galeata and D. galeata x hyalina withthe dominant D. hyalina taxa makes recent hybridization, aswell as back-crossing, plausible. Males and ephippial femalesof D. hyalina were not back-crossed as were the parthenogeneticfemales. The low number of sexual clones of the hybrid D. galeatax hyalina might reflect its reduced fertility, although thesefew clones were detected in high densities. Only hybrid-clonesthat had a back-cross genotype (towards D. hyalina) exhibitedephippial females and males. This indicates that male and ephippialfemale production within the Daphnia taxa is not random, whichmight increase the chance for the parental Daphnia species toremain distinct.     

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.     

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.     

Instar-dependent mortality rates of coexisting Daphnia species in Lake Vechten, The Netherlands

In Lake Vechten, population parameters were determined for twocoexisting Daphnia species, D.hyalina and D.cucullata, fromApril 23 to December 10, 1986. Daphnia hyalina is dominant inspring and D.cucullata in summer and autumn. Size frequencydistributions were converted into instar frequency distributionsby using experimental growth curves and field measurements onnewborn size and the size at first reproduction. A discreteevent computer model was used to calculate the instar mortalityrates. Relative importances of instar mortality rates for thepopulation mortality rate were calculated and discussed. Themortality of the juvenile stages was found to be more importantthan adult mortality during most of the sampling season, despiteegg mortality when egg-bearing females are killed. Only in Septemberand October was the mortality of adult instars more important.Correlation coefficients between differences in the rates ofincrease and differences in mean brood size and instar mortalitiesshow that the former are caused mainly by differences in juvenilemortality, the main factor determining the replacement of D.hyalinaby D.cucullata during early summer.     

Zooplankton-induced unicell-colony transformation in Scenedesmus acutus and its effect on growth of herbivore Daphnia

The effects of colonial ecomorphs of the green alga Scenedesmus acutus on growth of Daphnia cucullata and D. pulex were examined. In ecologically relevant densities (up to 200 animals l^-1) the relatively small D. cucullata did not induce colony formation in Scenedesmus acutus, whereas the larger congener D. pulex significantly promoted colony formation. Both clearance rate and population growth rate (r) were significantly lower in D. cucullata when fed colonial Scenedesmus then when fed unicellular food. However, for D. pulex no effects of food type were observed. These results show that large Daphnia may influence its food in such a way that smaller congeners and competitors are negatively affected.     

Morphological Variation in Daphnia Cucullata Sars with Progressive Eutrophication of a Polymictic Lowland Reservoir

Morphological variation in Daphnia cucullata Sars was studied from 1994 to 2001 when fast increasing water trophy of the Siemianówka Reservoir was related to high TP and chlorophyll concentrations. During this period, D. cucullata biomass, as well as its contribution to the total cladocerans biomass clearly decreased. We found that among the six morphological parameters analyzed in D. cucullata, body length, height of the head and length of the tail spine had a tendency to decrease. The maximum values of these traits and carapace length also decreased. The morphology of D. cucullata appeared to be influenced by factors such as food availability, fish predation and an exclusive presence of D. cucullata among Daphniidae species. Increased water mixing in the reservoir may have also affected the changes in the head inclination of D. cucullata.     

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.     

Plasticity in filtering screens of Daphnia cucullata×galeata hybrids and parental species at two food concentrations

The coexistence of Daphnia cucullata × galeata hybrids with the parental species D. galeata and D. cucullata was investigated by measuring areas and mesh sizes of filtering structures of these herbivorous zooplankton taxa cultivated at low and high food concentrations. The clearance rates and somatic growth rates were also determined. When reared at low food concentration, all taxa had larger filtering areas. Larger filtering areas also resulted in higher clearance rates. Differences between taxa in both filtering area and clearance rate were caused mainly by interspecific size differences. Hybrids had the largest absolute mesh sizes, and the parental species had smaller mesh sizes. Hybrids also showed heterosis in somatic growth rate at high food concentration. The observed taxon-specific differences in mesh size and somatic growth rate contribute to resource partitioning between the taxa and thus to their successful coexistence in lakes. Received: 1 March 1999 / Accepted: 25 May 1999     

Instar-specific mortalities of coexisting Daphnia species in relation to food and invertebrate predation

Instar-specific mortalities of Daphnia hyalina and D.cucullatawere studied from May 19 to September 29, 1988 in combinationwith invertebrate predator and phytoplankton abundance. Simultaneouslife-table experiments were conducted under semi-natural conditionsin the laboratory to estimate juvenile mortality in a predator-freeenvironment. Juvenile mortality by predation was calculatedas the difference between juvenile mortality in the field andin the experiments and was the most important factor for thedifferences in abundance of the two species. For D.hyalina juvenilemortality was higher in early summer and probably caused byselective predation by Chaoborus flavicans. Predation by Leptodorakindtii was probably more important during the rest of the summer.Estimated mortality by predation adequately explained juvenilemortality, except for a 3-week period in August. Decreasingflagellate densities in July were accompanied by increased juvenilemortalities of D.hyalina and D.cucullata in the life-table experimentsin August and coincided with a Daphnia population decrease.     

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.     

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.     

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.     

Population dynamics and production of Daphnia hyalina Leydig and Daphnia cucullata Sars in Tjeukemeer

As part of a research programme on the food chains in Tjeukemeer, the Daphnia hyalina and Daphnia cucullata populations were studied for three successive years. To analyse the factors regulating the production of these two species, their population parameters (density, size distribution, fecundity) and population dynamics (birth rate, mortality rate) were studied and related to environmental factors. Since Daphnia in Tjeukemeer shows continuous recruitment, the population dynamics model INSTAR was developed and used to integrate field data with laboratory data on development rates and length-weight relationships. The dynamics of the Daphnia species are mainly regulated by temperature and fish predation, the latter affects both birth rate and mortality. Total annual Daphnia production was 3.1–6.9 g org. dry wt M^–2, and annual P/B ratio ranged from 25 to 40 for D. cucullata and from 45 to 49 for D. hyalina.     

Extreme helmet formation in Daphnia cucullata induced by small-scale turbulence

Although Daphnia cucullata is used as a textbook example forcyclomorphosis, distinct helmet development, as shown in thefield, has not been demonstrated in the laboratory until now.We show for the first time that small-scale turbulence is ableto induce the maximum response of morphological plasticity inD. cucullata. Helmet elongation reached magnitudes as extremeas observed in the field. Different modes of generating small-scaleturbulence caused different levels of helmet formation. Small-scaleturbulence alone may not be responsible for cyclomorphosis innature because field data show that two nearby lakes with similarmorphometry differ in cyclomorphosis patterns, while laboratoryexperiments show that there is no difference in the abilityto form helmets in the clones of each lake. Although helmetformation in D. cucullata is inducible with predator kairomones,helmet elongation is not as strong as that induced by turbulence.We discuss the possible role of helmets under turbulent conditions.     

Combined effects of food shortage and oxygen deficiency on life history characteristics and filter screens of Daphnia

Daphnia magna were reared in environments with different concentrationsof food and dissolved oxygen, and their life history characteristicsand filter-screen areas were analyzed. Both low food concentrationand low oxygen concentration degraded Daphnia fitness (as measuredby individual growth and reproduction), and synergism was detectedstatistically for the effects of the two factors together. Lowfood concentration induced the development of large filter screensin Daphnia, but this development was suppressed by low oxygenconcentration, suggesting that low oxygen concentration reducesthe filtering rate of Daphnia in a food-limited environment.Therefore, energy uptake by Daphnia seems to be reduced by foodshortages and oxygen deficiencies jointly, and this reductionin energy uptake may be a possible mechanism causing the synergisticeffects of the two environmental stresses on the life historyof Daphnia.     

Synergistic effects of different food species on life-history traits of Daphnia galeata

In this paper we describe the life history consequences of feeding Daphnia galeata with different food types in different concentrations. We fed the animals with four concentrations of two green algae Scenedesmus obliquus and Chlamydomonas globosa, given separately as well as in a 1:1 mixture. Growth, reproduction and condition of the animals and consequently the intrinsic rate of population increase, r, increased with the concentration of the food for all three experimental series. The life history parameters measured were lowest in the animals raised on C. globosa. Animals fed on S. obliquus showed higher values for r. We observed a distinct synergistic effect of the two algal species: growth, reproduction, condition and r were significantly higher for the animals raised on the mixed media as compared to the animals raised on the mono-algal media. We conclude that the two algal species represented complementary food resources.