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.     

Differential life history responses of several pelagic Daphnia clones differing in migratory behaviour

Females of the hybrid Daphnia galeata × hyalina were isolated from a natural population in Lake Maarsseveen and from these individuals clones were cultured in the laboratory. Some clones were assumed to be migrating and some non-migrating. Life history experiments were performed with these clones in the presence and absence of fish kairomones. Results show that clones differed in life history strategy in concert with their presumed migration strategies which suggests a link between life history and behaviour. However, no sharp border exists between migrating and non-migrating clones since a gradual response pattern was found for the traits investigated. In the absence of fish kairomones, the relation between length at maturity and number of eggs was found to differ between the clones of the non-migrating and the migrating group. The latter had a lower number of eggs at similar length values. In the presence of fish kairomones, the length–clutch size relation was the same. However, non-migrating clones were smaller at the age at maturity and had fewer eggs. Although clones of both groups were thus found to differ significantly, overlap existed. Age at maturity was found to be the same.     

The influence of a deep-storage reservoir on the physicochemical limnology of a Central Texas River

A study was conducted to determine a physicochemical profile of a new deep-storage reservoir and to determine the influence of impoundment and thermal stratification in the reservoir on the physicochemical limnology of the parent river. The presence of thermal stratification from May through November caused the most significant change in water conditions. Of 23 parameters studied, 12 remained unchanged, 10 improved, and 2 deteriorated (Table VI).The greatest downstream changes in water conditions from those upstream from the reservoir were a decrease in temperature, an increase of ammonia, and the presence of hydrogen sulfide during the period of thermal stratification. Ammonia did not increase to a level considered to be toxic to aquatic species. It could, however, serve as a nutrient for certain species of plants and result in a change in community structure.Water tempeature downstream from the reservoir was always within the annual temperature range of the river upstream from the reservoir; however, the summer maximum in the tailrace was decreased to a temperature that could interfere wih the normal life cycle of many species.     

Reproductive effort in Daphnia galeata: comparison of reservoir and pond populations

Populations of Daphnia galeata from the deep stratified moderately eutrophic reservoir and the shallow highly eutrophic carp ponds differed in the reproductive effort of the first laboratory generation cultivated at a rich algal diet. The reservoir animals were smaller at the first adult instar and exhibited longer time between hatching of one brood and forming the eggs of the next one.     

How could thermal stratification affect horizontal distribution of depth-integrated metazooplankton communities in the Gulf of Gabes (Tunisia)?

The spatial and temporal variability of metazooplankton communities in relation to environmental and trophic factors were studied in the Gulf of Gabes during three periods of increasing thermal stratification in June, July and September. Non-metric multi-dimensional scaling (NMDS) revealed the time and space succession of three zooplankton communities in relation to the degree of thermal stratification. A co-inertia analysis showed the association between these communities and the trophic conditions (abundance and composition of microplankton) driven by stratification. Three divergent communities were evident. The first, characterized by annelid larvae and coastal copepods, was associated with shallow mixed and microplankton-rich water (June, coastal). The second, characterized by euphausiids, amphipods, doliolids and several deep-water copepods, corresponded to highly stratified microplankton-poor water (September). The third, characterized by the most ubiquitous and abundant species, was linked to intermediate water (June deep and July). Temperature and salinity conditions also explained the distribution of key species. Clausocalanus furcatus, Oithona plumifera and Triconia conifera were clearly associated with warm and highly stratified water (September). Enhanced thermal stratification led to higher surface salinity, explaining the emergence of euryhaline taxa such as Oithona plumifera.     

Dietary essential amino acids affect the reproduction of the keystone herbivore Daphnia pulex

Recent studies have indicated that nitrogen availability can be an important determinant of primary production in freshwater lakes and that herbivore growth can be limited by low dietary nitrogen availability. Furthermore, a lack of specific essential nitrogenous biochemicals (such as essential amino acids) might be another important constraint on the fitness of consumers. This might be of particular importance for cladoceran zooplankton, which can switch between two alternative reproductive strategies--the production of subitaneously developing and resting eggs. Here, we hypothesize that both the somatic growth and the type of reproduction of the aquatic keystone herbivore Daphnia is limited by the availability of specific essential amino acids in the diet. In laboratory experiments, we investigated this hypothesis by feeding a high quality phytoplankton organism (Cryptomonas) and a green alga of moderate nutritional quality (Chlamydomonas) to a clone of Daphnia pulex with and without the addition of essential amino acids. The somatic growth of D. pulex differed between the algae of different nutritional quality, but not dependent on the addition of dissolved amino acids. However, in reproduction experiments, where moderate crowding conditions at saturating food quantities were applied, addition of the essential amino acids arginine and histidine (but not lysine and threonine) increased the total number and the developmental stage of subitaneous eggs. While D. pulex did not produce resting eggs on Cryptomonas, relatively high numbers of resting eggs were released on Chlamydomonas. When arginine and histidine were added to the green algal diet, the production of resting eggs was effectively suppressed. This demonstrates the high, but previously overlooked importance of single essential amino acids for the reproductive strategy of the aquatic keystone herbivore Daphnia.     

水蚤滞育的数量遗传学研究:遗传与环境控制及遗传作用

在单性生殖循环水蚤群体中,滞育卵由有性生殖产生．在一系列实验中用到了不同种群和种类的水蚤,通过这些实验来观察：1)有性生殖和滞育卵复苏的遗传和环境控制;2)生活在相同区域中,但有不同生存微环境的近缘种类的有性生殖的光周期反应;3)在群体遗传结构上有性生殖的遗传效应(基因型均值和生活史性状遗传方差)．结果发现：1)遗传作用和环境作用,以及两者的相互作用都对有性生殖和滞育卵的孵化有显著的影响．GE显著的相互作用对环境中观察到的有性生殖来说,有助于维持其较高的遗传方差;2)在相同区域中,不同生存微环境的近缘种类的有性生殖的光周期反应有所不同．这有助于进一步区别近缘的水蚤种类,这也可能是一个水环境中同素异形的物种形成的例子;3)在有性生殖上,生活史性状平均值和遗传方差变化与前代选择造成的均值和遗传方差相反(遗传滑阻),这会造成暂时的适应不良(遗传滑阻和隐藏的遗传变异的表达),应对它补偿滞育的进化优势．     

Invasibility of a reservoir to exotic Daphnia lumholtzi: experimental assessment of diet selection and life history responses to cyanobacteria

SUMMARY 1. In reservoirs of the south-central United States, the exotic cladoceran Daphnia lumholtzi is common during warm midsummer conditions, when cyanobacteria are abundant and native Daphnia are rare. In the current study, we employed surveys, field experiments, and a life table experiment to investigate the role of food quality in explaining the distribution and phenology of D. lumholtzi , relative to two native species ( Daphnia parvula and Daphnia mendotae ).
2. During May–September 2000 in eutrophic McDaniel Lake, Missouri U.S.A., cyanobacteria (primarily Oscillatoria ) first appeared at 6-m depth and then became abundant throughout the epilimnion.
3. During the May field experiment, D. lumholtzi , D. parvula and D. mendotae all consumed a similar diet of algae, showing positive selection for small greens (chlorophytes and cryptophytes <20 μm). During the July experiment, when the epilimnion exceeded 25 °C and cyanobacteria were common in the lake, D. lumholtzi consumed significantly more total algae and more cyanobacteria than the two native species. Although the Daphnia selected against cyanobacteria, all three species consumed about 25% of this food in their diet.
4. A life table experiment compared the responses of D. lumholtzi and D. parvula with variation in density of high-quality food ( Ankistrodesmus ) and concentration of a toxic strain of cyanobacteria ( Anabaena flos-aquae ). Both Daphnia species showed reduced survivorship, fertility and intrinsic rates of increase in response to elevated concentrations of cyanobacteria, particularly at the higher food level.
5. The results suggest that D. lumholtzi shows similar inhibition from cyanobacteria as does the native Daphnia . However, their continued high in situ feeding rates imply that D. lumholtzi is less affected by midsummer conditions in warm-water reservoirs than are native Daphnia .     

Plasticity of growth rate and metabolism in Daphnia magna populations from different thermal habitats

The aim of this study was to evaluate the effect of temperature on growth and aerobic metabolism in clones of Daphnia magna from different thermal regimes. Growth rate (increment in size), somatic juvenile growth rate (increment in mass), and oxygen consumption were measured at 15 and 25 degrees C in 21 clones from one northern and two southern sites. There were no significant differences in body size and growth rate (increase in length) at both 15 and 25 degrees C among the three sites. Clones from southern site 2 had a higher mass increment than clones from the other two sites at both temperatures. Clone had a significant effect on growth (body length) and body size at both temperatures. As expected, age at maturity was lower at 25 degrees C (4.5 days) than at 15 degrees C, (11.6 days) and body sizes, after the release of the third clutch, were larger at 15 degrees C than at 25 degrees C. Northern clones had higher oxygen consumption rates and specific dynamic action (SDA) than southern clones at 15 degrees C. By contrast, southern clones from site 1 had a higher oxygen consumption and SDA than subarctic clones at 25 degrees C. Clones from southern site 2 had high oxygen consumption rates at both temperatures. Our results reveal important differences in metabolic rates among Daphnia from different thermal regimes, which were not always reflected in growth rate differences.     

Chronic responses of different clones of Daphnia longispina (field and ephippia) to different food levels

Resting eggs are a fundamental reproductive strategy among freshwater cladocerans. Under adverse environmental conditions, whole Daphnia populations can disappear from a lake and a new community will arise from ephippial eggs. Since these new populations are subjected to genetic variation, their responses to environmental stress or contaminants can be different from the “original” population. In the present study, life history responses (reproduction and growth) of Daphnia longispina to different food concentrations was studied. Two Daphnia populations were tested: (a) field clones and (b) ephippial clones. Food (Selenastrum capricornutum) concentration was the stressor tested (absence of food, and low to high food concentrations). The results showed that reproductive responses of D. longispina to the tested food concentrations varied among clones, independently from their origin.     

Performance trade-off across a natural resource gradient

An important environmental factor determining both phytoplankton and zooplankton community composition is lake depth and thermal stratification. However, there is little information on how the interaction between zooplankton grazers and their phytoplankton food changes along an environmental gradient of lake depth. We contrasted resource availability for daphniid zooplankton populations living in two shallow, unstratified lakes and in two deep, stratified lakes using a novel growth bioassay. Stratified lakes had consistently lower resource richness than shallow unstratified lakes. To test whether resources were important in explaining differences in daphniid composition of shallow and deep lakes, we performed reciprocal transplant experiments. We raised daphniids typical of shallow (Ceriodaphnia reticulata) and deep (Daphnia dentifera) lakes in the resources from replicate shallow and deep lakes and monitored survival and reproduction. The two species exhibited a performance trade-off, measured by life table r and R ₀, across a gradient in natural resource richness. D. dentifera had higher relative fitness than C. reticulata when raised in the poorest resource environment from a deep lake. However, under richer resource conditions typical of shallow lakes, C. reticulata outperformed D. dentifera. We further created a gradient in natural resource quantity (by dilution) to test whether this trade-off in species relative fitness involved aspects of resource quality. No trade-off in species performance was evident across the dilution gradient, indicating that resource quality was important to the trade-off. We conclude that shifts in daphniid species composition along a gradient of lake depth involve an adaptive trade-off in ability to exploit rich versus poor resource quality. Received: 11 May 1998 / Accepted: 15 January 1999     

Vertical distribution of pelagial zooplankton in a middle-sized dimictic valley reservoir

Our observations indicate the vertical distribution of zooplankton and its seasonal changes in Dubník II reservoir (Slovakia) are determined mainly by the thermal regime of the reservoir, by transparency, and by fish and invertebrate predation. During periods of circulation, zooplankton vertical distribution in the whole water column was more homogeneous, whilst during summer temperature stratification zooplankton concentrated in the epilimnion — rotifers in higher layers than crustaceans. During summer stagnation a steep thermal gradient occurred at the boundary of the epi-and hypolimnion and low temperature and low dissolved oxygen in hypolimnion offered a refuge for Chaoborus flavicans larvae against fish, enabling coexistence of vertebrate and invertebrate predation. This evidence supports our previous findings concerning dominance of rotifers in zooplankton and representation of crustaceans by small-bodied species in the study reservoir. Steep thermal gradient and the presence of Chaoborus larvae caused very low zooplankton abundance in the lower part of the water column and a reduction of cladocerans refuges against fish to layers of thermocline or closely under thermocline where Daphnia cucullata and Daphnia parvula were found. Our previous assumptions about the high density of zooplanktivorous fish in Dubník II reservoir are supported by the fact that these small cladocerans are represented by smaller individuals in the upper layers and bigger individuals in deeper layers.     

Life histories and habitat selection in Daphnia: divergent life histories of D. magna clones differing in phototactic behaviour

To test the hypothesis of co-adaptation of life histories and daytime vertical distribution (vertical migration behaviour) in Daphnia, life history characteristics were analysed for two positively, three negatively, and four intermediately phototactic Daphnia magna clones. Clones with different phototactic behaviour were found to have divergent life history strategies, with positively phototactic clones being good exploiters under the non-limiting conditions provided in the laboratory, i.e. low density (1 ind./1), high food concentration (6,5–7 10⁵ Scenedesmus cells/ml, restored daily) and high temperature (20° C). They realized a high intrinsic rate of increase at a small adult body size through rapid development, at a cost of producing small neonates. Negatively and intermediately phototactic clones had larger adult body sizes, and produced larger neonates that were more starvation-resistant than those of positively phototactic clones. Selection for high intrinsic rate of increase in intermediately phototactic clones was mediated through the production of large clutches.     

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.     

Coexistence of similar genotypes of Daphnia magna in intermittent populations: response to thermal stress

We investigated the diversity and thermal response of a fitness related trait, juvenile growth rate, in seasonal population samples of Daphnia magna from two temperate ponds. Both populations were intermittent, i.e. they disappeared from the water body and recolonized seasonally by hatching from resting eggs in the sediment.
Temporally isolated clones of Daphnia magna showed the typical asymmetric response for growth rate with temperature and a sharp decline after the maximum response at 26°C (TMR). There was no evidence for genetically adapted seasonal groups. Despite significant genetic variation among clones and for phenotypic plasticity (G×E interactions without genetic correlations), seasonal groups of clones showed no shift in TMR and mean temperature reaction norms were similar among groups and both populations. Heritabilities remained similar among temperatures despite a large increase in genetic variance at stressfully high temperatures of 29°C and 32°C, due to simultaneous increase in environmental variance. Further, heritabilities remained high among sample periods and were not eroded during several months of asexual reproduction.
Regular diapause, an intrinsic feature of intermittent Daphnia populations, may replace the need for physiological temperature adaptation and promote maintenance of diversity through phenotypic similarity by reducing the time over which competitive interactions occur. Such populations are unlikely to be directly affected by elevated temperatures. They have a large potential for phenotypic plasticity as their TMR is higher than the temperature normally encountered.     

Habitat Characteristics of Eurytemora lacustris (Poppe, 1887) (Copepoda,Calanoida): The Role of Lake Depth,Temperature,Oxygen Concentration and Light Intensity

Field observations, laboratory experiments and a literature survey were conducted to evaluate the habitat characteristics of Eurytemora lacustris (Poppe 1887), a freshwater calanoid copepod species. Combined effects of temperature and oxygen concentration in the deep water of thermally stratifying lakes seem to be the ultimate factors governing the occurrence of the species throughout its home‐territory. E. lacustris is largely restricted to relatively deep lakes (>30 m) providing a hypolimnetic refuge characterised by low temperatures (<∼10 °C) and oxygenated water during summer. Therefore, although the species is spread over much of Europe it was only found in a small number of lakes. Long‐term records in different lakes revealed E. lacustris to be perennial with relatively high biomasses occurring from May to September. During thermal stratification on average 87% of the nauplii and 72% of the copepodite biomass was found in hypolimnetic waters colder than 10 °C. Diurnal vertical migration was observed for the copepodid stages, but the migration amplitude clearly decreased from May to September. The migration amplitude was significantly related to light intensity. According to its special habitat requirements, E. lacustris might be considered a glacial relict sensitive to temperature increase and oxygen depletion. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Evolution of predator avoidance in a Daphnia population: evidence from the egg bank

Dormant propagule pools (egg banks) of zooplankton populations that accumulate in sediments provide biological archives of past conditions and enable the investigation of evolutionary changes in populations over relatively long periods of time (many decades). This study examined the egg bank of a Daphnia pulicaria population in a lake that has been stocked annually with rainbow trout (a zooplanktivore) since 1961. Resting eggs from sediments from the 1920s to 2001 (determined by ²¹⁰Pb dating) were hatched and established as isofemale clonal lines. The phototactic behavior (a proxy for vertical migration behavior) of clones was assessed in the presence and absence of fish kairomones to evaluate the hypothesis that the heavy and consistent level of predation imposed by the stocked trout would have selected for clones that are more negatively phototactic. In addition, exposure to kairomones was expected to induce stronger negative phototaxis for all clones relative to control conditions. The behavior of clones derived from resting eggs from the trout era (after 1961) was significantly more negatively phototactic than it was for pre-trout era clones. Kairomone exposure induced a more negative phototactic response in clones from both eras, but the response was much greater for the pre-trout era clones. These results suggest that the consistently high level of predation by trout over several decades has selected for a conservative (less plastic) vertical migration strategy in which Daphnia maintain a deep daytime distribution with or without the presence of chemical cues from fish predators.     

PHENOTYPE FIXATION AND GENOTYPIC DIVERSITY IN THE COMPLEX LIFE CYCLE OF THE APHID PEMPHIGUS BETAE

The aphid Pemphigus betae typically shows a complex life cycle, with annual alternation between cottonwood trees, where it forms leaf galls, and herbaceous plants, where it lives on roots. Distinct phenotypes are associated with each phase. In a population in Utah, aphid clones vary in their tendencies to undergo the cottonwood phase of the life cycle, with certain clones rarely producing the winged migrants that initiate the cottonwood phase. To examine the extent of genotypic variability in life cycle traits and to determine the consequences of phase deletion for fitness in the remaining phase of the life cycle, I compared performances of sets of clones that differed in their tendencies to delete the cottonwood phase. Performances of 35 aphid clones were compared on root hosts in the laboratory, by quantifying size (a correlate of fecundity), developmental time, and mortality for clonal sublines. Clones that tend to delete the cottonwood phase and remain on roots developed faster, showed lower mortality, and were larger as compared to clones that tend to undergo an annual migration to cottonwood. These laboratory comparisons of individual clones agree with earlier findings from field experiments. Performance also varied significantly among clones within sets, with the set of clones not previously subjected to selection on root hosts showing higher variance than clones isolated following selection in the root-inhabiting phase. These data suggest that selection in the cottonwood phase opposes selection in the root phase. Thus, even in cases of polyphenism, in which divergent developmental pathways confer some independence of phenotypic expression in alternative phases, antagonistic pleiotropy limits adaptation within a phase. These findings on the correlates of intrapopulational variation in frequency of phase expression provide some of the best evidence for character release following reduction of a complex life cycle, and they support the hypothesis that the advantages of polyphenism may be limited by negative genetic correlations among fitness traits of alternative phenotypes.     

广东沿海与粤北山区中小型水库浮游生物群落比较分析

于丰水期和枯水期对广东山区水库(赤石迳水库)、沿海水库(大水桥、赤沙水库)三个水库的湖泊区和河流区进行研究,分析水质和浮游生物群落结构。三个水库之间的浮游生物群落均有所区别,地理位置和地形不同而引起的温度差异以及外源性营养盐输入是造成浮游生物种类和数量差异的主要原因。沿海水库不分层,而山区水库丰水期温跃层明显。各水库的氮磷比均较高,chl.a水平主要受磷限制,沿海水库则多喜暖性的种类,如裸藻、隐藻、沙居剑水蚤等,山区水库发现较多喜冷性浮游生物,如金藻、透明溞等,丰水期的甲藻丰度也相对高于枯水期。     

Influence of Temperature on Exotic Daphnia lumholtzi and Implications for Invasion Success

We investigated the effects of water temperature on the exoticcladoceran Daphnia lumholtzi in a eutrophic Kansas reservoir(USA) and under laboratory conditions. Daphnia lumholtzi demonstrateda distinct late summer appearance in the reservoir at temperaturesbetween 26 and 31°C, which corresponded with steep declinesin the densities of native Daphnia spp. Laboratory life-tableexperiments confirmed that D. lumholtzi performs well at elevatedwater temperatures. The intrinsic rate of increase (r), netreproductive rate (R_o), age at first reproduction, survivorship(l_x) and molting rates all demonstrate that D. lumholtzi hasa high temperature optimum between 20 and 30°C. A comparisonof literature-reported r-values indicates that the reproductiverate of D. lumholtzi is comparable with other Daphnia spp. between20 and 25°C, but also implies that D. lumholtzi may out-performsome Daphnia spp. at temperatures >25°C. Collectively,these results suggest that D. lumholtzi may be taking advantageof a late summer thermal niche, and that this invader may continueto colonize lakes and reservoirs in the southern US. However,life table data also indicate that D. lumholtzi performs poorlyat temperatures < 10°C, which may inhibit the range expansionof this invader into northern waters.