Respiratory electron transport system (ETS) activity as an estimator of the thermal tolerance of two Daphnia hybrids

The influence of temperature on the activity of the respiratoryelectron transport system (ETS) was measured in one clone ofDaphnia hyalina x galeata and one of Daphnia cucullata x galeata,isolated from Lake Bled (Slovenia). The ETS activity of ovigerousfemales acclimated to 7, 20 and 25°C, was measured at 5,10, 15, 20, 25 and 30°C. Population growth experiments showedthat D. cucullata x galeata grew better at high rather thanlow temperatures. Daphnia hyalina x galeata, however, grew moresuccessfully at low temperatures than did D. cucullata x galeata.The highest Q₁₀ of ETS activity of D. cucullata x galeata atthe lowest temperature range of 5–15°C indicated theabsence of enzymes that could function sufficiently well atlow temperatures. The ETS activity of the warm-acclimated hybridD. hyalina x galeata reached a maximum at an incubation temperatureof 20°C, while D. cucullata x galeata had maximal ETS activityat 25°C. Thus D. cucullata x galeata has a more efficientenzyme system than D. hyalina x galeata at the higher temperature.The higher Arrhenius activation energy (E_a) for D. cucullatax galeata than for D. hyalina x galeata indicates that enzymesfrom D. cucullata x galeata are more temperature sensitive thanthose from D. hyalina x galeata. In conclusion, the ETS of D.cucullata x galeata is adapted to a higher temperature and tonarrower temperature fluctuations than that of D. hyalina xgaleata.     

Behaviour plasticity without learning: phenotypic and genetic variation of naïve Daphnia in an ecological trade-off

The swimming behaviour of adult Daphnia largely governs their depth, which has a direct effect on both individual foraging success and predation avoidance. We treated individual swimming behaviour as a threshold character and used directional changes in average clonal depth within experimental tubes as a test for character plasticity. We compared the swimming behaviours of two cohabiting, phenotypically similar Daphnia (Daphnia galeata and Daphnia galeata-Daphnia rosea hybrid) to determine (1) whether there is inherited variation (H(2)) for different traits (responses to hunger and predator cues); (2) whether changes in genetic parameters (norm of reaction and character state) across four environments could be simulated by combinations of the presence or absence of a predator cue and high or low hunger levels; and (3) whether these Daphnia would respond to directional selection, particularly in a trade-off environment (high hunger and predator cue treatments). Responses of both D. galeata and the D. galeata-rosea hybrid to hunger and a predator cue and the trade-off environment were plastic. Daphnia galeata expressed significant genetic variation within (H(2)) and between environments (heritability of plasticity). Both the character state and norm of reaction estimates of heritable variation in the hybrid were close to zero. Genetic correlations were positive and stable across six environmental pairs in Daphnia galeata. Most hybrid genetic correlations were not significant. The phenotypic distributions of both D. galeata and the hybrid were bimodal in the trade-off environment. The D. galeata distribution was partly due to between-clone variation and the hybrid distribution was almost entirely due to within-clone variation. Genetic variation expressed by D. galeata in the trade-off environment appears to depend on both clone by environment interactions and stable inherited differences. These results indicate that while plastic phenotypic responses cause a similar opportunity for selection in D. galeata and the hybrid, their responses to selection would be different in the trade-off and in related environments. Copyright 2000 The Association for the Study of Animal Behaviour.     

Genetic assessment of species boundaries in the North American Daphnia longispina complex (Crustacea: Daphniidae)

Species boundaries in the North American Daphnia longispina group have proved difficult to establish on the basis of morphology alone. This confusion may be due to hydridization, phenotypic plasticity or the existence of sibling species. We therefore used genetic analysis to delineate species boundaries by examining 27 North American populations belonging to the longispina complex for variation at 15–26 allozyme loci. The populations consisted of Daphnia thorata from two western sites and two eastern sites, Daphnia galeata mendotae from its type location and seven sites across its range, and Daphnia rosea from eight temperate and seven arctic sites. Two populations from the Eurasian longispina complex were also included for reference. Populations assigned to D. galeata mendotae formed a genetically cohesive group, whereas a genetic dichotomy was found between temperate and arctic D. rosea , suggesting that this taxon includes two species. Genetic analysis also confirmed the distinctness of western D. thorata from other members of the longispina group. Unexpectedly, eastern populations resembling D. thorata were genetically more similar to temperate D. rosea than to any helmeted species ( D. galeata, Daphnia hyalina or D. thorata ). Our results suggest that the helmet character is a poor indicator of phylogenetic relationships, as the genetic ability to produce this feature has been lost or acquired several times in the evolution of the longispina group.     

基于16S rDNA和COⅠ基因序列探讨四种溞类的系统关系和分类地位

为了检验已记录的4个溞属种类（(隆线溞Daphnia carinata、透明溞D. hyalina、蚤状溞D. pulex和大型溞D. magna）的系统分类, 用试剂盒法分别提取4种溞类的基因组DNA。利用特异性引物, 通过PCR扩增了4种溞属种类的16S rDNA和COⅠ基因部分序列, 并与来自GenBank中每个种类相似度较高的同种序列进行分析。结果表明, 基于16S rDNA和COⅠ基因, 4个溞属种类的平均种间相似度分别为85.56%和80.67%, 碱基中A+T含量均明显高于G+C含量。基于COⅠ基因, 巢湖隆线溞与来自GenBank上的拟同形溞（(Daphnia similoides AB549199）)相似度为99%, 分歧度为0.4%0.9%。就16S rDNA和COⅠ基因而言, 巢湖透明溞更接近于盔形溞（Daphnia galeata）。基于16S rDNA和COⅠ基因构建的NJ树和贝叶斯树, 两者的主要分枝基本一致。结果暗示, 巢湖隆线溞与拟同形溞为一个种, 透明溞应属于盔形溞。由于缺乏核基因的研究, 有关巢湖溞属的分类地位还需进行更深入的探讨。       

Effects of food type on the life history of Daphnia clones from lakes differing in trophic state. I. Daphnia galeata feeding on Scenedesmus and Oscillatoria*

1. Effects of the green alga Scenedesmus obliquus and the filamentous cyanobacterium Oscillatoria limnetica as food sources on the growth and reproduction of nine Daphnia galeata clones were studied. A high concentration of these two algae species was fed to Daphnia clones collected from four lakes with varying trophic conditions in a life history experiment.
2. Although Oscillatoria is regarded as a low quality food for Daphnia because of its filamentous morphology and suspected nutritional inadequacy, the cyanobacterium sustained both body growth and reproduction in D. galeata. Growth and reproduction were, however, lower and the onset of reproduction was delayed on a diet of Oscillatoria compared to a diet of Scenedesmus. The intrinsic growth rate, r , for each D. galeata clone was positive on a diet of Oscillatoria , ranging from 0.06 to 0.16 mm day^–1, whereas the intrinsic growth rates on a diet of Scenedesmus ranged from 0.27 to 0.34 mm day^–1.
3. The degree of growth depression on the Oscillatoria diet varied among clones. Clonal responses were correlated with body size; clones with larger body size decreased their growth and reproduction more on a diet of Oscillatoria. The responses of different clones from the same lake to the two diets resembled each other, irrespective of geography or trophic state of the lake.     

Phytoplankton responses to grazing by Daphnia galeata in the biomanipulated Bautzen reservoir

We studied the response of phytoplankton to grazing by Daphnia galeata in the hypertrophic Bautzen reservoir (Dresden, Germany) from January 1995 to May 1996 during a long-term whole-lake biomanipulation experiment. The correlation between clearance rate of D. galeata and total phytoplankton biomass was negative only if biomass of Microcystis aeruginosa was excluded. This suggests that M. aeruginosa is the main grazing resistant phytoplankton species in the Bautzen reservoir. Except for M. aeruginosa and grazing-resistant Staurastrum quadridentatum spec. nov. (Scharf, 1995) no other phytoplankton species was able to reach a biovolume above 1 mm3 L-1 when the clearance rate of Daphnia exceeded 0.1 L L-1 d-1. There was significant positive correlation between mean cell or colony size of phytoplankters and clearance rate of D. galeata, showing an advantage of bigger cells or colonies at high grazing pressure. Cross-correlation indicated a time lag of about one month between changes in grazing pressure and a change in phytoplankton mean size. The phytoplankton species were divided into edible and inedible fractions, depending on their width and length. No edible species were able to reach high biovolumes during high biomasses of D. galeata but a positive correlation was found between the edible fraction of phytoplankton biovolume and the clearance rate of D. galeata. However, this relationship disappeared when the ‘ingestible edible’ fraction of M. aeruginosa was excluded, suggesting a rejection of ‘edible’ Microcystis colonies by D. galeata. A negative correlation was found between the inedible fraction of phytoplankton biovolume without M. aeruginosa and the clearance rate of D. galeata which might be due to superior competition of M. aeruginosa. We could clearly show that biomanipulation might not work well with respect to the reduction of total phytoplankton biomass under hypertrophic conditions and finally discuss a theoretical threshold of phosphorus (probably around 80 μg L-1), below which biomanipulation should become effective. This revised version was published online in July 2006 with corrections to the Cover Date.     

Daphnia hybridization along ecological gradients in pelagic environments: the potential for the presence of hybrid zones in plankton

The relative homogeneity of pelagic environments has been regarded as the reason for the absence of hybrid zones for hybridizing planktonic Daphnia (Crustacea: Cladocera); occasional dominance of interspecific hybrids over parental species was explained by their temporal superiority in fluctuating environments. However, water bodies with spatially varying environmental conditions might facilitate the formation of hybrid zones in plankton. We studied the distribution of species and hybrids of the Daphnia longispina complex in 11 canyon-shaped reservoirs, localities characterized by horizontal environmental gradients (particularly of food supply and size-selective predation); we also analysed patterns of carapace size and fecundity among coexisting taxa. Spatial distribution of taxa agreed with their ecological characteristics; those showing different affinities along longitudinal reservoir profiles differed in size according to the presumed fish predation gradient. Only hybrids of Daphnia galeata with Daphnia cucullata and D. longispina (=hyalina) were recorded. The latter two species preferred opposite ends of gradients, such spatial segregation probably explaining the absence of their hybrids. Distributional patterns were relatively stable in two consecutive summers, apart from a substantial decline of D. galeata X cucullata in the second year. The observed pattern of a hybrid-dominated zone in intermediate conditions suggests that local Daphnia hybrid zones may indeed form within reservoirs.     

Alternative antipredator responses of two coexisting Daphnia species to negative size selection by YOY perch

Our study showed for the first time in nature that two coexistingDaphnia adopted alternative life history and behavioural strategiesto cope with negative size-selection predation by gape-limitedyoung-of-the-year (YOY) perch. We evaluated the phenotypic plasticityin life history and behavioural traits of two coexisting Daphniaspecies, D. pulicaria (2 mm) and D. galeata mendotae (1.4 mm),in response to seasonal changes in predation by YOY yellow perch(Perca flavescens) in a mesotrophic lake. We expected that thelarge-sized D. pulicaria, the most likely subjected to size-selectivepredation by YOY perch, will show stronger antipredator responsesthan the small-sized D. galeata mendotae. To test this hypothesis,we examined changes in life history and behavioural traits injuveniles and adults of both species during four YOY fish predationperiods that were selected based on the presence of YOY perchin the pelagic zone and the relative abundance of Daphnia preyin their gut contents. Our study supports the scenario of negativesize-selective predation by gape-limited YOY perch on both Daphniaspecies. The electivity index indicated that no daphnids witha body length > 1.75 mm were predated by YOY yellow perch.Coexisting Daphnia exhibited phenotypic plasticity in theirantipredator defenses based on their vulnerability to seasonalchanges in size-selective predation of YOY perch. Juvenile Daphniawere the targeted prey and they responded by a decreased bodylength. Behavioural defenses were the dominant strategy usedby both adult Daphnia populations to withstand high predation.A decreased size at maturity was not employed by Daphnia, exceptat the very end of the predation period. Behavioural defensesare short-term strategy adopted to avoid predation. Both antipredatordefenses became unnecessary expenses and were no longer sustainedafter the predation period.     

Modes and mechanisms of a Daphnia invasion

Whether exotic species invade new habitats successfully depends on (i) a change in the invaded habitat that makes it suitable for the invader and (ii) a genetic change in the invading taxon that enhances its fitness in the new habitat, or both. We dissect the causes of invasions of Swiss lakes, north of the Alps, by Daphnia galeata (a zooplankter typical of eutrophic lakes, e.g. those south of the Alps, which are also warmer) by comparing the fitness performance of eight geographically distributed clones that were fed algal-food typical of oligotrophic versus eutrophic conditions at two temperatures. Daphnia longispina, native to oligotrophic Swiss lakes, served as a reference. Daphnia galeata requires eutrophic food to persist, whereas D. longispina survives and grows on oligotrophic food but does even better on eutrophic food. Invasion by D. galeata is further explained because invading clones from the north perform better on eutrophic food and at cooler temperatures than native clones from the south, suggesting a local response to countergradient selection. Our data support the hypothesis that populations of the invader in northern lakes are dominated by well-adapted genotypes. Our results illustrate how environmental change (i.e. eutrophication) and local adaptation can act together to drive a successful invasion.     

A taxonomic reappraisal of the Daphnia hyalina complex (Crustacea: Cladocera): an experimental and ecological approach

The paper is a taxonomic study of the limnetic Daphnia populations of the English Lake District, the populations upon which the most popular taxonomic scheme is based (Johnson, 1952).
Johnson treats these populations as three varieties of the species D. hyalina, namely D. hyalina s. str., D.h. var. galeata and D.h. var. lacustris .
Careful examination of the populations of Esthwaite reveals that what has previously been regarded as a population of a single variety of D. hyalina. includes three distinct forms. Experimental work confirms that these are genetically distinct. One of these forms corresponds to D. hyalina var. galeata , whilst the other two are designated the "toothed" and "round-headed" forms.
The first part of the paper is concerned with the taxonomic status of var. galeata . Brooks (1957) and some later workers treat this as a separate species (D. galeata) on morphological grounds. This is fully supported by the present work. Differences between var. galeata and co-existing forms in incidence of sexual reproduction, in vertical distribution and in seasonal abundance and average brood-size provide further support for the separation of D. galeata.
The second part of the paper is concerned with D. hyalina s.str. which, on the basis of morphology, ecology and reproductive behaviour, is shown to be at least as distinct from other members of the complex as is D. galeata . It is argued that other forms should therefore to be removed from D. hyalina . The taxonomic status and affinities of the remaining "toothed" and "round-headed" forms are discussed.     

On the cost of vertical migration: are feeding conditions really worse at greater depths?

SUMMARY 1. The ultimate explanation for diel vertical migration (DVM) of zooplankton is the avoidance of visual predation in surface waters. Studies on migrating zooplankton have shown that remaining in the cold and food-poor hypolimnion during the day, however, has demographic costs. Higher temperatures and greater food concentrations in the surface waters are thought to be the main reasons why Daphnia species move upwards at night.
2. In this study, we investigated the growth condition of daphniids raised on seston taken from different depths from a lake with and without a deep-water chlorophyll maximum.
3. Juvenile growth rates of Daphnia galeata x hyalina from the lake without a deep-water chlorophyll maximum were similar for all treatments. After temperature correction, however, growth rates were significantly higher on seston taken from the surface layers.
4. In contrast, in the lake with the deep-water chlorophyll maximum, D. galeata growth rates were higher in deeper strata, even after temperature correction. Although this lake had a weak temperature gradient, D. galeata left the food-rich strata at night and migrated into the surface food-poor environment. Invertebrate predation and oxygen depletion are probably not the reasons for the nocturnal upward migration into the surface strata. Therefore, we assume that D. galeata migrates upwards to take advantage of higher temperatures. Using several temperature–egg-development models, we could not, however, fully explain this behaviour.     

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.     

A spontaneous mutant of microcystin biosynthesis: genetic characterization and effect on Daphnia

Microcystis aeruginosa strain MRC is unique in its' possession of the mcyA-J gene cluster, which encodes microcystin synthetase, but its' inability to produce microcystins. M. aeruginosa strain MRD is genetically identical to MRC at numerous genomic loci examined, but produces a variety of microcystins, mainly with the amino acid tyrosine in the molecule. Zooplankton studies with Daphnia galeata and D. pulicaria , using the mutant (MRC) and its' wild type (MRD), showed for the first time that microcystins other than microcystin-LR can be responsible for the poisoning of Daphnia by Microcystis . Regardless of microcystin content, both Daphnia exhibited significantly reduced ingestion rates when fed with either strain of M. aeruginosa compared with the green alga Scenedesmus acutus . A disruption of the molting process in both Daphnia spp. was noted when these species were fed with MRC cells. Such symptoms on Daphnia have not been previously reported for cyanobacteria and may point to a bioactive compound, other than microcystin, which inhibits the hardening of protein–chitin complexes in Daphnia .     

Discrimination of hybrid classes using cross-species amplification of microsatellite loci: methodological challenges and solutions in Daphnia

Microsatellite markers are important tools in population, conservation and forensic studies and are frequently used for species delineation, the detection of hybridization and introgression. Therefore, marker sets that amplify variable DNA regions in two species are required; however, cross-species amplification is often difficult, as genotyping errors such as null alleles may occur. To estimate the level of potential misidentifications based on genotyping errors, we compared the occurrence of parental alleles in laboratory and natural Daphnia hybrids (Daphnia longispina group). We tested a set of 12 microsatellite loci with regard to their suitability for unambiguous species and hybrid class identification using F(1) hybrids bred in the laboratory. Further, a large set of 44 natural populations of D. cucullata, D. galeata and D. longispina (1715 individuals) as well as their interspecific hybrids were genotyped to validate the discriminatory power of different marker combinations. Species delineation using microsatellite multilocus genotypes produced reliable results for all three studied species using assignment tests. Daphnia galeata × cucullata hybrid detection was limited due to three loci exhibiting D. cucullata-specific null alleles, which most likely are caused by differences in primer-binding sites of parental species. Overall, discriminatory power in hybrid detection was improved when a subset of markers was identified that amplifies equally well in both species.     

Spatial distribution of the Daphnia longispina species complex and other planktonic crustaceans in the heterogeneous environment of canyon-shaped reservoirs

Canyon-shaped reservoirs are often characterised by longitudinalgradients of environmental factors (including trophic level,phytoplankton and zooplankton biomass and abundance of planktivorousfish) affecting the taxonomic composition of the pelagic community.We tested the hypothesis that the spatial distribution of differentspecies and interspecific hybrids of the Daphnia longispinaspecies complex is non-random under such conditions. Duringthe summer stratification, we sampled crustacean zooplanktonfrom 11 reservoirs, covering both longitudinal (upstream, middle,dam) and vertical (epi-, meta- and hypolimnion) environmentalgradients. Allozyme electrophoresis was used to discriminateamong different Daphnia taxa. All three frequently hybridizingEuropean species of the complex (galeata, cucullata, longispina= hyalina) and hybrids with Daphnia galeata were commonly recorded.Smaller-bodied Daphnia cucullata and its hybrids, when present,preferred mostly the nutrient- and food-rich upstream regions;D. longispina and its hybrids were more commonly found in thedownstream part, and often dominated in the meta- or hypolimnion.Redundancy analyses confirmed significant differences in theDaphnia taxon composition, as well as in spatial distributionof other crustacean species, along both gradients. For the firsttime, we demonstrate regular patterns in the horizontal distributionof Daphnia species and hybrids within a water body, thus acceptingour hypothesis. Such spatial distributional patterns may stronglyimpact local hybridization processes.     

Quaternary diversification in a sexual Holarctic zooplankter, Daphnia galeata

The effects of Quaternary glacial range partitioning on the diversification of Holarctic biota remain unclear. Glacial refugial lineages may form vicariant species, hybrid products, or merge after secondary contact. Here, we assess the effects of Quaternary glaciation on a Holarctic sexual zooplankter, Daphnia galeata, with apparently marked dispersal capacity and a widespread hybrid lineage in the New World. We collected samples of this species from 148 Holarctic lakes, analysed the nuclear and mitochondrial gene sequences, and tested predictions for hypotheses that account for the origin and spread of the New World D. galeata. We detected five nuclear phylogroups and four mitochondrial phylogroups, most of which were restricted to either the New World or the Old World. The oldest mitochondrial phylogroup was restricted to Japan. One major mitochondrial clade was distributed throughout the Holarctic, but only four haplotypes were shared among continents, and analysis of molecular variance indicated significant structure at the continental level. Haplotype sharing among continents could largely be attributed to anthropogenic introductions. Mismatch distributions, haplotype networks, phylogenetic trees, longitudinal haplotype diversity erosion and coalescence analyses are consistent with colonization from an Old World and a New World refugium. Our nuclear and mitochondrial DNA sequence evidence supports the hypothesis that the New World D. galeata underwent introgression with Daphnia dentifera, with dispersal being enhanced by glaciation. We conclude that Quaternary glaciation had a pronounced effect on the diversification of a Holarctic sexual zooplankter.     

Length–weight relationships of eight freshwater planktonic crustacean species in Japan

1. The relationships between body length and dry weight were examined for eight species of freshwater planktonic crustacea common in Japan: Eodiaptomus japonicus, Acanthodiaptomus pacificus, Daphnia galeata, D. similis, D. magna, Scapholeberis mucronata, Simocephalus exspinosus and Bosmina longirostris.
2. The relationships of two diaptomid species were similar and the approximate equation was ln W = 2.7 + 2.6*ln L , where L is prosome length (mm) and W is dry weight (μg). The carapace length–dry weight relationships in branchiopods were more variable, with the slope ranging from 1.9 to 2.9 and the intercept from 2.0 to 3.7.
3. The effects of food conditions on the relationship were examined in the laboratory, and the seasonal changes in the field were also studied.
4. Practical advice is presented for predicting crustacean weight from body length.     

南漪湖沉积物枝角类卵鞍密度的垂直变化及其与营养盐的关系

本文研究了南漪湖沉积物(1～28 cm)中枝角类卵鞍密度的垂直变化.共鉴定出中华拟同形溞、盔形溞、蚤状溞、网纹溞、微型裸腹溞、象鼻溞、船卵溞、粗刺大尾溞和盘肠溞9种枝角类的卵鞍.大型溞属种类和小型枝角类卵鞍密度的变化范围分别为0～2.0和0～10.0 ind·g^-1 DM.中华拟同形溞和网纹溞分别是南漪湖沉积物大型枝角类和小型枝角类的优势种.中华拟同形溞的卵鞍密度与1^#采样点沉积物中总磷含量呈显著负相关,与3^#采样点沉积物中总氮、总磷含量均呈显著正相关.在3个采样点中(除2^#采样点总氮外),网纹溞的卵鞍密度与沉积物中总氮、总磷含量均呈显著正相关.结果暗示,湖泊富营养化和鱼类捕食影响了南漪湖枝角类群落结构的历史演变.     

Effects of light on size-selectivity of kokanee feeding on Daphnia

Planktivory by fingerling kokanee Oncorhynchus nerka feeding on Daphnia became more size-selective with increased illumination (0·1 v . 15 lux). The average difference in length of Daphnia ( D. pulex and D. galeata ) in trial arenas before and after kokanee feeding was 0·107 mm ( P  = 0·0007) at 0·1 lux and 0·371 mm ( P  < 0·0001) at 15 lux.