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468 results for “Daphnia”
FIGURE 4 in A new species of the Daphnia sinevi group (Crustacea: Cladocera: Daphniidae) from Sakhalin Island, Russian Far East
FIGURE 4. Daphnia sakhaliensis sp.nov., parthenogenetic female from a puddle near River Tim', near Voskresenovka, Sakhalin Area, Russia, NMK 2432. A, adult, lateral view. B, its antero-dorsal view. C, head, lateral view. D–E, postabdominal claws with different morphology. F, armature of inner face of posterior valve margin. G, exopodite III. Scale bars denote 0.1 mm for A–E, I, and 0.01 mm for F–H.
Data from: Urbanization drives genetic differentiation in physiology and structures the evolution of pace-of-life syndromes in the water flea Daphnia magna
Natural and human-induced stressors elicit changes in energy metabolism and stress physiology in populations of a wide array of species. Cities are stressful environments that may lead to differential selection on stress-coping mechanisms. Given that city ponds are exposed to the urban heat island effect and receive polluted run-off, organisms inhabiting these ecosystems might show genetic differentiation for physiological traits enabling them to better cope with higher overall stress levels. A common garden study with 62 Daphnia magna genotypes from replicated urban and rural populations revealed that urban Daphnia have significantly higher concentrations of total body fat, proteins, and sugars. Baseline activity levels of the antioxidant defense enzymes superoxide dismutase (SOD) and glutathione-S-transferase (GST) were higher in rural compared to city populations, yet urban animals were equally well protected against lipid peroxidation. Our results add to the recent evidence of urbanisation-driven changes in stress physiology and energy metabolism in terrestrial organisms. Combining our results with data on urban life history evolution in Daphnia revealed that urban genotypes show a structured pace-of-life syndrome involving both life history and physiological traits, whereas this is absent in rural populations.
Data from: Variation in life-history traits among Daphnia and its relationship to species-level responses to phosphorus limitation
Currently organisms are experiencing changes in their environment at an unprecedented rate. Therefore the study of the contributions to and responses in traits linked to fitness is crucial, as they have direct consequences on a population's success in persisting under such change. Daphnia is used as a model organism as the genus contains keystone primary consumers in aquatic food webs. A life-history table experiment (LHTE) using four species of Daphnia was conducted to compare variation in life-history traits among species across two different environmental conditions (high and low phosphorous availability). Results indicate that the food quality environment had the most impact on life-history traits, while genetic contributions to traits were higher at the species-level than clonal-level. Higher trait variation and species-level responses to P-limitation was more evident in reproductive traits, while growth traits were found to be less affected by food quality and had less variation. Exploring trait variation and potential plasticity in organisms is increasingly important to consider as a potential mechanism for population persistence given the fluctuations in environmental stressors we are currently experiencing.
Data from: Contrasting patterns of divergence at the regulatory and sequence level in European Daphnia galeata natural populations
Understanding the genetic basis of local adaptation has long been the focus of evolutionary biology. Recently there has been increased interest in deciphering the evolutionary role of Daphnia's plasticity and the molecular mechanisms of local adaptation. Using transcriptome data, we assessed the differences in gene expression profiles and sequences within and between four European Daphnia galeata populations. To distinguish neutral from adaptive differentiation, we corrected for phylogenetic differentiation of Daphnia populations. We also applied a "transcriptome scan" approach to investigate the role of natural selection in shaping divergent expression profiles among populations. Furthermore, a SNP analysis allowed inferring population structure and the distribution of genetic variation. Using sequence information, the transcripts were annotated using a comparative genomics approach. In total, ~33% of 32903 transcripts were differentially expressed between populations. Among 10280 differentially expressed transcripts, 5209 transcripts deviated from neutral expectations and were likely involved in local adaptation. The population divergence at the sequence level was higher than at the gene expression level by several orders of magnitude and revealed very distinct clusters according to population origin. Our analysis revealed the respective roles of genetic drift and selection in the four Daphnia populations. This study is a first attempt to understand the genetic background of adaptation to environmental changes in a key species of aquatic ecosystems in absence of any laboratory induced stressor.
Data from: Does local adaptation to resources explain genetic differentiation among Daphnia populations?
Substantial genetic differentiation is frequently observed among populations of cyclically parthenogenetic zooplankton despite their high dispersal capabilities and potential for gene flow. Local adaptation has been invoked to explain population genetic differentiation despite high dispersal, but several neutral models that account for basic life history features also predict high genetic differentiation. Here, we study genetic differentiation among four populations of Daphnia pulex in east central Illinois. As with other studies of Daphnia, we demonstrate substantial population genetic differentiation despite close geographic proximity (< 50 km; mean θ = 0.22). However, we explicitly tested, and failed to find evidence for, the hypothesis that local adaptation to food resources in these populations. Recognizing that local adaptation can occur in traits unrelated to resources, we estimated contemporary migration rates (m) and tested for admixture to evaluate the hypothesis that observed genetic differentiation is consistent with local adaptation to other untested ecological factors. Using Bayesian assignment methods, we detected migrants in three of the four study populations including substantial evidence for successful reproduction by immigrants in one pond, allowing us to reject the hypothesis that local adaptation limits gene flow for at least this population. Thus, we suggest that local adaptation does not explain genetic differentiation among these Daphnia populations, and that other factors related to extinction / colonization dynamics, a long approach to equilibrium FST, or substantial genetic drift due to a low number of individuals hatching from the egg bank each season may explain genetic differentiation.
Data from: Divergent clades or cryptic species? Mito-nuclear discordance in a Daphnia species complex
Background: Genetically divergent cryptic species are frequently detected by molecular methods. These discoveries are often a byproduct of molecular barcoding studies in which fragments of a selected marker are used for species identification. Highly divergent mitochondrial lineages and putative cryptic species are even detected in intensively studied animal taxa, such as the crustacean genus Daphnia. Recently, eleven such lineages, exhibiting genetic distances comparable to levels observed among well-defined species, were recorded in the D. longispina species complex, a group that contains several key taxa of freshwater ecosystems. We tested if three of those lineages represent indeed distinct species, by analyzing patterns of variation of ten nuclear microsatellite markers in six populations. Results: We observed a discordant pattern between mitochondrial and nuclear DNA, as all individuals carrying one of the divergent mitochondrial lineages grouped at the nuclear level with widespread, well-recognized species coexisting at the same localities (Daphnia galeata, D. longispina, and D. cucullata). Conclusions: A likely explanation for this pattern is the introgression of the mitochondrial genome of undescribed taxa into the common species, either in the distant past or after long-distance dispersal. The occurrence of highly divergent but rare mtDNA lineages in the gene pool of widespread species would suggest that hybridization and introgression in the D. longispina species complex is frequent even across large phylogenetic distances, and that discoveries of such distinct clades must be interpreted with caution. However, maintenance of ancient polymorphisms through selection is another plausible alternative that may cause the observed discordance and cannot be entirely excluded.
Data from: Thermal evolution offsets the elevated toxicity of a contaminant under warming: a resurrection study in Daphnia magna
Synergistic interactions between temperature and contaminants are a major challenge for ecological risk assessment, especially under global warming. While thermal evolution may increase the ability to deal with warming, it is unknown whether it may also affect the ability to deal with the many contaminants that are more toxic at higher temperatures. We investigated how evolution of genetic adaptation to warming affected the interactions between warming and a novel stressor: zinc oxide nanoparticles (nZnO) in a natural population of Daphnia magna using resurrection ecology. We hatched resting eggs from two D. magna subpopulations (old: 1955-1965, recent: 1995-2005) from the sediment of a lake that experienced an increase in average temperature and in recurrence of heat waves but was never exposed to industrial waste. In the old 'ancestral' subpopulation, exposure to a sublethal concentration of nZnO decreased the intrinsic growth rate, metabolic activity and energy reserves at 24°C but not at 20°C, indicating a synergism between warming and nZnO. In contrast, these synergistic effects disappeared in the recent 'derived' subpopulation that evolved a lower sensitivity to nZnO at 24°C, which indicates that thermal evolution could offset the elevated toxicity of nZnO under warming. This evolution of reduced sensitivity to nZnO under warming could not be explained by changes in the total internal zinc accumulation but was partially associated with the evolution of the expression of a key metal detoxification gene under warming. Our results suggest that the increased sensitivity to the sublethal concentration of nZnO under the predicted 4°C warming by the end of this century may be counteracted by thermal evolution in this D. magna population. Our results illustrate the importance of evolution to warming in shaping the responses to another anthropogenic stressor, here a contaminant. More general, genetic adaptation to an environmental stressor may ensure that synergistic effects between contaminants and this environmental stressor will not be present anymore.
Data from: Parental and hybrid Daphnia from the D. longispina complex: long-term dynamics in genetic structure and significance of overwintering modes
In recent decades, hybridization has become a focus of attention because of its role in evolutionary processes. However, little is known about changes in genetic structure within and between parental species and hybrids over time. Here, we studied processes of genetic change in parental species and hybrids from the Daphnia longispina complex (Crustacea, Cladocera) over a period of six years across ten habitats. These cyclical parthenogens respond to fluctuating environments by switching from asexual to sexual reproduction. Importantly, sexually produced diapausing eggs, which resist extreme conditions such as low temperatures and serve as dispersal stages, are produced to a lower extent by hybrids. Long-term microsatellite data revealed clear differences between hybrids and parental species. In hybrids, clonal diversity values were lower, whereas heterozygosity and linkage disequilibrium values were higher compared to parental species. Clonal diversity of hybrids responded to the strength of the winter, with cold winters resulting in few genotypes in the following spring. In time windows when only asexual hybrid females survive, priority effects will favour the establishment of the hybrid offspring before hatchlings from parental diapause eggs can enter the community. The constant high levels of heterozygosity maintained by clonal reproduction in hybrids might lead to their successful establishment over time, when they are able to escape competition from both parental species. Although we found evidence that hybrids diversity depends on fluctuating environments, a direct link between hybrid abundance and the strength of winter was missing. Because of reduced adaptability in clonally reproducing hybrids, multiple factors must contribute to promoting their long-term success in fluctuating environments.
Data from: Variation in chronic radiation exposure does not drive life history divergence among Daphnia populations across the Chernobyl Exclusion Zone
Ionising radiation is a mutagen with known negative impacts on individual fitness. However, much less is known about how these individual fitness effects translate into population-level variation in natural environments that have experienced varying levels of radiation exposure. In this study, we sampled genotypes of the freshwater crustacean, Daphnia pulex, from the eight inhabited lakes across the Chernobyl Exclusion Zone (CEZ). Each lake has experienced very different levels of chronic radiation exposure since a nuclear power reactor exploded there over thirty years ago. The sampled Daphnia genotypes represent genetic snapshots of current populations and allowed us to examine fitness-related traits under controlled laboratory conditions at UK background dose rates. We found that whilst there was variation in survival and schedules of reproduction among populations, there was no compelling evidence that this was driven by variation in exposure to radiation. Previous studies have shown that controlled exposure to radiation at dose rates included in the range measured in the current study reduce survival, or fecundity, or both. One limitation of this study is the lack of available sites at high dose rates, and future work could test life history variation in various organisms at other high radiation areas. Our results are nevertheless consistent with the idea that other ecological factors, e.g., competition, predation or parasitism, are likely to play a much bigger role in driving variation among populations than exposure to the high radiation dose rates found in the CEZ. These findings clearly demonstrate that it is important to examine the potential negative effects of radiation across wild populations that are subject to many and varied selection pressures as a result of complex ecological interactions.
Data from: Initial genetic diversity enhances population establishment and alters genetic structuring of a newly established Daphnia metapopulation
When newly created habitats are initially colonized by genotypes with rapid population growth rates, later arriving colonists may be prevented from establishing. Although these priority effects have been documented in multiple systems, their duration may be influenced by the diversity of the founding population. We conducted a large-scale field manipulation to investigate how initial clonal diversity influences temporal and landscape patterns of genetic structure in a developing metapopulation. Six genotypes of obligately asexual Daphnia pulex were stocked alone (no clonal diversity) or in combination ('high' clonal diversity) into newly created experimental woodland ponds. We also measured the population growth rate of all clones in the laboratory when raised on higher-quality and lower-quality resources. Our predictions were that in the 3 years following stocking, clonally diverse populations would be more likely to persist than nonclonally diverse populations and exhibit evidence for persistent founder effects. We expected that faster growing clones would be found in more pools and comprise a greater proportion of individuals genotyped from the landscape. Genetic composition, both locally and regionally, changed significantly following stocking. Six of 27 populations exhibited evidence for persistent founder effects, and populations stocked with 'high' clonal diversity were more likely to exhibit these effects than nonclonally diverse populations. Performance in the laboratory was not predictive of clonal persistence or overall dominance in the field. Hence, we conclude that although laboratory estimates of fitness did not fully explain metapopulation genetic structure, initial clonal diversity did enhance D. pulex population establishment and persistence in this system.
Data from: Genes mirror geography in Daphnia magna
Identifying the presence and magnitude of population genetic structure remains a major consideration in evolutionary biology as doing so allows one to understand the demographic history of a species as well as make predictions of how the evolutionary process will proceed. Next-generation sequencing methods allow us to reconsider previous ideas and conclusions concerning the distribution of genetic variation, and what this distribution implies about a given species evolutionary history. A previous phylogeographic study of the crustacean Daphnia magna suggested that, despite strong genetic differentiation among populations at a local scale, the species shows only moderate genetic structure across its European range, with a spatially patchy occurrence of individual lineages. We apply RAD sequencing to a sample of D. magna collected across a wide swath of the species' Eurasian range and analyse the data using principle component analysis (PCA) of genetic variation and Procrustes analytical approaches, to quantify spatial genetic structure. We find remarkable consistency between the first two PCA axes and the geographic coordinates of individual sampling points, suggesting that, on a continent-wide scale, genetic differentiation is driven to a large extent by geographic distance. The observed pattern is consistent with unimpeded (i.e. no barriers, landscape or otherwise) migration at large spatial scales, despite the fragmented and patchy nature of favourable habitats at local scales. With high-resolution genetic data similar patterns may be uncovered for other species with wide geographic distributions, allowing an increased understanding of how genetic drift and selection have shaped their evolutionary history.
Data from: Maladaptation to acute metal exposure in resurrected Daphnia ambigua clones after decades of increasing contamination
Human environmental impacts have driven some of the strongest and fastest phenotypic changes recorded in wild animal populations. Across populations, this variation is often adaptive, as populations evolve fitness advantages in response to human-modified environments. Yet some populations fail to adapt to changing environments. Evidenced by declines in relative fitness, such seemingly maladaptive outcomes are less common, but may be more likely in human modified contexts. Further, our ability to investigate the dynamics of these adaptive and maladaptive responses over time is typically limited in natural systems. I combined resurrection ecology and paleolimnology approaches to examine evolutionary responses of the freshwater zooplankter Daphnia to exposure to heavy metal contamination over the past 50-75 years using animals hatched from diapausing egg banks. In contrast to the predicted trend of adaptation to metal exposure over time, I observed an increase in sensitivity to both copper and cadmium exposure associated with increasing historic contamination. This potentially maladaptive trend occurred in Daphnia populations in three lakes. Given that the release of toxicants such as heavy metals is widespread and other researchers have observed local maladaptation to toxicant exposure, it is important to understand the drivers and implications of this pattern.
Data from: Temporal genetic stability in natural populations of the waterflea Daphnia magna in response to strong selection pressure
Studies monitoring changes in genetic diversity and composition through time allow a unique understanding of evolutionary dynamics and persistence of natural populations. However, such studies are often limited to species with short generation times that can be propagated in the laboratory or few exceptional cases in the wild. Species that produce dormant stages provide powerful models for the reconstruction of evolutionary dynamics in the natural environment. A remaining open question is to what extent dormant egg banks are an unbiased representation of populations and hence of the species' evolutionary potential, especially in presence of strong environmental selection. We address this key question using the water flea Daphnia magna, which produces dormant stages that accumulate in biological archives over time. We assess temporal genetic stability in three biological archives, previously used in resurrection ecology studies showing adaptive evolutionary responses to rapid environmental change. We show that neutral genetic diversity does not decline with the age of the population and it is maintained in presence of strong selection. In addition, by comparing temporal genetic stability in hatched and unhatched populations from the same biological archive, we show that dormant egg banks can be consulted to obtain a reliable measure of genetic diversity over time, at least in the multi-decadal time frame studied here. The stability of neutral genetic diversity through time is likely mediated by the buffering effect of the resting egg bank.
Data from: Can Daphnia lumholtzi invade European lakes?
The cladoceran Daphnia lumholtzi is a subtropical and tropical zooplankter, and an invasive species in North America. Thus far, D. lumholtzi has not been detected in Europe. Here we investigated whether a hypothetical introduction to Europe could result in a successful invasion, either now or in the near future when facilitated by climate change. In laboratory experiments, we tested whether different clones of D. lumholtzi can invade a resident community consisting of native Daphnia from lake Klostersee, Germany, and how invasion success depends on temperature and the presence or absence of planktivorous fish. In some treatments, invasion success was consistently high, and D. lumholtzi reached densities similar to the native competitors by the end of the experiment. Surprisingly, the presence of a planktivorous fish inhibited the invasion of D. lumholtzi, and a clone with an inducible defense against fish predation was a more successful invader than a permanently defended clone. Of the three temperatures tested in this study (15, 20, and 24°C), invasion success was highest at 20°C. To understand the competitive interaction between native and introduced Daphnia, we fit a Lotka-Volterra-type competition model to the population dynamics. Our experimental and modeling results suggest that D. lumholtzi can invade European lakes and can cause substantial declines in the population size of native Daphnia, with potential consequences for higher trophic levels.
Data from: 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. In order 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 twelve microsatellite loci with regard to their suitability for unambiguous species and hybrid class identification using F1 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 multi-locus genotypes produced reliable results for all three studied species using assignment tests. D. galeata x 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.
Data from: Local human pressures influence gene flow in a hybridizing Daphnia species complex
Anthropogenic environmental changes are considered critical drivers of the genetic structure of populations and communities through, for example, the facilitation of introgressive hybridization between syntopic species. However, the mechanisms by which environmental perturbations trigger changes in the genetic structure of populations and communities, such as the processes that determine the directionality of hybridization and patterns of mitochondrial introgression over many generations, remain largely unexplored. In this study, the changes in genetic structure of hybridizing members of the Daphnia longispina species complex were reconstructed over the last 100 years for three large temperate lakes under strong anthropogenic pressures via paleogenetic analyses of resting egg banks. Drastic changes in the genetic structure of the Daphnia community, associated with hybridization events between D. longispina and D. galeata and subsequent introgression, were detected in Lakes Geneva and Bourget. In Lake Bourget, these changes were induced by the successful establishment of D. galeata with rising phosphorus levels and reinforced by the sensitivity of D. longispina to fish predation pressure. In Lake Geneva, the pattern of hybridization during eutrophication is more likely a function of the original taxonomic composition of the species complex in this lake. Lakes seem to require at least a meso-oligotrophic status to allow D. galeata populations to establish and accordingly no D. galeata genotypes were found in the egg bank of oligotrophic Lake Annecy. In contrast to the generally assumed pattern of unidirectional hybridization in this species complex, bidirectional hybridization was recorded in Lakes Geneva and Bourget. Our results also demonstrate complex genetic trajectories within this species complex and highlight the irreversibility of changes in the genotypic architecture of populations driven by local human pressures. Finally, we show that extensive hybridization and introgression do not necessarily result in a large and homogenous hybrid swarm.
Data from: Population structure of a microparasite infecting Daphnia: spatio-temporal dynamics
Background: Detailed knowledge of spatial and temporal variation in the genetic population structure of hosts and parasites is required for understanding of hostparasite coevolution. As hot-spots of contemporary coevolution in natural systems are difficult to detect and long-term studies are restricted to few systems additional population genetic data from various hostparasite systems may provide important insights into the topic. This is particularly true for parasites as these players have been under-investigated so far due to the lower availability of suitable molecular markers. Here we traced genetic variation (based on sequence variants in the internal transcribed spacer region ITS) among seven geographically isolated populations of the ichthyosporean Caullerya mesnili a common microparasite of the cladoceran Daphnia (here the D. longispina hybrid complex). At some sites we also studied parasite genetic variation over time. Then we tested for associations between parasite genotypes and host species. Results: Parasite (and host) populations were significantly structured across space indicating limited dispersal. Moreover the frequency of parasite genotypes varied significantly over time suggesting rapid evolutionary change in Caullerya. Finally the distribution of parasite genotypes was similar across different host species which might in turn have important consequences for parasite epidemiology. Conclusions: The approach proposed here can be applied to track spatial and temporal changes in the population structure of other microparasite species for which sequence variation in the ITS or other highly variable genome regions has been documented but other types of polymorphic markers are lacking. Screening of parasite sequence variants allows for reliable detection of cross-species infections and using advanced sequencing techniques in the near future for detailed studies of parasite evolution in natural hostparasite systems.
Data from: Evolution of asexual Daphnia pulex in Japan: variations and covariations of the digestive, morphological and life history traits
Background Several genetic lineages of obligate parthenogenetic Daphnia pulex, a common zooplankton species, have invaded Japan from North America. Among these, a lineage named JPN1 is thought to have started colonization from a single genotype several hundred to thousand years ago and subsequently produced many genotypes in Japan. To examine the phenotypic variations due to ecological drivers diverging the genotypes in new habitats, we measured heritability and variation in 17 traits, including life history, morphology and digestive traits, and the genetic distance among the D. pulex JPN1 genotypes in Japan. Results We found that most of the traits measured varied significantly among the genotypes and that heritability was highest in the morphological traits, followed by the digestive and life history traits. In addition, 93% of the variation in these traits was explained by the first three components in the principal component analysis, implying that variations of these heritable traits are not random but rather converged into a few directions. These relations among traits revealed the potential importance of predation pressures and food conditions as factors for diverging and selecting different genotypes. However, the magnitude of the difference in any single trait group did not correlate with the genetic distance. Conclusions Our findings show that the divergent traits evolved within D. pulex JPN1 without genetic recombination since their ancestral clone invaded Japan. Large variations and covariations of the phenotypic traits, irrespective of the genetic distance among the genotypes, support the view that the invasive success of D. pulex JPN1 was promoted by a genetic architecture that allowed for large phenotypic variations with a limited number of functionally important mutations without recombination.
Data from: The role of selection in driving landscape genomic structure of the waterflea Daphnia magna
The combined analysis of neutral and adaptive genetic variation is crucial to reconstruct the processes driving population genetic structure of natural populations. However, such combined analysis is challenging because of the complex interaction among neutral and selective processes in the landscape. Overcoming this level of complexity requires an unbiased search for the evidence of selection in the genomes of populations sampled from their natural habitats and the identification of demographic processes that lead to present-day populations genetic structure. Ecological model species with a suite of genomic tools and well-understood ecologies are best suited to resolve this complexity and elucidate the role of selective and demographic processes in the landscape genomic structure of natural populations. Here we investigate the water flea Daphnia magna, an emerging model system in genomics and a renowned ecological model system. We infer past and recent demographic processes by contrasting patterns of local and regional neutral genetic diversity at markers with different mutation rates. We assess the role of the environment in driving genetic variation in our study system by identifying correlates between biotic and abiotic variables naturally occurring in the landscape and patterns of neutral and adaptive genetic variation. Our results indicate that selection plays a major role in determining the population genomic structure of D. magna. First, environmental selection directly impacts genetic variation at loci hitchhiking with genes under selection. Secondly, priority effects enhanced by local genetic adaptation (cf. monopolization) affect neutral genetic variation by reducing gene flow among populations and genetic diversity within populations.
FIGURE 9 in A new Central European species of the Daphnia curvirostris complex, Daphnia hrbaceki sp. nov. (Cladocera, Anomopoda, Daphniidae)
FIGURE 9. Relationship among species of the Daphnia curvirostris complex and its position relative to other species complexes (represented by selected taxa) of the D. longispina group (nomenclature of the D. longispina complex follows Petrusek et al. 2008). Three members of the D. pulex group, including D. obtusa coexisting with D. hrbaceki, were used as outgroups. The tree was constructed by the Bayesian inference of phylogeny from a partial sequence of the mitochondrial ND2 gene. Node support is provided for Bayesian inference, Maximum Likelihood and Maximum Parsimony analyses, asterisks indicate sister species with support at least 99% in all three analyses. Vertical bars delineate species complexes, scale indicates 10% divergence.
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