Identifying hybridizing taxa within the Daphnia longispina species complex: a comparison of genetic methods and phenotypic approaches

Daphnia galeata Sars, D. longispina O. F. Muller and D. cucullata Sars (Crustacea: Cladocera) are closely related species which often produce interspecific hybrids in natural populations. Several marker systems are available for taxon determination in this hybridizing complex, but their performance and reliability has not been systematically assessed. We compared results from identifications by three molecular methods. More than 1,200 individuals from 10 localities in the Czech Republic were identified as parental species or hybrids by allozyme electrophoresis and the analysis of the restriction fragment length polymorphism of the internal transcribed spacer (ITS-RFLP); over 440 of them were additionally analyzed and identified by 12 microsatellite loci. Identification by microsatellite markers corresponded well with allozyme analyses. However, consistent discrepancies between ITS-RFLP and other markers were observed in two out of 10 studied localities. Although some marker discrepancies may have been caused by occasional recent introgression, consistent deviations between ITS-RFLP and other markers suggest a long-term maintenance of introgressed alleles. These results warn against its use as a sole identification method in field studies. Additionally, we quantitatively evaluated the discriminatory power of geometric morphometric (elliptic Fourier) analysis of body shapes based on photos of over 1,300 individuals pre-classified by allozyme markers. Furthermore, a randomly selected subset of 240 individuals was independently determined from photos by several experts. Despite a tendency for morphological divergence among parental Daphnia species, some taxa (especially D. galeata, D. longispina, and their hybrids) substantially overlapped in their body shapes. This was reflected in different determination success for particular species and hybrids in discriminant analysis based on shape data as well as from photograph

The diaptomid fauna of Israel (Copepoda, Calanoida, Diaptomidae), with notes on the systematics of Arctodiaptomus similis (Baird, 1859) and Arctodiaptomus irregularis Dimentman & Por, 1985 stat. rev

Background: To date, only scarce information is available about the diaptomid copepods of the Middle East despite the ecological and biogeographical importance of the family Diaptomidae in the inland waters of the Holarctic region. Moreover, the taxonomic status of some of the taxa occurring in the area is in need of revision. We studied crustaceans collected from temporary and permanent lentic water bodies in Israel with the aim of providing an updated census of the diaptomid copepods occurring in the country. Furthermore, we morphologically and genetically analysed samples of Arctodiaptomus similis s.l. to shed light on its taxonomy. Results: Five diaptomid taxa were collected during this survey. Among these, Phyllodiaptomus blanci is new for the country and the whole circum-Mediterranean area and might be an allochthonous species of eastern origin. Within the collected samples, we singled out two parapatric groups of populations within A. similis s.l.; these consistently differ both based on morphology (chaetotaxy of the left male antennule) and molecular data (divergence over 17% at the mitochondrial gene for the cytochrome b). We thus attribute the full species rank to Arctodiaptomus irregularis Dimentman & Por, 1985 stat. rev., originally described as a subspecies of the widespread species Arctodiaptomus similis (Baird, 1859). Conclusions: We critically evaluated all species hitherto reported for Israeli inland waters. Considering both the confirmed literature data and the new findings, Israeli diaptomid fauna is composed of at least seven species. However, the need for further surveys in the Middle East and for detailed systematic revisions of some controversial taxa is stressed. Our results on the systematics of A. similis s.l. illustrate the importance of implementing molecular analyses when investigating diversity patterns of groups which are difficult to resolve based on morphology alone

Apparent interspecific transmission of Aphanomyces astaci from invasive signal to virile crayfish in a sympatric wild population

The crayfish plague pathogen (Aphanomyces astaci) causes mass mortalities of European crayfish when transmitted from its original North American crayfish hosts. Little is known, however, about interspecific transmission of the pathogen between different American crayfish species, although evidence from trade of ornamental crayfish suggests this may happen in captivity. We screened signal and virile crayfish for A. astaci at allopatric and sympatric sites in a UK river. Whilst the pathogen was detected in signal crayfish from both sites, infected virile crayfish were only found in sympatry. Genotyping of A. astaci from virile crayfish suggested the presence of a strain related to one infecting British signal crayfish. We conclude that virile crayfish likely contracted A. astaci interspecifically from infected signal crayfish. Interspecific transmission of A. astaci strains differing in virulence between American carrier species may influence the spread of this pathogen in open waters with potential exacerbated effects on native European crayfish

Spatial insurance against a heatwave differs between trophic levels in experimental aquatic communities

Climate change-related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better-adapted taxa. Yet, experimental evidence for such predictions from multi-trophic communities and pulse-type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat-induced increase in metabolic costs, resulting in weaker top-down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer-lasting changes in ecosystem functioning

Strong differences in the clonal variation of two Daphnia species from mountain lakes affected by overwintering strategy

<p>Abstract</p> <p>Background</p> <p>The population structure of cyclical parthenogens such as water fleas is strongly influenced by the frequency of alternations between sexual and asexual (parthenogenetic) reproduction, which may differ among populations and species. We studied genetic variation within six populations of two closely related species of water fleas of the genus <it>Daphnia </it>(Crustacea, Cladocera). <it>D. galeata </it>and <it>D. longispina </it>both occur in lakes in the Tatra Mountains (Central Europe), but their populations show distinct life history strategies in that region. In three studied lakes inhabited by <it>D. galeata</it>, daphnids overwinter under the ice as adult females. In contrast, in lakes inhabited by <it>D. longispina</it>, populations apparently disappear from the water column and overwinter as dormant eggs in lake sediments. We investigated to what extent these different strategies lead to differences in the clonal composition of late summer populations.</p> <p>Results</p> <p>Analysis of genetic variation at nine microsatellite loci revealed that clonal richness (expressed as the proportion of different multilocus genotypes, MLGs, in the whole analysed sample) consistently differed between the two studied species. In the three <it>D. longispina </it>populations, very high clonal richness was found (MLG/N ranging from 0.97 to 1.00), whereas in <it>D. galeata </it>it was much lower (0.05 to 0.50). The dominant MLGs in all <it>D. galeata </it>populations were heterozygous at five or more loci, suggesting that such individuals all represented the same clonal lineages rather than insufficiently resolved groups of different clones.</p> <p>Conclusions</p> <p>The low clonal diversities and significant deviations from Hardy-Weinberg equilibrium in <it>D. galeata </it>populations were likely a consequence of strong clonal erosion over extended periods of time (several years or even decades) and the limited influence of sexual reproduction. Our data reveal that populations of closely related <it>Daphnia </it>species living in relatively similar habitats (permanent, oligotrophic mountain lakes) within the same region may show strikingly different genetic structures, which most likely depend on their reproductive strategy during unfavourable periods. We assume that similar impacts of life history on population structures are also relevant for other cyclical parthenogen groups. In extreme cases, prolonged clonal erosion may result in the dominance of a single clone within a population, which might limit its microevolutionary potential if selection pressures suddenly change.</p

The long-term consequences of hybridization between the two Daphnia species, D. galeata and D. dentifera, in mature habitats

<p>Abstract</p> <p>Background</p> <p>Ecological specializations such as antipredator defense can reinforce morphological and distributional divergence within hybridizing species. Two hybridizing species of <it>Daphnia </it>(<it>D. galeata </it>and <it>D. dentifera</it>) are distributed in both Japan and North America; however, these populations have a longer history in Japan than in North America due to the differing impact of the last glaciation on these two regions. We tested the hypothesis that this longer coexistence in Japan would lead to extensive genetic admixture in nuclear and mitochondrial DNA whilst the distinct morphological traits and distributional patterns would be maintained.</p> <p>Results</p> <p>The high level of correspondence among morphological traits, distribution, and mitochondrial and nuclear DNA types for the specimens with <it>D. dentifera </it>mtDNA indicated that the species distinction has been maintained. However, a discordance between mtDNA and nuclear ITS-1 types was observed for most specimens that had <it>D. galeata </it>mtDNA, consistent with the pattern seen between the two species in North America. This observation suggests nuclear introgression from <it>D. dentifera </it>into <it>D. galeata </it>without mitochondrial introgression.</p> <p>Conclusions</p> <p>The separation of morphological traits and distribution ranges of the two hybridizing species in Japan, as well as in North America, has been maintained, despite large differences in climatic and geographical histories of these two regions. Variations in environmental factors, such as predation pressure, might affect maintenance of the distribution, although the further studies are needed to confirm this.</p

Inducible Defenses with a "Twist": Daphnia barbata Abandons Bilateral Symmetry in Response to an Ancient Predator

Predation is one of the most important drivers of natural selection. In consequence a huge variety of anti-predator defenses have evolved in prey species. Under unpredictable and temporally variable predation pressure, the evolution of phenotypically plastic defensive traits is favored. These "inducible defenses", range from changes in behavior, life history, physiology to morphology and can be found in almost all taxa from bacteria to vertebrates. An important group of model organisms in ecological, evolutionary and environmental research, water fleas of the genus Daphnia (Crustacea: Cladocera), are well known for their ability to respond to predators with an enormous variety of inducible morphological defenses. Here we report on the "twist", a body torsion, as a so far unrecognized inducible morphological defense in Daphnia, expressed by Daphnia barbata exposed to the predatory tadpole shrimp Triops cancriformis. This defense is realized by a twisted carapace with the helmet and the tail spine deviating from the body axis into opposing directions, resulting in a complete abolishment of bilateral symmetry. The twisted morphotype should considerably interfere with the feeding apparatus of the predator, contributing to the effectiveness of the array of defensive traits in D. barbata. As such this study does not only describe a completely novel inducible defense in the genus Daphnia but also presents the first report of a free living Bilateria to flexibly respond to predation risk by abandoning bilateral symmetry

The prevalence of Aphanomyces astaci in invasive signal crayfish from the UK and implications for native crayfish conservation

The crayfish plague agent, Aphanomyces astaci, has spread throughout Europe, causing a significant decline in native European crayfish. The introduction and dissemination of this pathogen is attributed to the spread of invasive North American crayfish, which can act as carriers for A. astaci. As native European crayfish often succumb to infection with A. astaci, determining the prevalence of this pathogen in non-native crayfish is vital to prioritize native crayfish populations for managed translocation. In the current study, 23 populations of invasive signal crayfish (Pacifastacus leniusculus) from the UK were tested for A. astaci presence using quantitative PCR. Altogether, 13 out of 23 (56·5%) populations were found to be infected, and pathogen prevalence within infected sites varied from 3 to 80%. Microsatellite pathogen genotyping revealed that at least one UK signal crayfish population was infected with the A. astaci genotype group B, known to include virulent strains. Based on recent crayfish distribution records and the average rate of signal crayfish population dispersal, we identified one native white-clawed crayfish (Austropotamobius pallipes) population predicted to come into contact with infected signal crayfish within 5 years. This population should be considered as a priority for translocation

Mesozoic fossils (>145 Mya) suggest the antiquity of the subgenera of Daphnia and their coevolution with chaoborid predators

<p>Abstract</p> <p>Background</p> <p>The timescale of the origins of <it>Daphnia </it>O. F. Mueller (Crustacea: Cladocera) remains controversial. The origin of the two main subgenera has been associated with the breakup of the supercontinent Pangaea. This vicariance hypothesis is supported by reciprocal monophyly, present day associations with the former Gondwanaland and Laurasia regions, and mitochondrial DNA divergence estimates. However, previous multilocus nuclear DNA sequence divergence estimates at < 10 Million years are inconsistent with the breakup of Pangaea. We examined new and existing cladoceran fossils from a Mesozoic Mongolian site, in hopes of gaining insights into the timescale of the evolution of <it>Daphnia</it>.</p> <p>Results</p> <p>We describe new fossils of ephippia from the Khotont site in Mongolia associated with the Jurassic-Cretaceous boundary (about 145 MYA) that are morphologically similar to several modern genera of the family Daphniidae, including the two major subgenera of <it>Daphnia</it>, i.e., <it>Daphnia </it>s. str. and <it>Ctenodaphnia</it>. The daphniid fossils co-occurred with fossils of the predaceous phantom midge (Chaoboridae).</p> <p>Conclusions</p> <p>Our findings indicate that the main subgenera of <it>Daphnia </it>are likely much older than previously known from fossils (at least 100 MY older) or from nuclear DNA estimates of divergence. The results showing co-occurrence of the main subgenera far from the presumed Laurasia/Gondwanaland dispersal barrier shortly after formation suggests that vicariance from the breakup of Pangaea is an unlikely explanation for the origin of the main subgenera. The fossil impressions also reveal that the coevolution of a dipteran predator (Chaoboridae) with the subgenus <it>Daphnia </it>is much older than previously known -- since the Mesozoic.</p