Size distribution of Daphnia longispina in the vertical profile

D. longispina of the meromictic lake El Tobar is a round-headed form. It never has a helmet, but in summer a small proportion of immature individuals (0.9–1.2 mm females and males) have one or two neck teeth. The size structure of this Daphnia population, as well as the vertical distribution and migration of different size-classes, were studied in September and November of 1991 and April of 1992. The large variation in mean size and size at first reproduction, as well as the occurrence of different patterns of vertical migration are interpreted as responses to different predator situations. At the end of April, when Daphnia mortality by visually hunting predators is dominating, a typical nocturnal migration is adopted and size distribution is biased to smaller size classes. In November, when mortality is mainly attributed to the nocturnally migrating Chaoborus, Daphnia shows a reversed migration pattern. In September, when the population of Daphnia is responding to both visual (fish) and non-visual predators (Chaoborus), it adopts a pattern of twilight migration. The presence of neck teeth in vulnerable size classes in September might be an additional adaptative response to Chaoborus predation. In September, the size structure of the Daphnia population is shifted to larger classes and the vertical distribution of size classes shows a pronounced segregation between juveniles and adults. Juveniles are found closer to the surface, while adults dwell predominantly in the rich, deep waters near the oxicline. This suggests that an additional advantage of the ascent of the adult Daphnia exploiting those deep resources is the release of young in more oxygenated and warmer waters. The Daphnia population of lake El Tobar is known to be clonally diverse, and the changing frequency of genotypes could play an important part in the observed seasonal differences in behaviour and size.     

西藏湖泊长刺溞复合种的系统进化关系

本文通过对西藏湖泊长刺溞复合种(Daphnia longispina complex)中分布最广的3个物种, 即长刺溞(D. longispina)、盔形溞(D. galeata)和颈齿溞(D. dentifera)线粒体COI基因序列以及GenBank中欧洲的长刺溞、加拿大的颈齿溞和我国东部低海拔地区的盔型溞COI基因序列的比较分析, 研究了西藏湖泊长刺溞复合种的系统进化关系, 发现西藏地区的盔型溞、颈齿溞和长刺溞均已出现较大分化。颈齿溞种群内遗传差异度为0.33-2.32%, 盔型溞为0.33-2.74%, 长刺溞的遗传差异度最高, 为1.31-5.50%。基于COI基因序列构建的最大似然树和贝叶斯系统树均表明, 长刺溞复合种由3个进化分支组成, 分别对应长刺溞、盔型溞和颈齿溞, 三者之间的遗传差异度为9.40-16.98%(Kimura 2-parameter双参数模型)。基于COI基因单倍型(haplotype)所构建的网络关系也支持上述3个分支的存在。早期记录虽然显示长刺溞在我国分布较广, 但本次调查只在班公错有发现, 相比之下, 盔形溞和颈齿溞则分布更广。我们的研究表明, 由于形态学鉴定上的局限性, 早期的长刺溞记录很可能混杂了容易引起混淆的盔型溞或颈齿溞。     

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.     

Phenotypic plasticity and developmental noise in hybrid and parental clones of Daphnia longispina complex

Aquatic Ecology - According to the “temporal hybrid superiority hypothesis”, seasonal variability in environmental factors in temperate lakes gives hybrid clones within the D....     

Morphological differentiation within the Daphnia longispina group

Although morphological evolution is assumed to be slow within Daphnia species complexes, discontinuities in morphological space can be detected. Here, morphological data derived from females of genetically-defined clones (cf. Gießler et al., 1999) are presented, in order to estimate the genetic component of phenotypic variance under standardised laboratory conditions. Animals originated from clonal assemblages of pre-alpine lakes and ponds, and a remote lake in western Germany, covering a wide range of morphotypes known from the traditional species D. cucullata, D. galeata, D. hyalina, D. rosea, and a variety of interspecific hybrids. Phenotypic analyses were based on quantitative and qualitative morphological characters of females in the first and fifth instars. Morphological divergence between clones was analysed using discriminant analysis or multidimensional scaling and the significance of the morphological groupings was estimated using neighbour-joining trees and bootstrapping. All analyses confirmed that (a) phenotypic similarities among taxa change with instar, (b) in contrast to low genetic divergence, pronounced morphological divergence exists between animals separated on the lake/pond level favouring speciation by the habitat shift hypothesis.     

Analysis of reticulate relationships within the Daphnia longispina species complex. Allozyme phenotype and morphology

A two-step method is proposed to get reliable associations between morphology and genotype in clonal assemblages in which more than two predominantly parthenogenetic species are thought to coexist with hybrids. In dataset 1, the genetic relationships among clones of the Daphnia longispina hybrid complex from seven prealpine lakes in southern Germany were studied based on the variation at 21 enzyme loci. The spatial arrangement in the multidimensional scaling plot revealed a reticulate pattern among three presumably parental species, D. cucullata, D. galeata and D. hyalina, and three hybrid groups, D. cucullata/galeata, D. cucullata/hyalina and D. galeata/hyalina. The Got1 locus was believed to be a discriminating factor between species and hybrids (cf. Wolf and Mort, 1986). However, this locus is more variable, and 57% of the clones would have been misidentified using it. Moreover, the morphological variation within genetically defined groups is also higher than previously assumed. In dataset 2, the revision of morphological and genetic markers greatly improved the association between morphology and genotype in newly collected animals. The spatial arrangement of clones in multidimensional scaling plots and morphological asymmetries to parents suggest both, different degrees of introgression and bidirectional hybridization. Most unexpected genotypes were found in the cxh hybrid group, suggesting that F1-hybrids are fertile. The results showed (1) that the clonal diversity was very high (2) that detailed analyses of multiple morphological and allozyme markers are necessary to resolve taxonomic relationships within clonal assemblages consisting of multiple species, hybrids and differently introgressed backcrosses, and (3) that the three original species seem to have sufficient within-species recombination and a low enough rate of backcrossing to allow taxonomic identification. It must remain undecided if the present situation is locally restricted, if it is stable or represents a transient situation which could lead to either a consolidation of the three species by gradual elimination of the hybrids, to a taxonomic breakdown, or to hybrid speciation.     

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.     

Natural co-infection of Babesia caballi and Encephalitozoon-like microsporidia in the tick Anocentor nitens (Acari: Ixodidae)

The present paper reports the occurrence of natural co-infection of Babesia caballi and Encephalitozoon-like microsporidia in the tick Anocentor nitens. Engorged females of ticks, collected from a naturally B. caballi-infected horse, were incubated at 27 degrees C and relative humidity over 83%. After a 6-day incubation period, Giemsa-stained smears prepared from hemolymph were examined microscopically under oil immersion. B. caballi infected ticks were dissected and samples of midgut tissue were examined by transmission electron microscopy, through which free sporokinetes were seen in the cytoplasm of gut epithelial cells. In addition, Encephalitozoon-like microsporidia were observed inside the parasitophorous vacuoles in the same cell in which sporokinetes of B. caballi were found and also in some neighbour cells. They presented different morphological stages, suggesting a sequential phases of development.     

A taxonomic reappraisal of the European Daphnia longispina complex (Crustacea, Cladocera, Anomopoda)

The Daphnia longispina complex contains some of the most common water flea species in the northern hemisphere, and has been a model organism for many ecological and evolutionary studies. Nevertheless, the systematics and nomenclature of this group, in particular its Palaearctic members, have been in flux for the past 150 years; this hinders the correct interpretation of scientific results and promotes the erroneous use of species names. We revise the systematics of this species complex based on mitochondrial sequence variation (12S rDNA and COI) of representative populations across Europe, with a special focus on samples from type localities of the respective taxa. Combining genetic evidence and morphological assignments of analysed individuals, we propose a comprehensive revision of the European members of the D. longispina complex. We show that D. hyalina and D. rosea morphotypes have evolved several times independently, and we find no evidence to maintain these morphotypes as distinct biological species. Alpine individuals described as D. zschokkei are conspecific with the above-mentioned lineage. We suggest that this morphologically and ecologically plastic but genetically uniform hyalina–rosea–zschokkei clade should be identified as D. longispina (O. F. Müller, 1776). The valid name of Fennoscandian individuals labelled D. longispina sensu stricto in the recent literature is D. lacustris G. O. Sars, 1862. Additionally, we discovered another divergent lineage of this group, likely an undescribed species, in southern Norway. Our results present a solution for several prevailing taxonomic problems in the genus Daphnia , and have broad implications for interpretation of biogeographical patterns, and ecological and evolutionary studies.     

Intra-specific rDNA-ITS restriction site variation and an improved protocol to distinguish species and hybrids in the Daphnia longispina complex

A collaborative research effort was undertaken to evaluate the robustness of a recently developed genetic tool for species identification of members in the morphologically variable Daphnia longispina species complex. This genetic method, based on restriction fragment length polymorphism (RFLP) of the internal transcribed spacer region (ITS) of nuclear ribosomal DNA (rDNA) with restriction enzymes Mwo I and Sau96 I [Billiones et al., 2004. Hydrobiologia 526: 43–53], was applied to many different European populations. Results were compared with two or more independently obtained characters (morphology, allozymes, mitochondrial DNA (mtDNA), or cloned rDNA-ITS sequences). Individuals of most taxa were readily identified, but unexpected ITS-RFLP patterns were found in many individuals indicated by other markers to be D. galeata or one of its hybrids. Among 43 investigated D. galeata populations (902 specimen analysed by ITS-RFLP), deviant RFLP fragment patterns occurred in 26 (i.e., more than half) of the populations. The deviant patterns could be attributed to the loss of one single restriction site in the ITS2 region. This loss made the distinction of D. galeata from other species unreliable, and F1 hybrids could not be identified. Future users should be aware of this shortcoming of the Billions et al. [2004. Hydrobiologia 526: 43–53] protocol. As a solution to this problem, we present an improved genetic identification protocol based on a simple double digestion of the rDNA-ITS region with the restriction enzymes BsrB I and EagI. Sequence analyses of rDNA-ITS clones and preliminary testing indicate that the new protocol is unaffected by the rDNA variation which troubled the Mwo I/Sau96 I protocol. Further, the new protocol identifies all European species of the D. longispina complex, as well as their F1 hybrids. However, a wider screening is required to verify its general utility for all species, since yet unknown variation may occur. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Large Hardy-Weinberg equilibrium deviations in the Daphnia longispina of Lake El Tobar

The population structure of Daphnia longispina in Lake El Tobar, Spain was studied by measuring variation at the aldehyde oxidase (AO), phosphoglucose isomerase (PGI) and phosphoglucose mutase (PGM) loci in each of 1337 individuals from four collections. In 9 of the 12 comparisons between observed allele frequencies and those expected by Hardy-Weinberg equilibrium there was an excess of heterozygotes. We found 27 of the potential number of 54 composite electromorphs (clones) based on the three allozymes. Clone diversities were rather high in all collections. Three clones reached frequencies of over 25% and different clones were dominant in each of the four collections. Strong temporal variation was found in the genetic structure of this Daphnia population. This variation was driven by changes in the relative frequencies of the component clones in the lake rather than by a recruitment of novel clones into the population.We conclude with a consideration of the role of models relating allele and genotype frequencies in populations of cyclical parthenogens. Because the breeding system of these populations infrequently involves recombination between clones, models such as the Hardy-Weinberg have limited value in providing meaningful measures of population structure.     

Productivity of Daphnia longispina in a highly humic boreal lake

The development of the Daphnia longispina (O. F. Müller)population in a highly humic boreal lake was followed throughoutone growing season, and the amount of secondary production wasestimated in relation to primary production and available foodresources. The growth rate method was applied in the secondaryproduction measurements. Daphnia longispina did not appear inthe water column until 16 May, after which the animals werepresent throughout the growing season. The population showedthree density peaks; the first appeared in early June, and thesecond and third in mid-July and at the beginning of September,respectively. Somatic production followed a seasonal pattern,with highest production rates in midsummer. The maximum valueof 127 mg C m^–2day^–1 was recorded at the beginningof July. The total annual net production of D. longispina was7.9 g C m^–2. During most of the growing season, the primaryproductivity in the lake was well below 100 mg C m^–2 day^–1and the total annual productivity of photosynthetic algae was5.0 g C m^–2. We conclude that in this lake the zooplanktonpopulation did not rely on phytoplankton primary productionas a sole carbon source, but that most of the carbon must haveoriginated from bacterial production either directly or througha microbial loop.     

Clonal diversity,clonal persistence and rapid taxon replacement in natural populations of species and hybrids of the Daphnia longispina complex

Hybridization is common among cyclical parthenogens, especially in zooplankton species assemblages of the genus Daphnia. To explore hybridization dynamics and the extent of clonal diversity in the Daphnia longispina complex, we analysed population structure in eight permanent lakes. Based on 15 microsatellite loci, three major taxonomic units emerged: two species, D. galeata and D. longispina and their F1 hybrids, supported by factorial correspondence analysis and two Bayesian methods. At the same time, the detection of backcross classes differed between methods. Mean clonal diversity was lowest in the F1 hybrids, as expected from the high rate of asexual reproduction. Within taxa, replicated genotypes were of clonal origin, but clonal lineages persisted in subsequent years in only one of three resampled lakes. In another lake, the taxon composition changed from being dominated by hybrids to complete dominance by one parental taxon. Such a year‐to‐year taxon replacement has not been reported for the D. longispina complex before. Our data on this hybrid complex illustrate that high‐resolution genotyping is essential for the understanding of ecological and evolutionary outcomes of hybridization in partially clonal taxa.     

Spatial patterns in the abundance and distribution of parasites of freshwater fish

It is not yet known whether the abundance and distribution of freshwater fish parasites form spatial patterns. Here, Rita Hartvigsen and Odd Halvorsen review the main approaches to the study of spatial patterns of these parasites, and propose an alternative approach based on meta-population theory and landscape ecology.     

Structural variations of the eye linked to daily vertical migrations of Daphnia longispina

The role of the eyes, and more precisely that of the ommatidian pigments, in the control of daily rhythms of movement of Daphnia longispina is investigated. In the laboratory, under permanent light (LL), the pigmentary modifications observed are globally similar to those observed in situ, except in their timing, around dusk, LL cycles do not coincide with the ascent of daphnids but precede it. This is the expression of an endogenous free-running rhythm. Therefore, in D. longispina, an internal oscillator controls DVMs, which are circadian and not nycthemeral, and, in situ, illumination at dawn suffices to synchronize migrations on dark/light alternation.     

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

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

Ecological genetics of Norwegian Daphnia. II. Distribution of Daphnia longispina genotypes in relation to short-wave radiation and water colour

In the mountain range of South Norway, transparent and melanic members of the Daphnia longispina complex occur. Melanic populations inhabit clearwater lakes and ponds, while transparent populations are found in ponds with slightly humic water. Mixed populations have not been detected. The distribution patterns of the two morphs are related to ambient levels of short-wave radiation, and the light absorptive properties of the inhabited waters. The pigment present is probably melanin, which is deposited in the dorsally directed parts of the carapace, head shield and antennae. Allozyme studies indicate that these distinct morphs are only remotely related, the alpine transparent group being closer to lowland, likewise transparent, populations. A melanic population found at Svalbard is closely related to the melanic mainland populations. Clonal diversity in ponds and shallow lakes is very low, in contrast to populations of deeper lakes.Deceased.     

Filtering rates on natural bacteria by Daphnia longispina and Eodiaptomus japonicus in Lake Biwa

Filtering rates on [³H]thymidine-labelled natural unattachedbacteria and that on [¹⁴C]bicarbonate-labelled natural planktonwhich pass through the 25 µm-mesh-size screen were measuredfor Daphnia longispina and Eodiaptomus japonicus in Lake Biwa.Errors associated with the radioisotope technique, i.e the lossof labels after feeding trials and the self-absorption of thebeta emittance of ³H, were checked and corrected for the calculationof the filtering rates. It was suggested that Daphnia collectsbacteria efficiently, although the efficiency is somewhat variabledepending on food particle composition (i.e. presence and absenceof larger particles) and feeding condition (i.e. animal densityand physical disturbance). By contrast, copepodites of Eodiaptomuswere suggested to be less efficient bacteria feeders. Food resourceexploitation strategies of these two co-existing zooplanktersare discussed.     

Ecology of predator-induced morphological defense traits in Daphnia longispina (Cladocera,Arthropoda)

Oecologia - Inducible defenses against predators are widespread among plants and animals. For example, some Daphnia species form neckteeth against predatory larvae of the dipteran genus Chaoborus....