Does oxygen availability regulate sexual reproduction in local populations of the littoral cladoceran Alonella nana?

We investigated sexual reproduction patterns of a ubiquitous littoral cladoceran Alonella nana (Anomopoda, Chydoridae), using ecological and paleoecological approaches to clarify the forcing mechanisms behind its sexual reproduction. Contemporary sampling of A. nana populations in two environmentally different lakes showed abundant sexual reproduction in the lake with eutrophic, oxygen-deficient conditions and scarce gamogenesis in the oligotrophic, well-oxygenated lake. Sediment core studies from the same lakes indicated that comparable sexual reproduction patterns had prevailed for many centuries. However, in eutrophic Lake Hamptr?sk, A. nana increased its sexual reproduction from 1700 ad onward, consistently with decreasing trend in chironomid-inferred oxygen. The core samples, together with a surface sediment dataset of 25 lakes from southern and central Finland, showed a negative correlation between ephippia of A. nana and winter oxygen, although the surface sediment data per se did not show any significant correlation. When the dataset was scaled locally to include lakes in close proximity in southern Finland, the correlation became clearer. The results imply that the spatial and temporal variations in sexual reproduction of A. nana populations may partly be explained by differences in oxygen levels.     

Chydorid Ephippia as Indicators of Past Environmental Changes – A New Method

I present a new method, ephippium analysis, to investigate past environmental changes which may have affected chydorid Cladocera (Anomopoda, Chydoridae). I studied chydorid ephippia from sediments of two small lakes (Kaksoislammi and Rutikka) in southern Finland. The relative adundance of chydorid ephippia periodically increased during the Holocene. Ephippia were abundant during the late Pleistocene/early Holocene transition, reflecting the severe climate (short open-water season) that existed prior to the onset of rapid postglacial warming. Both lakes experienced an environmental change in the late Holocene. In Kaksoislammi there was a dramatic change in predator–prey relationships following a decline in pH. All chydorids responded with increased gamogenesis. Prehistoric anthropogenic activities caused a rise in the trophic state in Rutikka and probably caused changes in the predator/food web. One chydorid, Alona affinis, reacted first with increased gamogenesis and then almost disappeared. These first results of ephippium analysis indicate that the method has a great potential in detecting periods of environmental change during the Holocene.     

Reproductive strategy of the alien Ponto-Caspian cladocerans Cercopagis pengoi (Ostroumov, 1891) in the Baltic Sea (by example of the Vistula Lagoon)

Early and prolonged sexual reproduction has been registered in the population of Cercopagis pengoi within the Vistula Lagoon relative to its original habitat. Decrease in the role of parthenogenesis and increase in the frequency of gamogenesis have been registered in the population under study. It has been accompanied by reduction in the average number of parthenogenetic eggs per female and increase in the production of gamogenetic eggs relative to the Caspian Sea.     

Reproduction in Laboratory Stocks of Pleuroxus (Chydoridae,Cladocera) Under the Influence of Photoperiod and Light Intensity

The effects of photoperiod and light intensity on the reproduction of laboratory populations of three species in Pleuroxus from different geographic origins were determined statistically. There were highly significant differences in the numbers of gamogenetic individuals produced between the populations at different photoperiods but not at different light intensities. More males than ephippial females appeared at long-day photoperiods. Pleuroxus denticulatus stocks showed greater variation of gamogenetic response to photoperiod and geographic origin suggesting that local populations have evolved different reproductive patterns to meet the major environmental stresses of the region.     

Subfossil chydorid (Cladocera, Chydoridae) ephippia as paleoenvironmental proxies: evidence from boreal and subarctic lakes in Finland

Chydorids (Cladocera, Chydoridae) have two reproductive strategies: asexual reproduction that prevails during favorable environmental conditions and sexual reproduction that is induced by environmental stimuli associated with seasonal or aperiodic environmental stresses. These modes of reproduction can be recognized in the subfossil sedimentary records as parthenogenetic shells of females (asexual reproduction) and by ephippia (sexual reproduction). We studied the interrelations between subfossil chydorid ephippia and environmental variables by analyzing surface sediment samples obtained from 76 Finnish lakes across a latitudinal gradient (60–70°N). The results showed that the total chydorid ephippia (TCE) increases along the climate gradient from ~2 to 3% in the south to ~25% in the north and suggested a significant dependence (r ~ −0.8, P < 0.001) with several climate factors, especially that of mean July air temperature. We used this relationship to create a model for reconstructing past mean July air temperatures. A linear regression of the log₁₀ transformed TCE as a single independent variable explained 76% (SE ± 0.76°C) of the variance of the observed mean July air temperatures. Accordingly, we propose that this novel tool may be highly suitable for reconstructing paleotemperatures in cold-temperate environments.     

The impact of conifer plantation forestry on the Chydoridae (Cladocera) communities of peatland lakes

Conifer plantation forestry is recognised as a potential source of diffuse pollution to surface waters and represents a risk to their ecological status. In this study, the water chemistry and Chydoridae (Cladocera) communities of 26 small blanket bog lakes were investigated to assess the impact of plantation forestry. The study was conducted over a 12-month period in 2009?C2010 by comparing lakes with three distinct catchment land uses: (i) unplanted blanket bog only present in the catchment, (ii) mature conifer plantation forests only present in the catchment and (iii) catchments containing mature conifer plantation forests with recently clearfelled areas. All three catchment land uses were replicated across two geologies: sandstone and granite. Lakes with afforested catchments had very high concentrations of plant nutrients (P and N), total dissolved organic carbon (TDOC) and heavy metals (Al and Fe), the highest concentrations being recorded from the clearfelled lakes. Similarly, the chydorid communities differed between lakes of contrasting catchment land use. The dominance of Alonopsis elongata in the unplanted blanket bog lakes shifted to dominance by the smaller bodied Chydorus sphaericus, along with Alonella nana, Alonella excisa and Alonella exigua, in the plantation forestry-effected lakes, consistent with a shift in lake trophy. Our findings have shown that plantation forestry can have a profound impact on the water quality of small peatland lakes, especially at the clearfell stage. The response of the chydorid communities is consistent with plantation forestry exerting a trophic, rather than an acidic or toxic, effect on lake ecosystems.     

Demographic variation of dwarf birch (<Emphasis Type="Italic">Betula nana</Emphasis>) in communities dominated by <Emphasis Type="Italic">Ledum palustre</Emphasis> and <Emphasis Type="Italic">Vaccinium uliginosum</Emphasis>

The structure and demographic processes were compared in shrub communities to test the effects of vegetation succession on population growth, fecundity and abundance of the dwarf birch (Betula nana L.), which is a rare and endangered plant species in Poland and a glacial relict in Central Europe. The effects of Ledum palustre L. and Vaccinium uliginosum L. were studied in the Linje nature reserve in Chełmińskie Lake District (northern Poland), in three permanent plots on a peat bog. Vegetative growth and reproduction of B. nana were lower in plant communities dominated by L. palustre and V. uliginosum, than in a reference site. Fecundity was also lower, despite the fact that the percentage share of potentially fertile age groups was similar in all study sites. Mortality of ramets was independent of vegetation, both for juvenile and mature stages. The results confirm that B. nana is intolerant of shade, and it is more abundant in vegetation without competitors. Light limitation can lead to its decline, primarily by a decrease in vegetative growth. Sexual reproduction may be negatively affected by shade, but it plays only small role in population growth. Butterfly larvae can destroy inflorescences, and thus contribute to low effectiveness of sexual reproduction. Increasing density of shrubs and trees in peat bogs can reduce the abundance of dwarf birch, and can lead to the extinction of its local populations.     

Diverse,endemic and polyphyletic clones in mixed populations of a freshwater snail (Potamopyrgus antipodarum)

Understanding the source and diversity of clones is necessary to resolve the complicated issues surrounding the apparent evolutionary stability of sexual reproduction. The source of clones is important because present theory is based on an “all else equal” assumption, which is predicated on the idea that clonal mutants are derived from and compete with local sexual populations. Clonal diversity is important because it reduces the advantage of sexual reproduction under either soft selection (the Tangled Bank Hypothesis) or under strict frequency-dependent selection (the Red Queen Hypothesis). In the present study, protein electrophoresis was used to determine the source and diversity of clones in a freshwater snail (Potamopyrgus antipodarum) in four glacial lakes in which sexual and clonal females were thought to coexist. The results showed (1) that the populations were mixtures of diploid sexual and triploid asexual individuals, (2) that genotypic diversity of clonal populations is very high in all four lakes (but lower than in the sympatric sexual populations), and (3) that the clones are polyphyletically derived from their sympatric sexual populations. Consequently, repeated mutation to parthenogenetic reproduction since the Pleistocene has introduced a different and diverse set of clones in all four lakes. Such diversity may provide a challenge for the ecological theories of sex that rely on frequency-dependent selection.     

Benthic communities of exposed littoral sand-flats in eighteen Dutch lakes

Benthic communities of the exposed sandy littoral zone in eighteen Dutch lakes were sampled using a small, hand-held core (surface area 0.8 cm²). A community dominated by chironomid larvae, chydorid cladocerans, nematodes and copepods was found to be characteristic for this habitat, especially in large and/or deep lakes. Bray-Curtis similarities between sites were generally high (>70%). A hierarchical cluster analysis distinguished two clusters at a 75% similarity level only, varying in copepod and chydorid densities. The chironomidsCladotanytarsus andStictochironomus reached high densities in many lakes, whileRhynchotalona falcata, Monospilus dispar, Disparalona rostrata, Alona spp. andChydorus spp. (mainlyChydorus piger) were the most abundant chydorid taxa.     

Effect of waterlogging on regeneration in the dwarf birch (<Emphasis Type="Italic">Betula nana</Emphasis>)

The aim of this study was to determine the effect of waterlogging on annual regeneration in the dwarf birch (Betula nana L.), and to determine which demographic factors are responsible for the absence of B. nana in the waterlogged parts of a peat-bog. Growth parameters and demographic data were collected from 2001 to 2003 at four sites in the Linje peat bog in the Chełmno Lake District in northern Poland. Three of the sites represented waterlogged open bog communities. The fourth site served as the reference. The density of first-year ramets, annual shoot growth, and the proportion of flowering shoots was significantly lower at the waterlogged sites than at the reference site. No seedlings were found during the course of the study, even though flowering and fruit production took place at the reference site. The flowering shoots were mainly between nine and thirteen years old. Ramets in this age range were found only at the reference site due to high mortality in mature ramets at the waterlogged sites. Mean ramet height was significantly lower at the waterlogged sites than at the reference site, and was strongly correlated with mean ramet age. The distribution of B. nana in peat bogs is limited primarily by two demographic factors: the high mortality rate in older ramets, and the low rate of vegetative reproduction. Another contribution factor is that the proportion of older fertile ramets is low. The effects of fluctuations in the water table on distribution of B. nana are also discussed.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

樟叶蜂两性生殖与孤雌生殖方式下雌虫生殖适合度及子代生活史特征的比较

【目的】樟叶蜂Mesoneura rufonota是危害樟树Cinnamonum campora的重要食叶性害虫,该虫的繁殖策略包括两性生殖和孤雌生殖两种模式。本研究旨在明确孤雌生殖在樟叶蜂生活史中的生物学意义。【方法】在室内25℃恒温条件下,测定并分析了樟叶蜂孤雌生殖和两性生殖两种生殖方式在亲代生殖适合度(雌虫寿命、产卵量和卵孵化率)和子代生活史(各虫态发育历期、死亡率、子代性比和产卵量等)特征上的差异。【结果】孤雌生殖的樟叶蜂雌虫寿命显著长于两性生殖的雌虫寿命,而雌虫产卵量和卵孵化率在两种生殖方式间均无差异。子代各虫态的发育历期和死亡率以及子代单雌产卵量在两种生殖方式间均无差异,但子代成虫性比在两种生殖方式间存在显著差异,表现为孤雌生殖大多产雄性子代,而两性生殖大多产雌性子代。【结论】樟叶蜂的孤雌生殖延长了亲代雌虫的寿命,且为产雄孤雌生殖。这些研究结果表明,樟叶蜂的孤雌生殖不但具有自身建群的能力,同时在种群繁衍中可以提供大量的雄虫以弥补两性生殖后代雄性个体的不足。     

Taxonomic Status and Reproduction Dynamics of the Non-Indigenous Cercopagis in the Gulf of Riga (Baltic Sea)

Overwintering population of Cercopagis is represented by resting eggs which hatch generally in May. Although representatives of the first parthenogenetic generation (the spring form individuals) differ morphologically from individuals of the subsequent generations (the summer form individuals) and could be keyed to a separate species – Cercopagis (Apagis) ossiani – our analysis confirm that there is only one Cercopagis species in the Gulf of Riga: Cercopagis pengoi. Notable seasonal dynamics was observed for the gamogenetic mode of reproduction, being strongly associated with the total population density. Gamogenetic females develop since the end of June and reach the maximum absolute abundance in July whereas their proportion is generally the highest in fall. The gamogenetic absolute fecundity drops to the lowest level late July but increased afterwards until the late August. One resting egg was, an average, found in 45.4%, two in 53.4% and three in 1.2% of females. Parthenogenetic fecundity was significantly higher in May–June compared to other months studied. Brood pouch of parthenogenetic females was found to contain 11.6 ± 1.0 and 10.2 ± 0.3 embryos in the spring and summer form individuals, respectively. Variation in the two modes of reproduction and fecundity is probably solely not controlled by temperature, but also by food availability and population density.     

Food quality triggers the reproductive mode in the cyclical parthenogen <Emphasis Type="Italic">Daphnia</Emphasis> (Cladocera)

Cyclic parthenogenesis (heterogony) is a widespread reproductive mode found in diverse taxa such as digenean trematodes, gall wasps, gall midges, aphids, cladocerans and rotifers. It is of particular interest as it combines the advantages of asexual reproduction (rapid population growth) and sexual reproduction (recombination). Usually sexual reproduction is initiated when, or slightly before, environmental conditions deteriorate, and often results in the production of resting stages. The optimal timing of diapause induction must thus be under strong natural selection. Using the cladoceran Daphnia as a model system, we show here for the first time that the switch from parthenogenetic to sexual reproduction in a cyclical parthenogenetic organism can be influenced by the chemical composition of food. Under crowding conditions Daphnia reproduced parthenogenetically with subitaneous eggs when fed the algal species Cryptomonas sp., but started the production of resting eggs when fed with the green algal species Scenedesmus obliquus. Supplementation experiments with lipids and especially proteins showed that the induction of resting egg production in two clones of different Daphnia species was due to a dietary deficiency in the green alga. Hence, the low food quality induced a switch in the reproductive mode that may contribute to optimal timing of the sexual reproduction of Daphnia in nature. Furthermore, our results have two other major implications: first, they suggest that protein compounds should be added to the list of diet constituents potentially limiting or influencing Daphnia reproduction. Second, we show that the role of food quality goes far beyond the up to now documented effects of food quality on somatic growth and trophic transfer efficiency of herbivores: due to its effects on sexual reproduction and the production of resting eggs, food quality might influence genetic diversity and long-term persistence of Daphnia in lakes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Cladocera in space and time: Analysis of lake sediments

Shells of Bosminidae and Chydoridae are quantitatively preserved in lake sediments. The chronological deposition of these remains provides the means for longterm observation of these Cladocera, both in terms of species and communities. Chydorid analysis, as based on subfossil assemblages, is an analysis of community and provides direct observation of community dynamics over extended periods of time. It has proved to be a valuable method to obtain information on the influence of environmental factors and time on community characteristics. Morphological variation inBosmina (Eubosmina) has been followed for some thousand years. This is of special interest for the evaluation of taxonomic rank (species, forms) if closely related taxa have co-existed. Bosmina successions, as well as shifts in the chydorid fauna, are related to environmental change. Thus, cladoceran analysis of lake sediments provides information on the developmental history of lakes and allows observation of the effects of longterm environmental changes, such as climatic changes and eutrophication.     

Cladocera assemblages from the surface sediments of south-central Ontario (Canada) lakes and their relationships to measured environmental variables

Fossil cladoceran remains preserved in surface sediment samples from 44 oligotrophic lakes in south-central Ontario were examined to evaluate the relationships between species assemblages and measured environmental variables. Differences in cladoceran assemblages were related to physical and chemical variables using multivariate techniques. Redundancy Analysis (RDA) identified five environmental variables as significantly influencing assemblage composition: sulphate (SO₄²⁻), calcium (Ca²⁺), pH, maximum lake depth (Z _max) and dissolved organic carbon (DOC). There was a distinct separation of lakes and taxa along the ion gradient based on SO₄, Ca and pH. Additionally, cladoceran communities in coloured, shallow lakes had relatively higher abundances of littoral chydorid species and the pelagic taxa Holopedium spp., and the Daphnia pulex complex. Deep, clear lakes had relatively higher abundances of other pelagic taxa. Predation by fish (measured as presence–absence) and Chaoborus (measured as density) were less significant than some of the physico-chemical variables in influencing cladoceran assemblage structure. However, this could be due to the limited resolution of the predation data that was available at the time of this study. The distribution of cladocerans in the surface sediment, and their relation to these important environmental variables, suggests that there is considerable potential for the use of sedimentary cladoceran remains as environmental indicators in south-central Ontario lakes. Handling editor: J. Saros     

Distribution of cladoceran assemblages across environmental gradients in Nova Scotia (Canada) lakes

Cladocera sub-fossils have widely been recognized as useful environmental indicators. Nevertheless, investigations into the distribution and environmental controls on Cladocera are scarce in North America, hindering their use in paleolimnological studies. Here, we examine cladoceran assemblage and size structure from the surface sediments of 49 soft-water Nova Scotia (Canada) lakes to provide ecological data on this key taxonomic group, the first study of its kind for the northern Atlantic coast of North America. We found that Nova Scotia lakes contain a diverse chydorid assemblage, but were generally dominated by the smaller pelagic herbivore Eubosmina longispina. Daphnia, a larger pelagic herbivore, was relatively uncommon in these lakes. Redundancy analyses (RDA) identified maximum lake depth and dissolved organic carbon (DOC) as the environmental variables that best explained the structuring of these cladoceran assemblages. Generalized linear models were then used to better characterize the ecological associations for individual taxa. Body size of the bosminids in these lakes was significantly correlated only to total nitrogen (TN), with larger bosminids in lakes with lower TN values. Bosminid mucro length, an indication of invertebrate and fish predation pressure, was significantly related to TN, maximum lake depth, and lake surface area. The ecological information provided by this study should assist the interpretation of paleolimnological assessments of environmental change in these and other similar lakes.     

Regional community assembly drivers and microbial environmental sources shaping bacterioplankton in an alpine lacustrine district (Pyrenees,Spain)

Microbial communities in natural ecosystems are subject to strong ecological rules. The study of local communities along a regional metacommunity can reveal patterns of community assembly, and disentangle the underlying ecological processes. In particular, we seek drivers of community assembly at the regional scale using a large lacustrine dataset (>300 lakes) along the geographical, limnological and physico-chemical gradients in the Pyrenees. By using high throughput amplicon sequencing of the 16S rRNA gene, and inferring environmental sources of bacterial immigrants, we showed that surface aquatic bacterial assemblages were strongly influenced by terrestrial populations from soil, biofilms or sediments, and primarily selected by a pH-alkalinity gradient. Indeed, source proportions explained 27% of the community variation, and chemistry 15% of the total variation, half of it shared with the sources. Major taxonomic groups such as Verrucomicrobia, Actinobacteria and Bacteroidetes showed higher aquatic affinities than Parcubacteria, Gammaproteobacteria, Alphaproteobacteria or Betaproteobacteria, which may be recruited and selected through different hydrographic habitats. A regional fingerprint was observed with lower alpha diversity and higher beta diversity in the central Pyrenees than in both ends. We suggest an ecological succession process, likely influenced by complex interactions of environmental source dispersal and environmental filtering along the mountain range geography.     

Spatial distribution of subfossil Chironomidae in surface sediments of a large, shallow and hypertrophic lake (Taihu, SE China)

Spatial heterogeneity of benthic communities has clear implications for estimating lake production, biodiversity as well as identifying representative sites for palaeolimnological studies. This study investigates chironomid variability and the controlling factors (i.e., environmental and spatial variables) in surface sediments from Taihu Lake (2,338 km²), a hypertrophic lake in the Yangtze delta in eastern China. The spatial distribution of chironomids shows distinct heterogeneity. Microchironomus tabarui-type and Tanypus dominate the midge communities around the estuaries, while Cricotopus sylvestris-type and Polypedilum nubifer-type are the predominant taxa in the East Bays and the East Taihu Lake. Redundancy analysis was used for exploring the relationships between chironomid variability and environmental and spatial stressors. Four variables were identified as significant factors that influence chironomid community structures. The high nutrient concentrations around the estuarial areas favor the development of nutrient-tolerant taxa. Water depth-related oxygen depletion in the open lake during algae blooms prohibits the survival of many organisms, except for a few hypoxic-resistant species. High transparency in the East Bays and the East Taihu Lake indirectly creates a favorite microhabitat for macrophyte-associated chironomid species through aquatic plants. Space per se is a significant forcing factor for organism community and distribution at scales of >1,000 km². It might be important to consider spatial variables more explicitly in future studies of chironomids in large lakes where multiple stressors make the interactions within the ecosystem more complicated. This study aims to illustrate the ecological characteristics of specific chironomid taxa related to a “microecosystem” which is contributed by the multiple environmental gradients within a large lake, and to provide empirical support for interpretation of palaeochironomid data.     

Population dynamics of Stylaria lacustris (Linnaeus, 1767) (Oligochaeta,Naididae) in Los Talas,Argentina

Population dynamics of Stylaria lacustris were analyzed over 2 years in a pond located at Los Talas, Argentina. In both years, the peak in abundance, due to intense asexual reproduction, fell at the end of winter, and was followed by mature individuals. The species was not collected during summer. Temperature was the main factor regulating the population through both sexual and asexual reproduction. In the second year, density was lower and correlated with a lower water level and a reduced vegetation development. Individuals were randomly distributed in periods of high abundance and contagious when density was low. Generation time was 15 days.