New microsatellite primers for Daphnia galeata mendotae

In order to study genetic changes in populations of Daphnia galeata mendotae, I characterized seven polymorphic microsatellite loci. Primers to amplify these loci were tested on individual eggs from the resting egg bank of Onondaga Lake, NY. Levels of polymorphism and cross‐amplification in D. g. galeata indicate that they will be useful markers for ecological genetic studies on both adults and diapausing eggs of these species.     

Effects of winter conditions on Daphnia dynamics and genetic diversity in a dimictic temperate reservoir

1. Winter conditions shape plankton dynamics and community composition in temperate regions, but their effect on dynamics and genetic composition of cyclical parthenogens like Daphnia is largely unclear. 2. For 5 years, we studied the dynamics, hatching from resting eggs and genetic structure of a D. galeata × longispina hybrid complex in a dimictic, temperate reservoir. Our main hypothesis was that higher spring densities and an earlier population peak will be observed after warmer winters, with a lower genetic diversity because of a lower contribution of resting eggs to population growth. 3. The study period could clearly be categorised into cold‐winter years (n = 3) and warm‐winter years (n = 2). Daphnia densities at the end of spring overturn were ～10‐fold lower after cold winters than after warm ones, but no pattern emerged concerning the timing and the height of the population peak in early summer. 4. Hatching intensity from resting eggs was higher and contributed up to 8.5% to Daphnia abundance in a cold‐winter year compared to a negligible contribution in a warm‐winter year. Consistent with this finding, new multilocus genotypes (MLGs) adding to the overwintering stock after the end of spring overturn and presumably originating from resting eggs increased genetic diversity and attained high frequencies within the population only after a cold winter. New MLGs were recorded also after warm winters, but they never gained dominance and no shift in genetic diversity was observed. However, genetic diversity was not generally reduced after warm winters. 5. Our results confirm earlier findings that winter conditions have only a limited effect on the main growth phase and the peak of Daphnia during late spring and early summer. However, winter conditions determine the contribution of resting eggs to the population development, which may profoundly alter the genetic composition of the population compared to the previous season.     

The importance of hatching from resting eggs for population dynamics and genetic composition of Daphnia in a deep reservoir

1. Cyclical parthenogens, organisms switching between sexual and asexual mode of reproduction, are common in aquatic habitats. In permanent waterbodies with year‐round persistence of populations, the relevance of sexually produced resting eggs for population dynamics and structure is still largely unexplored. 2. The aim of this study was to investigate the importance of recruitment from resting eggs to Daphnia spp. in a deep dimictic reservoir with overwintering populations (Saidenbach Reservoir, Germany). We linked field studies on Daphnia population dynamics, hatching rates and diapausing egg abundances in the sediment with allozyme analysis of the genetic variability of both the pelagic population and hatchlings from resting eggs. 3. We found higher hatching rates at intermediate depths (10–18 m), although highest diapausing egg abundances were found in the deeper parts of the reservoir. From the extensive field data, we calculated a population budget for the main basin of the reservoir, which showed that hatched daphnids accounted for <1% of the total abundance and explained only a tiny fraction of population growth. 4. The genetic composition (MultiLocus Genotypes, MLG) of the pelagic population was relatively stable throughout the stratified period with up to five among 39 detected MLGs attaining frequencies >5% per date. From 11 MLGs differentiated among hatched daphnids, three were recovered in the pelagic population in perceptible, albeit low frequencies (1–4%). 5. We conclude that the contribution of hatched animals to the spring population dynamics is negligible in situations with an overwintering population present and will only become relevant if densities are extremely low. In this type of system, the significance of the resting egg bank is presumably mainly related to the maintenance of genetic diversity.     

Russian Wheat Aphid,Diuraphis noxia in the Czech Republic – cause of the significant population decrease

Diuraphis noxia is a newly established alien pest in the Czech Republic. There were two population explosions during the period 1993–2012. A successful overwintering of the anholocyclic populations of D. noxia was the cause of a population explosion in the oncoming growth season. Viviparous parthenogenetic females of the anholocyclic populations and diapausing eggs of holocyclic populations could overwinter at the same time. Analysis of the course of winter temperatures showed that a complete winter mortality of anholocyclic populations depended mainly on the duration of the freezing period, when temperature dropped below ?5°C, and on the intensity of the frost. We proposed the cumulative effective freeze (in minus degree days) as a key parameter for the aphid anholocyclic population's mortality during overwintering. This parameter equalled ?10.1 degree days for relatively warm winter period 2006–2007, whereas in the relatively cold winter period of 2009–2010, this parameter reached ?87.1 DD, which caused complete mortality of anholocyclic populations. A cumulative effective freeze could be used for short‐time prognosis of D. noxia population explosion.     

Local variation in overwintering stages ofPanonychus akitanus Ehara (Acarina: Tetranychidae)

The overwintering behaviour ofPanonychus akitanus Ehara was compared in a population (Sapporo) overwintering as the egg and the adult female and in a population (Tomakomai) overwintering in the egg stage only. In both populations, winter eggs were larger in size and darker in colour than summer eggs, and females laying winter eggs showed clear differences in body colour and behaviour from summer females. Both populations produced winter eggs in response to short photoperiods, and the critical photoperiod was between 13.0 and 13.5 h in the Sapporo population and between 13.5 and 14.0 h in the Tomakomai population. The hatchability of winter eggs of the Sapporo population collected from the field began to increase in November, and that of the Tomakomai population in December. In the Sapporo population, a 40-day or more exposure to 5°C promoted hatching of winter eggs, and a 50-day exposure of females which had previously laid some winter eggs to 5°C induced the production of summer eggs. Females of the Sapporo population laid winter eggs in autumn, and then themselves overwintered as well, and produced summer eggs in the following spring. These females were called winter-habit females.The winter eggs of both populations were diapausing. However, the winter-habit females of the Sapporo population were not reproductively inactive and their condition of diapause was not certain. The ecological significance of the winter-habit females in the Sapporo population is discussed.     

Long-term genetic shifts in a microcrustacean egg bank associated with anthropogenic changes in the Lake Constance ecosystem

Diapausing egg banks of aquatic zooplankton have the potential to remain viable for decades or even centuries, and can thus harbour potentially high levels of genetic variation. Diapausing (ephippial) eggs from the Daphnia galeata–hyalina complex (Crustacea: Anomopoda) in Lake Constance (Bodensee), Germany, were isolated from sections of dated sediment cores, hatched in the laboratory, and established as a clone bank. We used cellulose acetate electrophoresis at four polymorphic enzyme loci (Pgm, Pgi, Ao, and Got) to examine long-term temporal changes in the genetic composition of the hatchling pool. Our results indicate that significant shifts have occurred in the genetic structure of this population, which parallel concomitant shifts in the trophic state of Lake Constance during the past 25–35 years. Here we discuss the utility of egg bank propagules as good model organisms to study microevolution, as related to past environmental change.     

Resting eggs of lake-Daphania II. In situ observations on the hatching of eggs and their contribution to population and community structure

SUMMARY. 1. To investigate the contribution of hatchlings from lake- Daphnia resting eggs (ephippia) to population and community structure, hatching was monitored in situ from April to November 1986 in two north German lakes. Hatching traps were placed on the sediments and the incidence, genetic and species composition of ephippial hatchlings determined twice-weekly.
2. Hatching began in April after the thawing of ice. and continued for 3–4 weeks until the lakes stratified. There was no obvious relationship between the onset and duration of hatching and the environmental variables recorded, namely Secchi depth, surface temperature, temperature and oxygen concentration at the sediment-water interface.
3. Among the hatchlings were D. galeata, D. hyalina and their interspecific hybrid.
4. In the lake which contained overwintering animals the number of ephippial hatchlings was approximately one third of the total number of juveniles present in the lake.
5. In the second lake, no adults were recorded during winter and the population was probably founded by ephippial hatchlings alone. There was good agreement, in the short term, between the proportion of each species represented among hatchlings and the subsequent species composition in the lake.
6. The hatchlings were genetically diverse, and alleles were representative of those alleles present in contemporary populations.     

Variation in ephippial buoyancy in Daphnia pulicaria

1. Freshwater zooplankton often produce diapausing eggs to survive environmental stress. The diapausing eggs of Daphnia (Crustacea, Cladocera) are encased in an ephippium that either floats at the surface or sinks to the sediment. These two types of ephippia may represent different strategies between spatial (floaters) and temporal (sinkers) dispersal of offspring. 2. We observed floating and sinking characteristics of ephippia obtained from eight lakes. We then conducted an experiment with 26 Daphnia pulicaria clones obtained from six of these lakes and observed the production of buoyant and non‐buoyant ephippia under high and low food conditions. 3. Ephippia were more often non‐buoyant than buoyant both from females caught in nature and those reared in the laboratory. The experiment revealed that each clone was able to produce both types of ephippia, but that there was considerable among‐clone variation in the percentage of non‐buoyant ephippia produced. 4. We conclude that production of non‐buoyant versus buoyant ephippia may be driven by both genetic and environmental factors.     

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.     

Population dynamics and seasonal variation in the embryonic dormancy of Pilophorus gallicus (Hemiptera: Miridae): ‘don't put all your eggs in one basket'

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Colonization of Daphnia magna in a newly created pond: founder effects and secondary immigrants

In habitats recently colonized by cyclical parthenogens, founder events lead to genetic differences between populations that do not erode quickly despite ongoing dispersal. By comparing the genetic composition during initial colonization with that of the diapausing egg bank at a local scale, we here present the relative contribution of the founding clones to the build-up of genetic diversity and differentiation of a newly established cladoceran population. We monitored the population genetic structure of Daphnia magna in one newly created pond as well as the diapausing egg banks of four water bodies in the neighbouring area. Our population was founded by four individuals. After the first growing season, the largest contribution to the sexually produced resting egg bank came from only two clones. Descendants of initially rare clones and potentially also additional immigrant clones profited from outbreeding vigour and increased their frequency during the first few years after colonization. Beyond this, no further significant changes in genetic structure were observed in the egg bank. At this point, priority effects became fully operational and led to sustained population genetic differentiation from nearby ponds. Our results support that colonization dynamics strongly influence within and among population genetic variation and evolutionary potential of populations.     

Diapause dynamics of two diaptomid copepod species in a large lake

Skistodiaptomus oregonensis and Leptodiaptomus minutus produce diapausing eggs in a large lake, Oneida Lake, in New York State. The timing of the switch from production of subitaneous (immediately hatching) eggs to diapausing eggs for both species is in October. This timing is consistent with a pattern, reviewed here, for other populations of diaptomid copepods: populations living in large lakes tend to begin production of diapausing eggs later in the season than those living in small lakes. Populations living in temporary ponds tend to switch still earlier in the season. All populations reviewed here live in the north temperate zone. The sediments of Oneida Lake contain densities of diaptomid diapausing eggs on the order of 10⁵ m^–2 per cm below the sediment surface down to 5 cm. Below this sediment depth, egg densities decline. The highest egg densities were found in sediments under the deepest water. Diapausing eggs of L. minutus survive in the sediments at least two years, as shown by the recovery of the population after a year in which no new diapausing eggs were produced, and probably for two or more decades. Long-term dormancy can have the effect of ensuring the continuation of a population through periods of poor recruitment, and can help create conditions for the coexistence of competing species. Other investigators have suggested that S. oregonensis and L. minutus are competitors in other lakes when they co-occur.     

Extended lifespan traded for diapause in Daphnia

1. Most freshwater crustaceans of the genus Daphnia are cyclically parthenogenetic organisms that are well adapted to unstable habitats due to their short life cycle, wide phenotypic plasticity, and the ability to produce protective diapausing eggs in anticipation of environmental deterioration. Short lifespan and heterogonic reproduction are typical features of Daphnia in a broad spectrum of freshwater habitats, from small temporary pools to large permanent lakes. However, in some locations, departures may be observed from this typical life history pattern to obligate asexuality or extended lifespan. 2. A 3‐year field study in a deep ultraoligotrophic fish‐free alpine‐type lake (Czarny Staw in the Tatra mountains in southern Poland) revealed the coexistence of two closely related asexual lineages of Daphnia of the pulex complex, which differ in body colour (transparent versus orange) and in their strategies for surviving long winters. 3. The ‘transparent’ clone of European origin exhibits an ephemeral lifestyle. It completes its life cycle within a single season, forming two generations of active specimens during the short summer and producing diapausing eggs late in the season. Transparent individuals live no longer than 5–6 months in this cold lake and survive winter exclusively in the form of diapausing eggs. 4. Individuals of the ‘orange’ lineage, which are closely related to eastern Nearctic Daphnia pulicaria, exhibit a biennial lifestyle unique to this genus. They do not form diapausing eggs or produce them only occasionally. Instead, they are active throughout the year and live for up to 13–14 months. Reproduction is postponed until the second year of life when food conditions have improved. Rich lipid reserves accumulated in the first season fuel them during the long winter and permit early reproduction the following spring. 5. Possible reasons for the evolution of obligatory parthenogenesis and long lifespan in Daphnia are discussed.     

Sexual reproduction and diapause of Hexarthra sp. (Rotifera) in short-lived ponds in the Chihuahuan Desert

1. In the life cycle of monogonont rotifers it is generally assumed that diapausing eggs invariably hatch into amictic stem females which produce female offspring parthenogenetically. Diapausing eggs are only produced by later generations after sexual reproduction has been induced by environmental cues. 2. We show that populations of an undescribed Hexarthra species inhabiting small temporary ponds in the Chihuahuan Desert deviate from this life cycle pattern. These ponds may dry within days and up to 85% of females were mictic. Females producing male offspring and diapausing eggs were observed 1 or 2 days, respectively, after ponds had filled with water. 3. Under laboratory conditions, 7–46% of females hatching from re‐hydrated sediments were sexual. Male offspring of these females can fertilise other mictic stem females leading to diapausing egg formation. In laboratory experiments, females produced fully developed diapausing eggs within 1.9 days at 20 °C and 1.2 days at 30 °C. 4. In addition, embryonic development time (1.1–0.3 days at temperatures between 12 and 30 °C) and juvenile period (2.1–0.5 days for the same temperature range) are shorter than those of other rotifer species. In short‐lived habitats, the potential for rapid population development and production of new diapausing eggs may be crucial in the long‐term survival of populations.     

Suppression of net transpiration by multiple mechanisms conserves water resources during pupal diapause in the corn earworm Helicoverpa zea

One critical aspect of an insect's ability to overwinter successfully is the effective management of its water resources. Maintenance of adequate water levels during winter is challenging because of the prevailing low relative humidity at that time of year and the short supply of environmental water that is not in the form of ice. These issues are further exacerbated for insects overwintering as pupae, comprising an immobile stage that is unable to move to new microhabitats if conditions deteriorate. The present study compares the water balance attributes of diapausing and nondiapausing pupae of the corn earworm Helicoverpa zea Boddie, aiming to identify the mechanisms used by diapausing pupae to maintain water balance during winter. Diapausing pupae are 10% larger than nondiapausing individuals. Water loss rates for nondiapausing pupae are low (0.21 mg h^?1) and are suppressed (0.01 mg h^?1) in diapausing pupae. Cuticular lipids, which serve to waterproof the cuticle and thus suppress cuticular water loss, are more than two‐fold more abundant on the surface of diapausing pupae, and oxygen consumption rates during diapause drop to almost one‐third the rate observed in nondiapausing pupae. Water gain can be accomplished only when atmospheric water content is near saturation or during contact with free water. At moderate relative humidities (20–40%), water loss rates are very low for diapausing pupae, suggesting that these moth pupae have robust mechanisms for combating water loss. The exceptional ability of H. zea to suppress water loss during diapause is probably a result of the combined effects of increased size, more abundant cuticular lipids and decreased metabolic rates.     

Population dynamics and production of Daphnia hyalina and Bosmina longirostris in a shallow, eutrophic reservoir

SUMMARY. 1. The population densities, dynamics and production of Daphnia hyalina and Bosmina longirostris were studied over a 2-year period in a shallow eutrophic reservoir in eastern England. The diet of the two species was assessed and their ecology was compared in relation to environmental factors. 2. Daphnia hyalina was characterized by small overall body size, probably due to heavy size-selective predation by fish. The first generation produced from the overwintering population appeared in April and Depopulation peaked in late spring and early summer. Adult survival was poor. Peak egg production varied between years, while mean brood size fell in summer probably due to food limitation during the cyanobacterial bloom. At this time selection for individual algal species was high and many guts were empty. Reproductive ratios were inversely related to population density. Bosmina longirostris exhibited a similar annual cycle. 3. Patterns of production were very different in the 2 years in D. hyalina. In 1981 production was high from August to early October, but in 1982 the peak was in spring. Annual production was 32.4 g C m^?2 in 1981 and 13.3 g C m^?2 in 1982. In 1981 B. longirostris production peaked in spring and late summer; in 1982 there was a high production in summer only. Annual production was 3.2 g C m^?2 in 1981 and 2.2 g Cm^?2 in 1982. 4. The number of adult D. hyalina was highly correlated with water temperature and total phytoplankton biomass, but these were negatively correlated with the number of eggs, mean brood size and instantaneous birth rate. In contrast, clutch size of B. longirostris was positively correlated with water temperature while birth rate was positively correlated with both temperature and phytoplankton biomass. The biomass of both species was positively related to total phytoplankton biomass, but neither relationship explained much of the variation in zooplankton biomass. 5. The significance of the observations on plankton in terms of managing the water quality in eutrophic reservoirs is discussed.     

Seasonal variability of the embryonic development time of three planktonic crustaceans: dependence on temperature,adult size,and egg weight

The seasonal variability of the temperature dependence of embryonic development in Daphnia galeata, D. hyalina and Eudiaptomus gracilis of Lake Constance was investigated. We found a significant correlation between the adult size and the variability of the temperature function in all three species, and only in case of D. hyalina a significant influence of egg weight on the temperature function. We could demonstrate no general trend in the influence of the thermal adaptation of parent animals on the temperature function, but some specifically different responses among the investigated species.     

Seasonal and geographical variation in diapause and cold hardiness of the Asian corn borer,Ostrinia furnacalis

Asian corn borer, Ostrinia furnacalis (Guenée), is a key corn pest in the Asian‐Western Pacific countries. It overwinters as full‐grown larvae in plant stalks or in a spun‐silk covering located in the plant debris in the temperate regions of China. Supercooling point (SCP) and survival rate after low sub‐zero temperature treatment were assessed for field‐collected populations in the laboratory using a cool bath with a 1°C/min cooling rate until ?40°C. Mean SCPs were varied among geographical populations, with a significant decline from ?22.7°C of Haikou, the multivoltine tropical population in the south, to ?28.5°C of Gongzhuling, the univoltine temperate population in the northeast of China. In addition, there was more than 1°C difference in SCP between Gongzhuling univoltine and bivoltine populations that were from the same geographic origin. Mean SCPs of the Guangzhou population fluctuated over the year, with significantly lower SCPs in winter than in other seasons, which correlated with a significantly higher proportion of diapausing larvae in winter than in other seasons. Over 41% of overwintering larvae from the northeast population could withstand to be supercooled for a few minutes to the low sub‐zero temperature of ?40°C, but only 6.7% of their southern counterparts did so. The findings from this study suggest that O. furnacalis mostly takes advantage of freeze avoidance as diapausing larvae for overwintering in the southern region, whereas it exhibits freeze tolerance in diapause in the northeastern region.     

水蚤滞育的数量遗传学研究:遗传与环境控制及遗传作用

在单性生殖循环水蚤群体中,滞育卵由有性生殖产生．在一系列实验中用到了不同种群和种类的水蚤,通过这些实验来观察：1)有性生殖和滞育卵复苏的遗传和环境控制;2)生活在相同区域中,但有不同生存微环境的近缘种类的有性生殖的光周期反应;3)在群体遗传结构上有性生殖的遗传效应(基因型均值和生活史性状遗传方差)．结果发现：1)遗传作用和环境作用,以及两者的相互作用都对有性生殖和滞育卵的孵化有显著的影响．GE显著的相互作用对环境中观察到的有性生殖来说,有助于维持其较高的遗传方差;2)在相同区域中,不同生存微环境的近缘种类的有性生殖的光周期反应有所不同．这有助于进一步区别近缘的水蚤种类,这也可能是一个水环境中同素异形的物种形成的例子;3)在有性生殖上,生活史性状平均值和遗传方差变化与前代选择造成的均值和遗传方差相反(遗传滑阻),这会造成暂时的适应不良(遗传滑阻和隐藏的遗传变异的表达),应对它补偿滞育的进化优势．     

Salinity tolerance in <Emphasis Type="Italic">Daphnia magna</Emphasis>: characteristics of genotypes hatching from mixed sediments

The hatching of diapausing eggs is a means of temporal dispersal that can provide populations with genotypes adapted to different environments. In a salinity-variable shallow lake, we predicted that the mixing of different age-classes of eggs in the sediment may yield genotypes with different salinity optima. The alternative would be the absence of local adaptation and the presence of a homogenous population of salt-tolerant genotypes with high phenotypic plasticity. We tested these alternatives by isolating Daphnia magna resting eggs from different sediment depths, exposing them to hatching cues at different salinity levels and measuring the performance of hatched individuals. Results revealed a homogeneous sediment with generally broad-tolerance genotypes and some genotypes with low salt tolerance, which supports the second hypothesis. However, the disturbed character of the sediment hampered historical reconstruction. The absence of local adaptation in the diapausing egg bank may be the result of various scenarios in the response of D. magna populations to severe salinity changes in the lake.