Reproduction strategies of <Emphasis Type="Italic">Daphnia pulicaria</Emphasis> population in a high mountain lake of Southern Spain

For three consecutive years, a population study of Daphnia pulicaria was undertaken in Río Seco Lake, a small high-mountain lake, in order to elucidate the reproductive strategies adopted by Daphnia in this system. Daphnia appears to colonize this lake every spring by hatching from ephippia and reproduce by means of subitaneous (non-diapausing) and ephippial (diapausing) eggs. D. pulicaria in this lake is an obligate parthenogenetic population. There is a short time period for subitaneous egg production and a much longer period for ephippial egg production. The contribution of subitaneous eggs to Daphnia population density and structure appears to be low. Diapause onset showed a high temporal synchronization in the three studied years in Río Seco Lake, and day-length emerged as the main cue triggering diapause onset and the main explanatory factor for the proportion of ephippial females observed. The development and reproduction of D. pulicaria in Río Seco Lake involves taking a gamble on resting forms to guarantee inter-annual Daphnia persistence in the lake, giving priority to investment in future generations. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

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.     

Response of diapausing eggs hatching to changes in temperature and the presence of fish kairomones

It has been hypothesized that the production of diapausing eggs in Daphnia can be induced by fish kairomones. A population of Daphnia could survive severe predation using this predator avoidance strategy. However, in changing environments, diapausing eggs experience various temperature conditions, and hatchlings at emergence may be exposed to the same predation risks as their mothers. Therefore, staying in diapause or an immediate response upon hatching to available environmental information could be important for hatchling survival. For this study, we investigated the impact of water temperature (10, 15, 20, and 25°C) in the presence and absence of fish kairomones (Lepomis macrochirus) on the hatching success of resting eggs (D. galeata). Results show that no diapausing eggs hatched at the lowest temperature (10°C), and the highest hatch percentage occurred at 15°C. Although higher water temperatures reduced hatching success, diapausing eggs hatched more quickly. The number of hatchlings was significantly higher after exposure to fish kairomones, and this was more noticeable at higher temperatures (20 and 25°C). The present results suggest that the diapausing eggs were produced as a predator avoidance strategy in Daphnia; however, the presence of fish works as a positive signal to increase hatchlings when the diapausing stage is terminated.     

Are they still viable? Physical conditions and abundance of <Emphasis Type="Italic">Daphnia pulicaria</Emphasis> resting eggs in sediment cores from lakes in the Tatra Mountains

All species of Daphnia (Cladocera) produce, at some stage in their life cycle, diapausing eggs, which can remain viable for decades or centuries forming a “seed bank” in lake sediments. Because of their often good preservation in lake sediment, they are useful in paleolimnology and microevolutionary studies. The focus of this study was the analysis of cladoceran resting eggs stored in the sediment in order to examine the ephippial eggs bank of Daphnia pulicaria Forbes in six mountain lakes in the High Tatra Mountains, the Western Carpathians (northern Slovakia and southern Poland). Firstly, we analyzed distribution, abundance and physical condition of resting eggs in the sediment for their later used in historical reconstruction of Daphnia populations by genetic methods. To assess changes in the genetic composition of the population through time, we used two microsatellite markers. Although DNA from resting eggs preserved in the High Tatra Mountain lake sediments was extracted by various protocols modified for small amounts of ancient DNA, DNA from eggs was not of sufficient quality for microsatellite analyses. Distribution curves of resting eggs from sediment cores correspond to the environmental changes that have occurred in the High Tatra Mountains area during last two centuries (atmospheric acid deposition, fish introduction) and demonstrate their influence on natural populations. Evaluation of ephippia physical condition (the most common category was empty ephippial covers) suggests that the majority of resting eggs hatched to produce a new generation of Daphnia or may be due to failed deposition of resting eggs by Daphnia to the chitinous case. In conclusion, age, low quantity and poor physical condition of resting eggs from these Tatra lake sediments proved to be unsuitable not just for use in genetic analyses, but also the possibilities of autogenous restoration of Daphnia populations from the resting egg banks in the Tatra sediments are negligible.     

Temperature,recreational fishing and diapause egg connections: dispersal of spiny water fleas (<Emphasis Type="Italic">Bythotrephes longimanus</Emphasis>)

The spiny water flea (Bythotrephes longimanus) is spreading from Great Lakes coastal waters into northern inland lakes within a northern temperature-defined latitudinal band. Colonization of Great Lakes coastal embayments is assisted by winds and seiche surges, yet rapid inland expansion across the northern states comes through an overland process. The lack of invasions at Isle Royale National Park contrasts with rapid expansion on the nearby Keweenaw Peninsula. Both regions have comparable geology, lake density, and fauna, but differ in recreational fishing boat access, visitation, and containment measures. Tail spines protect Bythotrephes against young of the year, but not larger fish, yet the unusual thick-shelled diapausing eggs can pass through fish guts in viable condition. Sediment traps illustrate how fish spread diapausing eggs across lakes in fecal pellets. Trillions of diapausing eggs are produced per year in Lake Michigan and billions per year in Lake Michigamme, a large inland lake. Dispersal by recreational fishing is linked to use of baitfish, diapausing eggs defecated into live wells and bait buckets, and Bythothephes snagged on fishing line, anchor ropes, and minnow seines. Relatively simple measures, such as on-site rinsing of live wells, restricting transfer of certain baitfish species, or holding baitfish for 24 h (defecation period), should greatly reduce dispersal.     

Population differences in the timing of diapause: adaptation in a spatially heterogeneous environment

Summary Populations of the planktonic copepod, Diaptomus sanguineus, live in permanent and temporary freshwater ponds in Rhode Island. All ponds in which they occur become uninhabitable at some time during the year, but the nature and timing of the harsh period varies both spatially and temporally. Females produce discrete clutches either of subitaneous eggs which hatch immediately or of diapausing eggs which hatch the following season. The two egg types show distinct chorion morphologies under transmission electron microscopy. In permanent ponds the copepods start making diapausing eggs in March, one month before rising water temperatures induce planktivorous sunfish to become active. In temporary ponds diapausing eggs are produced, in a complex pattern from May to July, before the water disappears in late summer or early fall.We investigated the spatial scale at which D. sanguineus is adapted to this complex environment. In a reciprocal transfer experiment between temporary and permanent bodies of water, female copepods placed in new ponds made subies of water, female copepods placed in new ponds made subitaneous and diapausing eggs in the same sequence as control females retained in their home ponds. The copepod populations enter diapause at times appropriate for the local habitat conditions they experience, but inappropriate for other, nearby ponds. Transplanted females were unable to sense a change in pond type or to adjust egg production accordingly. We conclude that D. sanguineus populations are adapted to the specific conditions of isolated ponds rather than to a broader geographical region containing several pond types.     

Resilience to changes in lake trophic state: Nutrient allocation into Daphnia resting eggs

During past decades, many lakes underwent drastic human‐caused changes in trophic state with strong implications for population dynamics and food web processes. We investigated the influence of trophic state on nutrient allocation into Daphnia resting eggs. The production of resting eggs is an important survival strategy, allowing Daphnia to cope with unfavorable environmental conditions. Allocation of essential nutrients into resting eggs may crucially influence embryonic development and offspring survival and thus is of great ecological and evolutionary interest. The capacity of Daphnia to adjust the allocation of nutrients into resting eggs may depend on the dietary nutrient supply, which may vary with trophic state‐related changes in the phytoplankton community composition. Resting eggs were isolated from sediment cores taken from Lake Constance, a large prealpine lake with a distinct eutrophication and reoligotrophication history, and analyzed for elemental (carbon, nitrogen, and phosphorus) and biochemical (sterols and fatty acids) nutrients. Carbon allocation into Daphnia resting eggs continuously decreased over time, irrespective of changes in trophic state. The allocation of nitrogen into Daphnia resting eggs followed the changes in trophic state, that is, nitrogen concentrations in resting eggs increased with eutrophication and decreased again with reoligotrophication. The allocation of phosphorus, sterols and long‐chain polyunsaturated fatty acids, such as eicosapentaenoic acid, into Daphnia resting eggs did not change significantly over time. Changes in trophic state strikingly influenced all trophic levels in Lake Constance. However, nutrient allocation into Daphnia resting eggs was mostly resilient to changes in lake trophic state.     

Status of Daphnia Resting Egg Banks in Bohemian Forest Lakes Affected by Acidification

We studied the egg banks of Daphnia gr. longispina in four acidified lakes in the Bohemian Forest (Czech Republic, Europe). Daphnia had become extinct in three of the studied lakes due to anthropogenic acidification and we assessed the possibility of autochthonous recovery of the population after recent amelioration of the lake environment. We determined the ephippia distribution in eleven cores 10–30 cm long, and the state of Daphnia resting eggs in over 13 000 ephippia from eight cores. Apparently well-preserved eggs were used for hatching experiments and for DNA amplification. Vertical profiles of ephippia densities in several cores did not agree with historical data on the Daphnia presence in the lakes, as the sediment had been repeatedly disturbed by human activities in the past. Ephippia are present near the surface of the sediment in all lakes, and they might therefore receive hatching cues. We were not able, however, to prove that viable eggs are present in the ephippia banks of the lakes. The percentage of the well-preserved eggs in all but one core was below 1%. In addition, no egg hatched in the hatching experiments and we were not able to amplify DNA from the preserved eggs of lakes where Daphnia is extinct, although amplification from relatively young eggs from the fourth lake was successful in 60% of the cases. We conclude that the recovery of Daphnia populations in studied lakes from autochthonous sources is unlikely.     

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.     

No Evidence for Adaptive Micro-Evolution to a Decrease in Phosphorus-Loading of a Daphnia Population Inhabiting A Pre-Alpine Lake

Many polluted lakes in Europe are being restored and phosphorus concentrations have dropped dramatically in these lakes. We studied the genetic structure of Daphnia galeata over the past 30 years in Lake Greifensee, Switzerland, a period during which the phosphorus concentration in the lake reduced dramatically. Distinct genotypes of D. galeata were hatched from diapausing eggs extracted from six different time horizons in dated sediments. We compared juvenile growth, size and egg production of D. galeata reared on high-P and low-P algae to investigate whether Daphnia have evolved to grow better on phosphorus-limited algae. Our results indicate that life histories of D. galeata differed significantly between both food types. We also found significant clone effects for size and egg number. But we found no significant interaction between the depth from which the clones were selected and food quality. This means that we found no evidence for adaptive micro-evolution in response to P reduction in the lake. We discuss our results in relation to other studies that found evidence for adaptive micro-evolution in comparable time frames.     

To sink or float: the fate of dormant offspring is determined by maternal behaviour in Daphnia

• 1 As the ephippia (chitinous shells enclosing diapausing eggs) of pelagic crustaceans of the genus Daphnia have been occasionally reported to float at the water surface, we considered that this might be an adaptation promoting their passive dispersal. We investigated the mechanisms by which ephippia appear at the water surface.
• 2 While field surveys revealed that floating Daphnia ephippia are often numerous in various freshwater habitats, laboratory tests showed that newly formed ephippia are not buoyant initially. Once transferred to the surface by whatever means, however, they may remain there due either to surface tension or gas absorption.
• 3 Video recordings showed that all ephippia at the water surface in laboratory vessels were shed there by ephippial females when moulting (despite the attendant risk of exposure to UV radiation). This implies that the moulting behaviour of female Daphnia may determine the fate of their dormant offspring, predetermining whether they remain in the natal environment (when the ephippium is released into the water column) or disperse (when it is deposited at the water surface).
• 4 Our findings reveal a potential mechanism underlying the high dispersal capacity of freshwater cladocerans inhabiting island‐like aquatic habitats.

     

Role of temperature in diapause response to fish kairomones in crustacean Daphnia

The effect of non-lethal thermal conditions on the diapause response to a simulated threat of fish predation was tested in the freshwater planktonic crustacean Daphnia magna.From an early developmental stage, female Daphnia were either exposed or not exposed to fish kairomones that notified of the threat of fish predation at the benign growth temperatures of 18, 22 or 26 °C. The proportion of females switching from the production of subitaneous to diapausing eggs and their rate of reproduction were recorded.Besides the faster development and more intense reproduction observed at higher temperatures, a smaller proportion of the females formed protective, diapausing eggs when exposed to fish kairomones than was the case in colder conditions. The production of diapausing eggs was not observed when the threat of fish predation was absent. These results indicate an interactive effect of fish kairomones and thermal conditions on diapause induction in D. magna.We interpret these findings in the context of strategies for the maximisation of reproductive success. The production of well protected diapausing eggs (which assures low yet stable gains under diverse thermal conditions) may be a more beneficial life history strategy at lower temperatures, where the chances of survival of active individuals until maturation and successful reproduction are low. Higher temperatures permit faster maturation and more intensive reproduction that may surpass numerical losses of active descendants due to predation, making diapause a less rewarding option.     

Fish as vectors in the dispersal of Bythotrephes cederstroemi: diapausing eggs survive passage through the gut

• 1 Bythotrephes cederstroemi (Crustacea: Onychopoda: Cercopagidae) is an introduced invertebrate predator currently spreading through the Laurentian Great Lakes region of North America. We examined a previously unsuspected way in which B. cederstroemi may be dispersed by fish by their consumption of diapausing eggs.
• 2 Ninety‐four percentage of the mature B. cederstroemi diapausing eggs consumed by fish were egested apparently intact. This proportion is considerably above previous estimates for the ephippial eggs of Daphnia. The hatching success of diapausing eggs was compared among four categories: (a) eggs released naturally by B. cederstroemi (control, 73% hatched) (b) eggs released during ‘stressful confinement’ (46% hatched) (c) eggs dissected from dead females (13% hatched) and (d) eggs recovered from faecal pellets following consumption by fish (viable gut passage experiment, 41% hatched).
• 3 Samples of small fish and B. cederstroemi were collected simultaneously. Examination of gut contents revealed that fish contained B. cederstroemi diapausing eggs and that B. cederstroemi bearing resting eggs were consumed selectively over those without eggs. Moreover, fish selected B. cederstroemi bearing mature rather than immature diapausing eggs.
• 4 The fact that diapausing eggs survive gut passage is important for the dispersal of B. cederstroemi. Fish often move between the pelagic and littoral zones of lakes and may thus disperse diapausing eggs widely. Fish may also move between lakes connected by river systems and can be caught and passively dispersed by anglers or piscivorous birds. Our results demonstrate the potential for fish to act as vectors in the spread of B. cederstroemi.

     

Alternative life histories in the water strider Gerris lacustris: time constraint on wing morph and voltinism

The wing dimorphic water strider Gerris lacustris L. (Heteroptera: Gerridae) switches to a bivoltine life cycle under favorable climatic conditions. The switch in voltinism is accompanied by a reduction of wing development in the directly reproducing midsummer generation, while the diapausing generation has a high fraction of long‐winged individuals. We investigated whether the thermal energy (degree‐days) available in natural habitats constrains the combination of developmental pathway and wing morph. Offspring of G. lacustris were reared under quasi‐natural conditions at two temperature regimes to determine the thermal constant k required to complete adult development in either wing morph. The thermal constant for egg‐to‐adult development of the short‐winged morph was about 20% lower than of the long‐winged morph. Based on the results from the outdoor laboratory experiment, we calculated the total degree‐days necessary to complete the possible combinations of wing morph pattern and voltinism. Comparison of these estimates with the thermal energy actually available during the reproductive season of 2004 for various natural habitats (sun‐exposed field ponds and shaded forest ponds) suggests that voltinism as well as wing morph pattern is strongly limited by the number of degree‐days available in these habitats. On forest ponds, only univoltine life cycles were possible, whereas on field ponds temperature allowed bivoltine life cycles. However, only the eggs laid at the very beginning of the season had the potential to accumulate enough degree‐days to complete a bivoltine life cycle with both generations long‐winged. We conclude that thermal energy is the main environmental constraint limiting voltinism of populations in the two habitat types. Furthermore, the available thermal energy also seems to influence the determination of the seasonal wing pattern in G. lacustris.     

Chronic responses of different clones of Daphnia longispina (field and ephippia) to different food levels

Resting eggs are a fundamental reproductive strategy among freshwater cladocerans. Under adverse environmental conditions, whole Daphnia populations can disappear from a lake and a new community will arise from ephippial eggs. Since these new populations are subjected to genetic variation, their responses to environmental stress or contaminants can be different from the “original” population. In the present study, life history responses (reproduction and growth) of Daphnia longispina to different food concentrations was studied. Two Daphnia populations were tested: (a) field clones and (b) ephippial clones. Food (Selenastrum capricornutum) concentration was the stressor tested (absence of food, and low to high food concentrations). The results showed that reproductive responses of D. longispina to the tested food concentrations varied among clones, independently from their origin.     

Starvation tolerance of rotifers produced from parthenogenetic eggs and from diapausing eggs: a life table approach

Monogonont rotifers have to cope with environmental variationby producing diapausing stages. Cyclically, parthenogeneticrotifers produce females by (i) parthenogenetic eggs and (ii)sexually produced eggs. Parthenogenetic eggs hatch shortly afterovoposition, frequently while they are carried by their mothers.Sexual eggs go into dormancy and hatch in the sediment, in responseto specific environmental cues. Therefore, it is expected thatpost-diapause and parthenogenetic offspring would face differentenvironments. Moreover, resource allocation is higher in diapausingeggs than in parthenogenetic eggs. In this study, the responseto starvation of females obtained from parthenogenetic eggsand from diapausing eggs of the rotifer Brachionus plicatiliswas compared. Starved individuals showed unexpectedly long maximumlongevity (12 days). The average lifespan and the lifespan varianceof individuals hatched from parthenogenetic eggs were higherthan those of individuals hatched from diapausing eggs. Contrastingwith a previous experimental finding, our results do not supportthe hypothesis that high resource allocation in diapausing eggscauses high starvation tolerance. Lower survival of individualshatched from diapausing eggs could be caused by the diversionof energy to other functions (i.e. maintenance during diapauseor the hatching event), or the result of the variance in theallocation of resources in parthenogenetic eggs. We hypothesizethat resource allocation in diapausing eggs follows a conservativepattern, while it is more opportunistic in parthenogenetic eggs. This paper was presented at Plankton Symposium III, held atFiguera da Foz, Portugal between 17 and 20 March 2005, underthe auspices of the University of Coimbra and the Universityof Aveiro, and coordinated by Mário Jorge Pereira andUlisses M. Azeiteiro.     

Does the Number of Diapausing Eggs Laid on Bark Affect the Population Dynamics of the Spider Mite Panonychus mori?

We studied the relationship between number of diapausing eggs produced by the spider mite Panonychus mori and the subsequent population trend for a period of 3 years. Panonychus mori showed a single population density peak on its host plant moonseed, Cocculus trilobus. The position and height of this peak were correlated with the density of diapausing eggs around the moonseed leaf buds produced the winter before. In 1994 the density of diapausing eggs measured in February was 4.3/bud, which was 6–14 times higher than the density of diapausing eggs for the same period in 1995 (0.3/bud) and 1996 (0.7/bud). The subsequent population density peak in 1994 occurred in mid June and was about 2.5 times higher than the peaks in 1995 and 1996, which both occurred early September. Thus, the present study showed a positive correlation between the density of diapausing eggs on the host plant and the start and the extend of the population increase the next growing season. Predators associated with the spider mite population were phytoseiid mites, especially Amblyseius eharai was well synchronized with the spider mite density in 1994. Field observations revealed that P. mori produced diapausing eggs in response to short photoperiod in early October each year, which corresponded with the timing predicted by the critical photoperiod around 13 h at 18°C, as assessed in laboratory trials. Diapause ended by early April when egg hatchability attained about 50% and eggs took 9 days to hatch at 25°C and a 16L:8D photoperiod. Hatching in early April was twice faster than in late February.     

Depth selection and life history strategies as mutually exclusive responses to risk of fish predation in <Emphasis Type="Italic">Daphnia</Emphasis>

We tested if pelagic crustaceans of the genus Daphnia use different anti-predator defences in environmental conditions that do or do not offer deep refuge from planktivorous fish. We kept Daphnia catawba in two series of 9-m deep enclosures with and without caged cyprinid fish Phoxinus eos. In one series of enclosures, Daphnia could select its depth of residence and hide in deep dark water layers to avoid anticipated fish predation, while in another series of enclosures, a plankton net barrier fixed at 2-m depth forced them to stay in subsurface zone exposed to fish kairomones. We compared depth residence and migratory behaviour strategies with life history strategies (body size and size at first reproduction, diapause induction) in Daphnia exposed or not to fish kairomones with or without deep refuge. In deep enclosures with fish, Daphnia spent daytime hours in deep dark layers while at night, they resided closer to the water surface. Yet, no change in life history parameters of migrating individuals was observed compared to the fish-free conditions. In enclosures with fish, where the net barrier forced Daphnia to reside in subsurface zone, they produced smaller offspring, matured at smaller size and achieved lower maximum body length compared to the fish-free conditions. However, they did not produce diapausing eggs. Our experimental study supports the hypothesis that diel vertical migration behaviour with daytime residence in deep, dark water are the preferred antipredator strategy chosen by Daphnia facing anticipated fish predation over life history changes such as reduced size and low growth rate which are used when dark deep refuge is not present or accessible.     

Identifying century-old long-spined <Emphasis Type="Italic">Daphnia</Emphasis>: species replacement in a mountain lake characterised by paleogenetic methods

Mountain lakes often harbour morphologically or genetically unique populations of zooplankton species, including cladocerans. Daphnia lacustris Sars, predominantly found in Fennoscandia but also known from two Central European lakes in the Tatra Mountains, is one of such taxa. This Daphnia species often forms morphotypes with extremely long tailspines. Historical literature from a century ago documented similar morphs from another lake in the Tatra mountain range, presently inhabited by the phenotypically very different D. galeata. Using a paleogenetic approach (partial sequencing of the mitochondrial gene for 12S rRNA from preserved ephippial eggs in the lake sediment), we tested the hypothesis that Daphnia species composition changed in the lake due to anthropogenic disturbances, and that long-spined morphs were actually another relict population of currently extinct D. lacustris. Ephippia with extremely long spines were successfully retrieved from sediment cores. Despite being morphologically very well preserved, intact eggs were found in less than 2% of analysed ephippia. Genetic analyses, benefiting in most cases from amplification of short 12S fragments using internal primers, proved that long-spined ephippia belonged to D. longispina, which apparently coexisted with D. galeata in the mid-twentieth century. Our results confirm that paleogenetic methods are useful for studying the recent population structures of zooplankton species forming dormant egg banks but lacking reliably identifiable remains in sediments, and show that the extreme development of tailspines in mountain-lake Daphnia is associated with as-yet unclear environmental factors rather than taxonomic status.     

Sensitivity of Daphnia species to phosphorus-deficient diets

The life history of freshwater cladocerans such as Daphnia spp. is strongly affected by their environment. Factors such as temperature, food quantity and even the presence or absence of predators influence growth, reproduction and morphology of individuals. Recently, it has also become clear that the quality of the food can affect various life history traits of Daphnia. More specifically, the effect of the elemental composition of algae, expressed as the C:P ratio, has been studied intensively. Daphnia species differ in their response to differences in the C:P ratio of their food. Until now, it has been unclear whether these species differences are driven by phylogenetic constraints or by adaptation to particular environmental conditions. Here we present laboratory experiments with 12 Daphnia species from three different subgenera originating from a broad range of habitats. We compared somatic growth rates and sensitivity to variation in the nutrient stoichiometry of the food with habitat parameters, taking into account the phylogenetic history of the species. No associations between fitness and habitat parameters were detected. However, we found a trade-off between sensitivity to P-deficient diets and the maximum growth rate on a P-sufficient diet. In several cases, this trade-off helps to explain the association between species distribution and habitat parameters. We observed no correlation of the sensitivity to P limitation with the phylogenetic history of the genus Daphnia. Thus, we conclude that the differential responses among Daphnia species to variation in P content in food were driven mainly by adaptations to their local habitats, and are not constrained by deep evolutionary patterns.