Exploitation of a deep-water algal maximum by Daphnia: a stable-isotope tracer study

The exploitation of a deep algal maximum by Daphnia in the absence of fish predation was studied in large indoor mesocosms. Facing the dilemma of low food but high temperature in the epilimnion vs. high food but low temperature in the hypolimnion, Daphnia distribute above and below the thermocline in order to optimise their fitness. Labelling hypolimnetic algae with ¹⁵N revealed that the vertical distribution of Daphnia is dynamic, i.e., all individuals traverse the thermocline and allocate a certain proportion of their time to feeding in the cold water. The overall energy gain from the deep-water algal maximum is lower than from the same algal concentration in the epilimnion due to the low temperature and the limited time an individual spends in the hypolimnion. The results provide mechanistic support for the hypothesis that Daphnia chose their habitat according to an Ideal Free Distribution with Costs model.     

Impact of food concentration on diel vertical migration behaviour of <Emphasis Type="Italic">Daphnia pulex</Emphasis> under fish predation risk

Vertical migration of Daphnia represents the best-studied predator-avoidance behaviour known; yet the mechanisms underlying the choice to migrate require further investigation to understand the role of environmental context. To investigate the optimal habitat choice of Daphnia under fish predation pressure, first, we selected the individuals exhibiting strong migration behaviour. The animals collected from the hypolimnion during the daytime were significantly larger, being more conspicuous, and in turn performed stronger diel vertical migration (DVM) when exposed to fish cue. We called them strong migrants. Second, we provided the strong migrant D. pulex with food at high and intermediate (1 and 0.4 mg C l⁻¹, respectively) levels, which were well above the incipient limiting level and of high quality. They traded the benefits of staying in the warm water layer and moved down to the cold water in response to fish cue indicating fish predation. The availability of food allowed the animals to stay in the cold hypolimnion. However, at the low food level (0.1 mg C l⁻¹), which is an additional constraint on fitness, Daphnia moved away from the cold hypolimnion. Poor food condition resulted in strong migrant Daphnia to cease migration and remain in the upper warmer water layer. Although temperature is known to be a more important cost factor of DVM than food, our results clearly show that this is only true as long as food is available. It becomes clear that food availability is controlling the direction of vertical positioning when daphnids experience a dilemma between optimising temperature and food condition while being exposed to fish cue. Then they overlook the predation risk. Thus, the optimal habitat choice of Daphnia appears to be a function of several variables including temperature, food levels and fish predation. Handling editor: S. I. Dodson     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Significance of a low oxygen layer for a Daphnia population in Lake Yunoko,Japan

Population dynamics and vertical migration of Daphnia longispina in Lake Yunoko were studied. The Daphnia population was small in spring and early summer, probably because of high predation pressure by fish. The population grew in midsummer, when thermal stratification developed and the dissolved oxygen became very low in the deeper layer of the hypolimnion. In this season, adults of D. longispina concentrated in the daytime near the lake bottom, where fish were absent because of the anoxic conditions, but ascended at night to the upper layer of the hypolimnion, where food was most abundant. The low oxygen layer near the bottom kept out the predators and protected Daphnia from predation, and consequently contributed to the built-up of its population. However, the low oxygen layer was unfavorable for reproduction of Daphnia, as reflected in the low egg ratio and high percentage of males in the population. The population decreased in the fall, when thermal stratification disappeared and predation pressure seemed to increase.     

Depth distribution of Daphnia in response to a deep-water algal maximum: the effect of body size and temperature gradient

1. In the absence of fish predation, Daphnia exploiting a deep‐water algal maximum are faced with a trade‐off. They can either dwell in the epilimnion where development in the warm water is fast, but food shortage causes low egg production, or in the hypolimnion, where food availability is high but development is slow because of low temperatures. 2. We tested the hypotheses that (i) depth distributions of various ontogenetic stages (size classes and egg‐bearing females) differ because daphnids react to light with size‐specific diel vertical migration (DVM) even in the absence of fish (residual predator avoidance hypothesis) and (ii) differently sized daphnids select different depths because the relative importance of temperature and food varies for ontogenetic stages (physiological hypothesis). We used large indoor mesocosms (Plankton Towers) to test these hypotheses experimentally. 3. Temperature was the strongest factor governing the distribution, with larger proportions of the population dwelling in the food‐rich hypolimnion if the temperature gradient was shallow. There were small but significant differences between ontogenetic stages during the day, but not at night. This suggested the existence of a ‘residual’ effect of light on depth distribution in the absence of a fish cue. 4. Although large individuals exhibited greater amplitude of DVM, the physiological hypothesis had to be rejected. A stage‐specific physiological effect is unlikely to be directly triggered by light, hence vertical movement of the individuals should not be synchronised. Rather, being forced into deeper layers by the residual light response during the day, large and egg‐bearing females experience a lower average temperature during day than juveniles. They probably compensate for this by spending longer time periods in warm waters at night.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Small fish use the hypoxic pelagic zone as a refuge from predators

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Food quality influences habitat selection in Daphnia

1. The vertical distribution of Daphnia in stratified lakes strongly depends on the depth profiles of temperature and food resources. However, ecological requirements for these factors are slightly different for juvenile and adult Daphnia. 2. Here, I investigated whether food quality influences the habitat selection of Daphnia pulicaria at night and whether the habitat selection of juvenile and adult D. pulicaria is different. Daphnia were allowed to choose their optimal habitat in large, stratified water columns (plankton towers, Plön) that held either the green alga Scenedesmus obliquus (high quality) in the cold hypolimnion (Hypo‐treatment) or S. obliquus in the warm epi‐ and cold hypolimnion (SCEN‐treatment) or the non‐toxic cyanobacterium Synechococcus elongatus (low quality) in the warm epilimnion and S. obliquus in the cold hypolimnion (SYN treatment). 3. When food (S. obliquus) was present only in the hypolimnion (Hypo‐treatment), juveniles and adults distributed similarly in the water column and spent most of their time in the interface between the warm and the food rich layer. 4. When food was present in the epilimnion and hypolimnion (SCEN‐ and SYN‐treatments), juvenile and adult D. pulicaria moved into the warm and now also food‐rich epilimnion, however, the magnitude of this shift depended on the food type and age class of Daphnia. Adult and juvenile D. pulicaria spent most of their time in the epilimnion when food there was of a high quality (S. obliquus; SCEN‐treatment). However, compared to the juveniles, adult Daphnia spent significantly more time in the colder hypolimnion when epilimnetic food was of a low quality (S. elongatus; SYN‐treament). 5. Therefore, habitat selection of adult D. pulicaria was affected by food quality whereas the habitat selection of juveniles was not. 6. Additional growth and reproduction experiments show that the food quality is likely to be responsible for the different habitat selection of juveniles and adults in the SYN‐treatment. 7. In conclusion, my experiments show that D. pulicaria behaviourally reacts to the quality of its food source.     

Fitness optimization of Daphnia in a trade-off between food and temperature

In thermally stratified lakes with a deep chlorophyll maximum (DCM), Daphnia face a trade-off between food availability and optimum development temperatures. We hypothesize that Daphnia optimize their fitness by allocating the time spent in the different vertical habitats depending on the distribution of algal resources and the temperature gradient. We used the plankton towers (large indoor mesocosms) to study the vertical distribution of a population of Daphnia hyalina×galeata in three different temperature gradients with a DCM. Additionally, we determined the fitness of Daphnia in the epilimnion and hypolimnion by transferring water from these layers into flow-through systems where we raised Daphnia and assessed their juvenile growth rate as a measure of fitness. The fitness distribution was correlated with the vertical distribution. The vertical distribution most likely reflected the proportions of time Daphnia allocated to dwelling in the two vertical habitats.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Sediments,not plants,offer the preferred refuge for Daphnia against fish predation in Mediterranean shallow lakes: an experimental demonstration

1. Different behavioural responses of planktonic animals to their main predators, fish, have been reported from shallow lakes. In north temperate lakes, large‐bodied zooplankton may seek refuge from predation among macrophytes, whereas in subtropical lakes, avoidance of macrophytes has been observed. The prevalent behaviour probably depends on the characteristics of the fish community, which in Mediterranean lakes is typically dispersed in both the open water zone and in the littoral, as in temperate lakes, and is dominated by small size classes, as in subtropical lakes. 2. We performed ‘habitat choice’ experiments to test the response of Daphnia magna to predation cues at both the horizontal and vertical level by mimicking a ‘shallow littoral’ zone with plants and a ‘deeper pelagic’ zone with sediments. 3. Initial separate response experiments showed that natural plants, artificial plants and predation cues all repelled D. magna in the absence of other stimuli, while sediments alone did not trigger any significant response by D. magna. 4. The habitat choice experiments showed that, in the presence of predation cues and absence of plants, Daphnia moved towards areas with sediment. In the presence of both plants and sediments, Daphnia moved away from the plants towards the sediments under both shallow and deep water treatment conditions. 5. Based on these results, we suggest that Daphnia in Mediterranean shallow lakes avoid submerged macrophytes and instead prefer to hide near the sediment when exposed to predation risk, as also observed in subtropical shallow lakes. This pattern is not likely to change with water level alterations, a common feature of lakes in the region, even if the effectiveness of the refuge may be reduced.     

Migrations of haemoglobin-rich Daphnia longispina in a small,steeply stratified,humic lake with an anoxic hypolimnion

Migrations of Daphnia longispina were studied in a small humic lake with an exceptionally shallow oxic epilimnion. Horizontal distributions showed clear avoidance of the shoreline, which might be explained by the lower density of predators (Chaoborus sp. and Notonecta sp.) in the central parts of the lake. In early summer all size classes of D. longispina exhibited upward nocturnal vertical migration, descending to the upper hypolimnion in daytime. Later in summer, when the nocturnally migrating Chaoborus sp. had grown large enough to graze on small Daphnia, the latter seemed to shift towards twilight migration. However, large Daphnia individuals showed no synchronized migration; rather their bimodal vertical distributions suggested asynchronous vertical migration. Large individuals showed a particular tendency to concentrate near to the oxycline, close to the dense phytoplankton and bacteria populations in the upper part of the anoxic hypolimnion. According to vertical trap experiments, large D. longispina visited the anoxic hypolimnion and might harvest its abundant food resources. The high haemoglobin content of large individuals seems a specific adaptation to allow access to low oxygen water and hence to maximize grazing potential, in both epi- and hypolimnion, and minimize predation pressure. By staying predominantly in cooler water near the oxycline, Daphnia might also minimize its energy consumption to adjust to low food availability while sustaining a sufficiently high population density to exploit those unpredictable short periods with abundant food which are common in small headwater lakes. It is suggested that migrations of zooplankton are a complex behavioural adaptation which may not be explained by any single factor. In humic lakes with shallow stratification, vertical migrations seem to offer particularly high potential advantages, because of the short distances between dramatically different environments in the water column. In further studies more emphasis should be placed on migrations of individuals rather than populations, and migrations should be considered as a dynamic part of the structure and function of the whole planktonic ecosystem.     

What goes down must come up: symmetry in light-induced migration behaviour of Daphnia

During a short period of the year, Daphnia may perform a phenotypically induced diel vertical migration. For this to happen, light-induced swimming reactions must be enhanced both at dawn and at dusk. Enhanced swimming in response to light intensity increase can be elicited by fish-associated kairomone in the laboratory, if food is sufficiently available. However, during the light change at dusk the Daphnia are still in the hypolimnion, where no fish kairomone is present and both temperature and food availability is low. Still, what goes down must come up. This raises questions about how Daphnia tunes its light-induced swimming behaviour to prevailing conditions such that a normal diel vertical migration can be performed. We investigated the symmetry in behavioural mechanism underlying these diel vertical migrations in the hybrid Daphnia galeata × hyalina (Cladocera; Crustacea), with special interest for the environmental cues that are known to affect swimming in response to light increase. That is, we tested whether fish- associated kairomone, food availability, and temperature affected both swimming in response to light intensity increase and decrease similarly. We quantified swimming behaviour during a sequentially increased rate of light change. Vertical displacement velocity was measured and proved to be linearly related to the rate of the light change. The slope (PC) of the function depends on the value of the factors kairomone concentration, food availability, and temperature. The changes of the PC with kairomone concentration and with temperature were similar both at light intensity increases and decreases. The PC also increased with food concentration, although during light increases in a different way from during light intensity decreases. Low food availability inhibited swimming in response to light intensity increase, but enhanced swimming in response to light intensity decrease. Hence, ascent from the deep water layers with low food concentration at dusk is facilitated. These causal relations are part of a proximate decision-making mechanism which may help the individual Daphnia to tune migration to predation intensity and food availability.     

Distribution of Daphnia in a trade-off between food and temperature: individual habitat choice and time allocation

1. In a thermally stratified water column with a deep‐water algal maximum, Daphnia face a trade‐off between food (high fecundity) and temperature (fast development). Recent studies showed that Daphnia populations move up and down the entire water column to take advantage of both, but the proportion of time allocated by individuals to the epilimnion, metalimnion and hypolimnion with their specific food and temperature conditions is not yet known. 2. In a system of 1 m deep, vertical perspex tubes, I established three temperature gradients with 2, 5 and 10 °C differences between the surface (epilimnion) and the bottom layer (hypolimnion). Algae were added to the hypolimnion to simulate a deep‐water algal maximum. 3. The migration behaviour of individual neonate and egg‐bearing Daphnia hyalina × galeata was monitored in order to measure the proportions of time the individuals allocated to the different vertical habitats and to assess the frequency of their shifts between epilimnion and hypolimnion. 4. Neonates stayed continuously at the surface, taking advantage of the higher temperature, possibly because feeding was less important for them because of egg yolk reserves. In contrast, egg‐bearing females spent more time feeding in the hypolimnion when the temperature gradient was weak, but also migrated into the epilimnion to take advantage of the higher temperature. In the steepest temperature gradient, the egg‐bearing females either shifted between epilimnion and hypolimnion, or dwelled constantly in the metalimnion with intermediate conditions.     

Enhanced growth at low population density in Daphnia: the absence of crowding effects or relief from visual predation?

1. It has been suggested that chemical information from crowded populations of an animal such as Daphnia carries a cue indicating imminent food limitation, and we suggest that in the presence of fish kairomones, it may also convey a hint of the need to enhance antipredation defences. 2. We performed two‐factorial experiments with Daphnia grown in flow‐through plankton chambers in medium containing high levels of Scenedesmus food plus chemical information on either low or high population density levels and in the presence or absence of fish chemical cues (kairomones) and recorded (i) the effects on Daphnia growth rate and reproduction, and (ii) the effects on Daphnia depth selection. Further depth‐selection experiments were performed to test the reaction of Daphnia to crowding information at different Daphnia concentrations and to test its effect on daytime and night‐time depth selection by different Daphnia instars in the presence of kairomones. 3. The effects of crowding information alone (in the absence of kairomones) were weak and were not significantly strengthened by the addition of kairomones. The effects of kairomones alone (in the absence of crowding information) were much stronger and were increased by the presence of crowding chemicals: Daphnia selected greater depths in daylight (the later the instar and the larger its body size, the greater the depth), their body growth was slower and daily reproductive investment reduced, compared with Daphnia grown in the absence of crowding information. This suggested that crowding chemicals carry a cue indicating the need to invest more into antipredation defences. 4. The adaptive significance of these effects was confirmed by the differential vulnerability to predation of the Daphnia when offered as prey to live roach after being grown for 6 days either in the presence (higher vulnerability) or in the absence (lower vulnerability) of information on high density. 5. The strong interaction between crowding information and fish kairomones may be explained either as the reaction to a cue indicating impending food stress or as the reaction to a signal of increased predation risk. While the former scenario is already known from crowding studies, the latter is a novel idea that stems from the old concept of ‘low‐density anti‐predation refuge’. The two scenarios are not mutually exclusive: each stems from the need to invest in survival rather than in growth and reproduction [Corrections were made to this paragraph after first online publication on 4 April 2012].     

Trade-offs in the vertical distribution of zooplankton: ideal free distribution with costs?

Zooplankton vertical migratory patterns are a classic example of optimal habitat choice. We hypothesize that zooplankton distribute themselves vertically in the water column according to an ideal free distribution (IFD) with costs such as to optimize their fitness. In lakes with a deep-water chlorophyll maximum, zooplankton are faced with a trade-off, either experiencing high food (high reproductive potential) but low temperature (slow development) in the hypolimnion or high temperature and low food in the epilimnion. Thus, in the absence of fish predation (e.g. at night) they should allocate the time spent in the different habitats according to fitness gain dependent on the temperature gradient and distribution of food. We tested this hypothesis with a Daphnia hyalina x galeata clone in large indoor columns (Plön Plankton Towers) and with a dynamic energy budget model. In the tower experiments, we simulated a deep-water algal maximum below the thermocline with epilimnetic/hypolimnetic temperature differences of 2, 5 and 10 degrees C. Experimental data supported the model. We found a significantly larger proportion of daphniids in the hypolimnion when the temperature difference was smaller. Our results are consistent with the concept of IFD with costs originally developed for stream fishes. This concept can be applied to predict the vertical distribution of zooplankton in habitats where fish predation is of minor importance.     

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.     

Causes of Daphnia midsummer decline in two deep meromictic subalpine lakes

1. Daphnia are key organisms in pelagic food webs, acting as a food resource for fish and predatory zooplankton and regulating phytoplankton through grazing. Its population dynamic follows regular seasonal patterns, with spring peaks followed by summer population declines (midsummer declines, MSDs). Midsummer declines show high inter-annual variation, which has been attributed to different causes. However, the mechanisms controlling the MSD remain poorly understood, especially in deep stratified lakes.
2. We tried to disentangle the factors causing Daphnia MSDs in Lake Lugano and Lake Iseo (in Switzerland and Italy), two deep peri-alpine lakes with similar trophic status and vertical mixing dynamics, characterised by phosphorus accumulation in the hypolimnion and variable mixing during late-winter turnovers.
3. Specifically, we assessed the effects of three different hypothetical pathways according to which: (1) winter air temperature controls MSDs by influencing mixing depth during turnovers and epilimnetic phosphorus replenishment; (2) vernal air temperature influences MSD by accelerating the timing of spring population peak; and (3) summer temperature influences MSDs by increasing fish predation. We assessed the relative strength of these pathways using structural equation modelling on long-term datasets for the two lakes (29 years for Lake Lugano and 19 years for Lake Iseo).
4. Between the hypothesised pathways, the one driven by winter air temperature (through P replenishment) influenced Daphnia abundance in spring in both lakes, but the effects propagated to summer Daphnia abundance only in Lake Lugano. Additionally, summer Daphnia abundance was influenced by the summer air temperature through a positive (although weak) effect. By comparison, vernal air temperature had no detectable effects on summer Daphnia abundance.
5. The results revealed marked differences between the meromictic study lakes and the shallow hypertrophic water bodies that were the focus of previous research on Daphnia MSD, and also between the two study lakes. The influence of epilimnetic P replenishment on the summer Daphnia abundance in Lake Lugano, which was recovering from past eutrophication, may have reflected the greater susceptibility of deep, stratified lakes to P depletion after spring compared to shallow hypertrophic lakes or reservoirs. This effect might not have been detected in Lake Iseo because P was more consistently depleted during the study period (i.e. variance in the predictor was too low to detect an effect).
6. This study highlighted the complexity of the effects of climate variability on Daphnia MSD in deep lakes, showing that the responses can differ even between two neighbouring lakes with similar vertical mixing dynamics and trophic status. At the same time, the results suggest that future increases in winter air temperature, caused by global warming, may cause critically low densities of Daphnia during spring and summer and compromise the ability of zooplankton to control phytoplankton biomass.

     

Depth-selection behavior and longevity in Daphnia: an evolutionary test for the predation-avoidance hypothesis

The classic evolutionary theory of senescence predicts that long lifespan evolves under low risk of extrinsic mortality. As lakes present planktonic animals with vertical gradients of mortality risk associated with fish predation, we expected the individual lifespan of Daphnia of the “hypolymnetic” clones to be longer than that of “epilimnetic” Daphnia. In order to test this prediction, we performed a laboratory study on 14 clones from the D. longispina species complex, taken during the daytime either from epilimnion or hypolimnion of three mesotrophic lakes. “Epilimnetic” Daphnia started reproduction earlier, aged faster, and lived shorter than their “hypolimnetic” conspecifics. Our results indirectly support the predation-avoidance hypothesis as the ultimate explanation for depth-selection behavior in Daphnia.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.