Diel vertical migration of zooplankton: optimum migrating schedule based on energy accumulation

Zooplankton perform diel vertical migration (DVM) to avoid predators at the upper water layer, but often stay in the upper water layer throughout the day seeking food in spite of the presence of predators. This difference in migrating behavior has been explained by differences in environmental conditions or genetic differences. We examined theoretically how nutritious conditions of zooplankton individuals relate to determining different migrating behavior. A simple optimization model, maximizing the population growth rate, demonstrates that zooplankton individuals change their migrating behavior depending on the amount of accumulated energy. Such energy accumulation and its investment in reproduction are repeated every reproductive cycle. Therefore, unless the reproductive cycle is synchronized among individuals, different migrating behaviors will be observed within a population even if no genetic differences exist. Our model demonstrates that such coexistence of the two migrating behaviors is possible in natural Daphnia populations, and suggests that internal conditions of zooplankton individuals may be important as a factor for determining migrating behavior of zooplankton. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Vertical distributions of zooplankton and population dynamics of Daphnia in a meromictic lake

Vertical distributions of zooplankton were studied in relation to profiles of temperature, oxygen and chlorophyll a in Roi Lake, a small meromictic lake in central Alberta. Zooplankton were distributed fairly evenly through the oxygenated part of the water column in early summer, but a gradual descent of several species became evident in June. The vertical distribution of chlorophyll was dominated by a huge peak at the 8- to 9-m-deep chemocline. the location of a plate of photosynthetic sulfur bacteria. Ambient concentration of chlorophyll was a poor predictor of the numbers of zooplankton and the fecundity of Daphnia pulicaria at different depths, and per capita birth rates of Daphnia were usually highest in the surface waters. The reproductively disadvantageous restriction of daphnids to deep water by late summer and their catastrophic decline in the face of high ambient concentrations of chlorophyll suggest that factors other than temperature and food supply are important in influencing the dynamics and distribution of zooplankton in this lake.     

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.

     

The impact of diel vertical migration of <Emphasis Type="Italic">Daphnia</Emphasis> on phytoplankton dynamics

Diel vertical migration (DVM) of large zooplankton is a very common phenomenon in the pelagic zone of lakes and oceans. Although the underlying mechanisms of DVM are well understood, we lack experimental studies on the consequences of this behaviour for the zooplankton’s food resource—the phytoplankton. As large zooplankton species or individuals migrate downwards into lower and darker water strata by day and upwards into surface layers by night, a huge amount of herbivorous biomass moves through the water column twice a day. This migration must have profound consequences for the phytoplankton. It is generally assumed that migration supports an enhanced phytoplankton biomass and a change in the composition of the phytoplankton community towards smaller, edible algae in the epilimnion of a lake. We tested this assumption for the first time in field experiments by comparing phytoplankton biomass and community assemblage in mesocosms with and without artificially migrating natural stocks of Daphnia hyalina. We show that DVM can enhance phytoplankton biomass in the epilimnion and that it has a strong impact on the composition of a phytoplankton community leading to an advantage for small, edible algae. Our results support the idea that DVM of Daphnia can have strong effects on phytoplankton dynamics in a lake.     

Diurnal vertical migration of Daphnia hyalina Leydig, 1860 (Crustacea: Cladocera) in Lake Bled (Slovenia) in relation to temperature and predation

Diurnal vertical migration (DVM) of Daphnia hyalina in Lake Bled was most intense during summer stratification. The extent of DVM varied with the size of the animal and its reproductive state. Migration distances were shortest in immature specimens and longest in ovigerous females. During daytime, ovigerous females stayed deeper in the water column than females without ova or immatures. The daytime temperature of water at the median depth of the ovigerous females did not exceed 10 °C, even in the warmest season. At night they migrated upward to an environment which was warmer by as much as 9 °C.Laboratory observations indicate that specimen's size and water temperature determine the velocity of passive sinking, such that morning descent of the different groups of Daphnia can be explained by passive sinking alone.Our hypothesis is that the distribution of different groups of D. hyalina in Lake Bled is influenced by two types of predators: fish (Perca fluviatilis L. and Rutilus rutilus (L.)) and larvae of Chaoborus flavicans (Meig.), the latter appearing in the epilimnion during the night. Fish predation has a key-role at the beginning of thermal stratification. Supposing that in spring the gene pool of Daphnia consists of a mix of different genotypes, distributed at different depths during the day, fish predation combined with a presence of fish chemicals favored genotypes with a lower day-depth during the spring/summer period.     

Diel Vertical Migration of zooplankton in a eutrophic bay of Lake Victoria

We studied the Diel Vertical Migration (DVM) of several zooplankton taxa and an important zooplanktivore Rastrineobola argentea in a eutrophic bay of Lake Victoria for a total of 3 months during wet and dry seasons. Zooplankton were sampled twice a month at full moon and new moon. The zooplankton community of this lake was numerically dominated by cyclopoid copepods (>80%) of which 80% were juveniles. The composition of zooplankton was not significantly different (P > 0.05) for the different sampling months (November 2000, March 2001 and July 2001), but total zooplankton abundance was significantly lower (P < 0.05) in July (dry season) than during March or November (wet seasons). DVM behaviour was observed in some of the zooplankton groups including Tropocyclops spp., Thermocyclops spp., Thermodiaptomus galeboides and Diaphanosoma excisum. In these groups, we also observed that they were significantly (P < 0.05) more abundant at full moon than at new moon in some months, a pattern not seen in non-DVM groups. The amplitude of migration tended to be higher at new moon than full moon for all DVM zooplankton but not significantly so (P > 0.05). The zooplanktivore R. argentea exhibits typical DVM behaviour, perhaps to avoid visual predation itself, but it also increases the predation pressure on zooplankton in deeper waters. Feeding times for R. argentea peak just after dusk and dawn. For the copepod calanoid T. galeboides, there was a strong relationship (P < 0.05) between the amplitude of migration and midday (12:00) water transparency, suggesting that these zooplankton may have descended deeper on days with higher water transparency. In the absence of temperature and food gradients but with predation pressure from a migrating zooplanktivore, the depth of descent of zooplankton may be well associated with water transparency.     

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.     

A comparison of the trophic transfer of fatty acids in freshwater plankton by cladocerans and calanoid copepods

1. Analyses of zooplankton fatty acid (FA) composition in laboratory experiments and samples collected from lakes in New Zealand spanning a wide gradient of productivity were used to assess the extent to which FAs might infer their diet. We used the cladocerans, Daphnia and Ceriodaphnia, and the calanoid copepod, Boeckella, as test organisms, and monocultures of cryptophytes, chlorophytes and cyanobacteria as food. Based on reproductive success, cryptophytes were the highest food quality, chlorophytes were intermediate and cyanobacteria the poorest. 2. Several FA groups were highly correlated between zooplankton and their diets. They were monounsaturated fatty acids (MUFAs), and ω3 and ω6 polyunsaturated fatty acids (PUFAs) for cladocerans, and saturated fatty acids (SAFAs) and ω3 PUFAs for copepods. Several FAs varied significantly less in the zooplankton than in their monoculture diets, e.g. MUFAs in Daphnia, and ω3 and ω6 PUFAs in Ceriodaphnia, despite clear dietary dependency for these FAs. 3. Zooplankton collected from lakes in New Zealand had more eicosapentaenoic acid (EPA) (Daphnia), more highly unsaturated ω3 and ω6 FAs (C₂₀, C₂₂; Daphnia, Ceriodaphnia, Boeckella) and less ω3 C₁₈ PUFAs (Daphnia, Ceriodaphnia, Boeckella) and ω6 C₁₈ PUFAs (Daphnia, Ceriodaphnia) than measured in the same species reared on phytoplankton in the laboratory. 4. Analyses of FA composition of seston and freshwater zooplankton globally showed that, in general, zooplankton had a significantly higher proportion of arachidonic acid and EPA than seston, and copepods also had a higher percentage of docosahexaenoic acid than seston. 5. These results suggest that zooplankton selectively incorporate the most physiologically important FAs. This could be a consequence of preferential assimilation, selective feeding on more nutritious cells or locating and feeding within higher food quality food patches.     

Morphological analyses allow to separate <Emphasis Type="Italic">Branchipus</Emphasis> species (Branchiopoda,Anostraca) from different geographic regions

Daphnia perform diel vertical migration (DVM), a predator-avoidance strategy to migrate towards deeper and colder layers in the water column in the morning and movement to the algae-rich surface layers in the evening. However, individuals performing DVM incur several trade-offs since they might suffer from resource limitation and a slower instantaneous birth rate in deeper depths. DVM patterns may be modified by abiotic factors such as temperature, food concentration, or pH and vary among different Daphnia species and genotypes. Furthermore, Daphnia host a variety of microparasites that might pose an additional factor influencing DVM behaviour. For infected individuals, migration into cooler temperature layers might slow down parasite growth. Moreover, parasites can increase opacity of their hosts. Non-migrating individuals might then be selectively purged from the upper layers by visually hunting predators. With these premises we asked, whether epidemics of the ichthyosporean parasite Caullerya mesnili affect or are affected by the DVM behaviour of Daphnia in Lake Greifensee, Switzerland by analysing the vertical distribution of Daphnia during day and night on two dates. Furthermore, we were interested whether a potential interaction depends on host genotype. We therefore studied the genotypic composition of the integrated population in regular sampling intervals over the course of one year and on a fine-grained vertical resolution during the Caullerya epidemic in late summer. Since Caullerya-infected Daphnia migrated equally well as uninfected ones, the findings of this study suggest that Caullerya epidemics neither affected nor were affected by the DVM behaviour of Daphnia. We observed clonal succession in the lake but could not link this succession to the Caullerya epidemic; all except one of the common multilocus genotypes were under-infected. In addition, outbreak and course of this Caullerya epidemic seemed to rely mainly on environmental cues. Because this first study only provides a snapshot of time, we hope that further studies will be done to verify our results.     

Can overwintering versus diapausing strategy in <Emphasis Type="Italic">Daphnia</Emphasis> determine match–mismatch events in zooplankton–algae interactions?

Mismatches between predator and prey due to climate change have now been documented for a number of systems. Ultimately, a mismatch may have far-reaching consequences for ecosystem functioning as decoupling of trophic relationships results in trophic cascades. Here, we examine the potential for climate change induced mismatches between zooplankton and algae during spring succession, with a focus on Daphnia and its algal food. Whereas the development of an overwintering population of daphnids may parallel shifts in phytoplankton phenology due to climate warming, changes in the photoperiod–temperature interaction may cause the emerging population of daphnids to hatch too late and mismatch their phytoplankton prey. A decoupling of the trophic relationship between the keystone herbivore Daphnia and its algal prey can result in the absence of a spring clear water phase. We extended an existing minimal model of seasonal dynamics of Daphnia and algae and varied the way the Daphnia population is started in spring, i.e., from free swimming individuals or from hatching resting eggs. Our model results show that temperature affects the timing of peak abundance in Daphnia and algae, and subsequently the timing of the clear water phase. When a population is started from a small inoculum of hatching resting eggs, extreme climate warming (+6°C) results in a decoupling of trophic relationships and the clear water phase fails to occur. In the other scenarios, the trophic relationships between Daphnia and its algal food source remain intact. Analysis of 36 temperate lakes showed that shallow lakes have a higher potential for climate induced match–mismatches, as the probability of active overwintering daphnids decreases with lake depth. Future research should point out whether lake depth is a direct causal factor in determining the presence of active overwintering daphnids or merely indicative for underlying causal factors such as fish predation and macrophyte cover. Priority program of the German Research Foundation—contribution 5.     

Feeding behaviour of major 0 + fish species in a shallow, eutrophic lake (Tjeukemeer, The Netherlands)

The forage base and the food selectivity of 0+ representatives of six abundant freshwater fish species were studied in a shallow, eutrophic Dutch lake. Most species relied on the zooplankton; the size-selective predation in early summer was directed to the smaller copepods and in late summer to larger cladocerans and copepods than concurrently present in the lake. Daphnia spp. and cyclopoid copepods were the main zooplankton taxa for smelt, perch and pikeperch. Energetically, the large cladoceran, Leptodora kindtii, was especially important for pikeperch. Bream and roach preyed upon smaller zooplankton than the other fish species. The influence of the zooplankton predation by abundant 0+ fish was clear from a small mean Daphnia size in September; this size is to be used as an indicator in fishery management. Neomysis integer, the most important macrofauna species, was consumed by perch, pikeperch and ruffe; pikeperch was most size-selective in this respect. The 0+ ruffe was à typically benthivorous fish. Only the 0+ pikeperch became piscivorous, especially in years when smelt was abundant.     

On non-Eltonian methods of hunting Cladocera,or impacts of the introduction of planktivorous fish on zooplankton composition and clear-water phase occurrence in a Mediterranean reservoir

Among the topics covered by Hutchinson’s Santa Rosalia article, the question of the shortening and lengthening of food webs occupies a central role. As Hutchinson realized, at the time scales of ecological studies, the impact of invader species on established food webs is the fastest shortcut to the shortening or lengthening of the food webs. The construction of thousands of dams in Spain during the last century has offered ecologists a good opportunity to test the effects of invader fish species on the plankton dynamics of these systems. In this article, a series of data related to the food web structure of Sau Reservoir is analyzed for the period 1997–2005. Parameters such as Secchi depth and chlorophyll concentration, as well as abundance and size structure of zooplankton, have been matched to the zooplankton dynamics in the reservoir. Most of the changes detected within this period are attributed to the introduction of zooplanktivorous fish in the reservoir. The Secchi depth measurements have showed a progressive diminution in the clear-water phase during recent years. These changes have been related to the decrease in the abundance of Daphnia and to the reduction of the size of zooplankton, which help to explain concomitant increases in the chlorophyll concentration in the same period. Other observed changes in the composition of the zooplankton community have been the substitution of Daphnia by Bosmina and the increase in the abundance of rotifers. Thus, the annual average abundance of Bosmina in 1997 was 70% of cladocerans, while in 2005 it reached 98%. In parallel, the percentage occurrence of individual rotifers was 40% of total zooplankton numbers but had risen to 85% at the end of the period. All these changes are attributed to the artificial expansion of the food web through stocking of the reservoir with zooplanktivorous fish (Rutilus rutilus and Alburnus alburnus). This study improves our understanding of the trophic relationships in the food web prior to the introduction of the fish.     

The longitudinal distribution and community dynamics of zooplankton in a regulated large river: a case study of the Nakdong River (Korea)

The longitudinal distribution and seasonal dynamics of zooplankton were examined along a 200-km section of the middle to lower Nakdong River, Korea. Zooplankton was sampled twice a month from January 1995 to December 1997 at five sites in the main river channel. There was considerable longitudinal variation in total zooplankton abundance (ANOVA, p < 0.001). All major zooplankton groups (rotifers, cladocerans, copepodids and nauplii) increased significantly with distance downstream along the river. There also were statistically significant seasonal differences in zooplankton abundance at the sampling sites (ANOVA, p < 0.01). Zooplankton abundance was high in spring and fall and low in summer and winter. The seasonal pattern of rotifers was similar to that of total zooplankton. This reflected the fact that rotifers (Brachionus calyciflorus, B. rubens, Keratella cochlearis and Polyarthra spp.) strongly dominated the zooplankton community at all locations. Among the macrozooplankton, small-bodied cladocerans (e.g. Bosmina spp.) dominated; the abundance of large-bodied cladocerans (e.g. Daphnia) was negligible (0–5 ind. l^–1). Among the environmental variables considered, partial residence time seemed to play the most important role in determining characteristics of the river zooplankton community.     

Seasonal shifts in the diet of lake zooplankton revealed by phospholipid fatty acid analysis

1. We analysed the phospholipid fatty acid (PLFA) profiles of seston and of the dominant zooplankter, Daphnia longispina, through the open water period in a small, dystrophic lake to investigate seasonal variation in the diet of Daphnia. Phytoplankton, heterotrophic bacteria, green sulphur bacteria and methane‐oxidizing bacteria (MOB) were all present in the water column of the lake, and previous studies have indicated that vertically migrating Daphnia can exploit all these potential food sources. 2. For adult Daphnia, although there was some correspondence between the PLFA profile of Daphnia and the concurrent seston PLFA profile, strongest correlations were between the Daphnia PLFA profile and those of potential food sources determined 7 days earlier. This interval presumably reflects the time it takes for adult Daphnia to turn over their fatty acid pool. 3. A correlation between the concentration of polyunsaturated fatty acids (PUFAs) in the epi‐ and metalimnion and measured primary production indicated that, within the total PLFA fraction, PUFAs can be useful biomarkers for phytoplankton in food‐web studies. Algal PUFAs contributed appreciably to total PLFAs in adult Daphnia during spring and summer, but less so in autumn. 4. Daphnia in the lake actually reached their highest biomass in autumn, when methanotrophic activity was also highest, and the highest magnitude of MOB‐specific PLFAs was recorded in both adult and juvenile Daphnia. A strong relationship existed between δ¹³C values of Daphnia reported previously and the proportion of MOB‐specific PLFAs in Daphnia. Autumnal mixing evidently stimulates bacterial oxidation of methane from the hypolimnion, and exploitation of the methanotrophic bacteria sustains a high Daphnia population late in the season. 5. Our results show that the PLFA composition of freshwater zooplankton like Daphnia corresponds rather well to that of their in situ diet of phytoplankton and bacteria, with a lag period of around 1 week in the case of adult animals. The PLFA profile of seston revealed the dominant available food sources, and relating these to the Daphnia PLFA profile provided insights into seasonal changes in Daphnia diet.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Summer depth selection in crustacean zooplankton in nutrient‐poor boreal lakes is affected by recent residential development

1. Strong vertical gradients in light, water temperature, oxygen, algal concentration and predator encounters during summer in stratified lakes may influence patterns of depth selection in crustacean zooplankton, especially Daphnia species. 2. To test how crustacean depth selection varies among lakes along a gradient of catchment disturbance by recent residential development and land use change, we calculated the weighted mean depth distribution of the biomass of crustaceans by day and night in eight nutrient‐poor boreal lakes. 3. Generally, the greatest biomass of crustaceans was located at the metalimnion or at the lower boundary of the euphotic zone during thermal stratification in July. The crustacean zooplankton avoided warm surface layers and tended to stay in colder deep waters by both day and night. They also remained at greater depths in lakes with a more extensive euphotic zone. 4. There was some evidence of upward nocturnal migrations of large Daphnia and copepods in some lakes, and one case of downward migration in a lake inhabited by chaoborid larvae. 5. Multivariate regression trees (MRT) were used to cluster crustaceans and Daphnia species in homogeneous groups based on lake natural and disturbance factors. For crustaceans, the depth of the euphotic zone, the sampling depth (epilimnion, metalimnion and hypolimnion), time (day or night) of sampling and the biomass of chlorophyll a were the main driving factors. For Daphnia species, the drainage area, the sampling depth, the cleared land surface area within the catchment and the concentration of total dissolved phosphorus were the main factors.     

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     

Cascading predation effects of Daphnia and copepods on microbial food web components

1. We performed a mesocosm experiment to investigate the structuring and cascading effects of two predominant crustacean mesozooplankton groups on microbial food web components. The natural summer plankton community of a mesotrophic lake was exposed to density gradients of Daphnia and copepods. Regression analysis was used to reveal top–down impacts of mesozooplankton on protists and bacteria after days 9 and 15. 2. Selective grazing by copepods caused a clear trophic cascade via ciliates to nanoplankton. Medium‐sized (20–40 μm) ciliates (mainly Oligotrichida) were particularly negatively affected by copepods whereas nanociliates (mainly Prostomatida) became more abundant. Phototrophic and heterotrophic nanoflagellates increased significantly with increasing copepod biomass, which we interpret as an indirect response to reduced grazing pressure from the medium‐sized ciliates. 3. In Daphnia‐treatments, ciliates of all size classes as well as nanoflagellates were reduced directly but the overall predation effect became most strongly visible after 15 days at higher Daphnia biomass. 4. The response of bacterioplankton involved only modest changes in bacterial biomass and cell‐size distribution along the zooplankton gradients. Increasing zooplankton biomass resulted either in a reduction (with Daphnia) or in an increase (with copepods) of bacterial biovolume, activity and production. Patterns of bacterial diversity, as measured by polymerase chain reaction–denaturing gradient gel electrophoresis (PCR–DGGE), showed no distinct grouping after 9 days, whereas a clear treatment‐coupled similarity clustering occurred after 15 days. 5. The experiment demonstrated that zooplankton‐mediated predatory interactions cascade down to the bacterial level, but also revealed that changes occurred rather slowly in this summer plankton community and were most pronounced with respect to bacterial activity and composition.