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.     

The genetics of phototaxis in Daphnia magna: Existence of three phenotypes for vertical migration among parthenogenetic females

Using a cloning technique, we revealed the existence of three phototactically distinct types in Daphnia magna, viz. negatives, positives, and gipsies. The latter migrate continuously between a low and a high light intensity. The expression of this behaviour is genetically determined, although positives and negatives can reversibly switch into gipsies.It is proposed that two opposite genotypes exist, with the gipsy behaviour epigenetically induced. All juveniles behave more or less uniformly, irrespective of their origin. The phototactic responses develop rather late during ontogenesis.     

Size distribution of Daphnia longispina in the vertical profile

D. longispina of the meromictic lake El Tobar is a round-headed form. It never has a helmet, but in summer a small proportion of immature individuals (0.9–1.2 mm females and males) have one or two neck teeth. The size structure of this Daphnia population, as well as the vertical distribution and migration of different size-classes, were studied in September and November of 1991 and April of 1992. The large variation in mean size and size at first reproduction, as well as the occurrence of different patterns of vertical migration are interpreted as responses to different predator situations. At the end of April, when Daphnia mortality by visually hunting predators is dominating, a typical nocturnal migration is adopted and size distribution is biased to smaller size classes. In November, when mortality is mainly attributed to the nocturnally migrating Chaoborus, Daphnia shows a reversed migration pattern. In September, when the population of Daphnia is responding to both visual (fish) and non-visual predators (Chaoborus), it adopts a pattern of twilight migration. The presence of neck teeth in vulnerable size classes in September might be an additional adaptative response to Chaoborus predation. In September, the size structure of the Daphnia population is shifted to larger classes and the vertical distribution of size classes shows a pronounced segregation between juveniles and adults. Juveniles are found closer to the surface, while adults dwell predominantly in the rich, deep waters near the oxicline. This suggests that an additional advantage of the ascent of the adult Daphnia exploiting those deep resources is the release of young in more oxygenated and warmer waters. The Daphnia population of lake El Tobar is known to be clonally diverse, and the changing frequency of genotypes could play an important part in the observed seasonal differences in behaviour and size.     

The interaction of Chaoborus size and vertical distribution determines predation effects on Daphnia

1. Larvae of Chaoborus, the phantom midge, are important pelagic planktivores in many freshwater lakes and ponds. The effect of Chaoborus on its prey depends on its size, especially mouth gape diameter, and vertical migration pattern, which affects predator–prey spatial overlap. These two features vary considerably in different Chaoborus species and instars. In this study, the interacting effects of both Chaoborus size and vertical distribution on population growth of Daphnia pulex was analysed with a field enclosure experiment and a matrix population model. 2. In the field experiment, Daphnia were grown in four replicated treatments that included a control (no Chaoborus) and three combinations of instar III and IV Chaoborus of two species (C. trivittatus and C. americanus). Parameters of the matrix model were based on differences between Chaoborus species and instars in capture and ingestion of Daphnia of differing sizes (prey vulnerability) and in vertical overlap with Daphnia in each treatment (density risk). 3. In comparison with the control, the two treatments containing the smaller, migratory C. americanus showed a significant effect on Daphnia population growth rate, while the treatment containing only the larger, non‐migratory C. trivittatus did not. The model accurately simulated these effects. 4. A Daphnia predation risk model, which uses prey vulnerability and density risk parameters, illustrated the individual and combined effect of the different Chaoborus types on Daphnia. Daphnia have a high prey vulnerability to the large C. trivittatus, but overall predation risk was low because of very little overlap. On the contrary, the smaller C. americanus affects only a small range of Daphnia instars, each with a low vulnerability, yet those instars that were vulnerable had a very high density risk because of an increased overlap. 5. This analysis of Daphnia predation risk parameters with coexisting Chaoborus species strongly supports an integrated approach using both size and vertical distribution to determine the ultimate predation effect on Daphnia.     

Evidence for a diurnal horizontal migration in Daphnia hyalina lacustris Sars

Diurnal sampling of a transect across a shallow, eutrophic gravel pit suggested that a diurnal horizontal migration was present in Daphnia hyalina lacustris, in which it remained in the littoral zone during the day and moved out into the open water at night. The avoidance of fish predation seemed the most likely ultimate cause of this migration.     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.     

Daphnia exhibit diurnal vertical migration in shallow rock-pools

We report diurnal changes in vertical distribution ofDaphnia longispina andD. magna in shallow (25–60 cm) rock-pools on Baltic islands, southern Finland. During the daytime shallow rock-poolDaphnia reside in water layers close to the sediment while at midnight they either are evenly distributed throughout the water column, or have a modal peak of distribution in the surface water. Contrary to this pattern, no changes were observed in vertical distribution ofD. longispina at noon and at midnight in a 150 cm deep pond. We suggest that the pattern ofDaphina vertical migration in shallow rock-pools is not satisfactorily explained by predation.     

Morphological Variation in Daphnia Cucullata Sars with Progressive Eutrophication of a Polymictic Lowland Reservoir

Morphological variation in Daphnia cucullata Sars was studied from 1994 to 2001 when fast increasing water trophy of the Siemianówka Reservoir was related to high TP and chlorophyll concentrations. During this period, D. cucullata biomass, as well as its contribution to the total cladocerans biomass clearly decreased. We found that among the six morphological parameters analyzed in D. cucullata, body length, height of the head and length of the tail spine had a tendency to decrease. The maximum values of these traits and carapace length also decreased. The morphology of D. cucullata appeared to be influenced by factors such as food availability, fish predation and an exclusive presence of D. cucullata among Daphniidae species. Increased water mixing in the reservoir may have also affected the changes in the head inclination of D. cucullata.     

Diel horizontal migration and swarm formation in Daphnia in response to Chaoborus

Dense swarms of Daphnia longispina (up to 4000 animals l^–1) were recorded along the littoral zone in a lake where Chaoborus flavicans is considered the main predator. D. longispina coexisted with D. pulex, but there were no D. pulex in the littoral swarms. Swarms were less dense at night (about 1/10 the density), and D. longispina exhibited diel horizontal migrations by aggregating in the littoral during the day and spreading out at night. Laboratory experiments showed that Chaoborus capture efficiency on juvenile daphnids was higher in the light compared to darkness, and that Daphnia exhibited a behavioural response to water that had previously contained Chaoborus. We conclude that predation from Chaoborus can be an important factor affecting the distribution patterns of Daphnia observed in this lake. The behavioural experiments indicated that this influence might be partly mediated by chemical agents.     

The effect of Daphnia on filament length of blue-green algae

Water from a hypertrophic lake rich in filamentous blue-green algae was passed through a continuous-flow system of aquaria containing Daphnia magna, and a control system without Daphnia. Daphnia caused a significant decrease in the blue-green algal density, and a two-fold reduction in filament length. It is suggested that feeding activity of Daphnia may result in an increase in the availability of blue-green filaments to filter-feeding cladocerans.     

On the cost of vertical migration: are feeding conditions really worse at greater depths?

SUMMARY 1. The ultimate explanation for diel vertical migration (DVM) of zooplankton is the avoidance of visual predation in surface waters. Studies on migrating zooplankton have shown that remaining in the cold and food-poor hypolimnion during the day, however, has demographic costs. Higher temperatures and greater food concentrations in the surface waters are thought to be the main reasons why Daphnia species move upwards at night.
2. In this study, we investigated the growth condition of daphniids raised on seston taken from different depths from a lake with and without a deep-water chlorophyll maximum.
3. Juvenile growth rates of Daphnia galeata x hyalina from the lake without a deep-water chlorophyll maximum were similar for all treatments. After temperature correction, however, growth rates were significantly higher on seston taken from the surface layers.
4. In contrast, in the lake with the deep-water chlorophyll maximum, D. galeata growth rates were higher in deeper strata, even after temperature correction. Although this lake had a weak temperature gradient, D. galeata left the food-rich strata at night and migrated into the surface food-poor environment. Invertebrate predation and oxygen depletion are probably not the reasons for the nocturnal upward migration into the surface strata. Therefore, we assume that D. galeata migrates upwards to take advantage of higher temperatures. Using several temperature–egg-development models, we could not, however, fully explain this behaviour.     

Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments

1. Invasion of tropical zooplankton into temperate lakes provides an interesting opportunity to explore habitat segregation in a thermal gradient. 2. We explored differing vertical positioning of native and exotic Daphnia (Daphnia mendotae and Daphnia lumholtzi) in a large indoor mesocosm system (Plön plankton towers) during 2 month‐long experiments. The two towers were manipulated to provide gradients of both temperature (15–29 °C) and algal food (0.05–0.58 mg C L^?1) and a day–night cycle. 3. Both juvenile and adult D. lumholtzi showed a ‘typical’ vertical migration pattern, with higher densities in the epilimnion at night than during the day. They avoided the food‐poor middle layer. In contrast, D. mendotae adults showed little tendency to migrate into the epilimnion at night, remaining in the cooler hypolimnion while juveniles migrated. The vertical distribution of D. mendotae appeared unaffected by the presence of D. lumholtzi. 4. The strong migration behaviour of D. lumholtzi in the absence of fish cues suggests that this behaviour may be a constitutive trait. Habitat partitioning of the two species is probably the result of different thermal tolerances, with D. mendotae constrained to remaining in deeper water by high temperatures in the epilimnion and the tropical D. lumholtzi able to use the warm epilimnion at night.     

Effects of food limitation,notonectid predation,and temperature on the population dynamics of Daphnia carinata

The relative contributions of invertebrate predation (Notonectidae: Anisops spp.), food limitation, and certain abiotic factors in driving the population dynamics of Daphnia carinata were studied for a two year period in a large farm dam in southern Victoria, Australia. Detailed measurements were made on the population densities of Daphnia and Anisops spp., the amount of food available, the nutritional status, and the size-specific fecundity of Daphnia. The density of the Daphnia population at the field site oscillated closely with water temperature. The amplitude of the population fluctuations varied seasonally, being much greater during the warmer months of the year and switching to fluctuations with low peaks when water temperature dropped below approximately 15°C. Anisops spp. densities were greatest in winter and declined during the spring of each year. Nymphs appeared in late spring and early summer and numerically dominated the population. During the warmer periods of the year, the daphnid population went through a series of rapid growth phases leading to over-exploitation of food resources and subsequent population collapses. Daphnia population densities were not correlated with Anisops numbers suggesting that predation was not a major regulatory factor during the warmer periods of the year. When water temperatures fell below ca. 15°C daphnid population densities remained low despite high food levels. During this period the impact of Anisops may have been greater. Two distinct phases were identified: a warm water period when food limitation was the main regulatory factor, and a cool water period when Anisops predation may have been the paramount factor. Low oxygen concentrations were associated with heavy rainfall in the spring and may have had a limiting effect on Daphnia for short periods. Daphnia may have had an important role in sustaining the Anisops population in the pond over each winter.     

Daphnia size structure,vertical migration,and phosphorus redistribution

The timing and magnitude of diel migration in two daphnid assemblages were determined from a series of vertical profiles of daphnid size distribution. Animals were collected concurrently for gut fullness determination. Only large daphnids (> 1.4 mm) migrated, but these animals could account for substantial vertical and diel differences in phosphorus excretion rate. Gut fullness measurements and time courses of diel vertical migration suggested that large Daphnia can cause a net downward flux of phosphorus during summer in thermally stratified lakes.     

Ovigerity,selective predation,and variable diel vertical migration in Euchaeta elongata (Copepoda: Calanoida)

Summary We present a statistical analysis of a previously published (Yen, 1983) but heretofore unanalyzed data set on the vertical distributions and diel vertical migration (DVM) of adult females of the marine planktonic copepod Euchaeta elongata in Dabob Bay, Washington, USA. Non-ovigerous females were strongly migratory on all four dates sampled, residing between 75–175 m during the day and at shallower depths during the night, commonly entering the upper 50 m of the water column. In contrast, ovigerous females were non-migratory or weakly migratory, largely remaining between 100–175 m both day and night, and entering the upper 50 m of the water column only rarely. Thus non-ovigerous females always migrated much more strongly, as measured by both amplitude of migration and the proportion of animals migrating, than did ovigerous females. These results led us to hypothesize that differential susceptibility to visually orienting predators was the cause of these differences in DVM behavior in female E. elongata, and we subsequently undertook an experimental study of the feeding selectivity of the copepod's natural predator, Pacific herring (Clupea harengus pallasi). Pacific herring exhibited a highly significant preference for ovigerous over nonovigerous adult female E. elongata. The demographic consequences of variable DVM in adult female E. elongata were investigated by way of life table analyses. Results indicated that under conditions of thermal stratification of the water column there is a distinct demographic disadvantage (reduced rate of realized population growth) incurred by non-migratory or weakly migratory ovigerous females due to delayed egg development at cooler subsurface temperatures. We conclude that ovigerous female E. elongata remain at depth both day and night to avoid visually orienting predators, and that such behavior must afford the copepod a demographic advantage of no less than a 26% reduction in adult mortality to offset the demographic cost of delayed egg development.     

The effect of parthenogenic eggs in Daphnia magna on prey location by the bluegill sunfish (Lepomis macrochirus)

The distance at which the bluegill sunfish (Lepomis macrochirus) can locate Daphnia magna with parthenogenic eggs is shown to be significantly greater than the reactive distance for non-gravid Daphnia of the same size. This difference is due to greater visibility of the gravid prey and not to different locomotor behavior, since there was no correlation between the number of eggs carried by a Daphnia and the antennal beat frequency. Based on this experiment and other observations, an explanation is given for selective predation of polymorphic cladoceran populations.     

Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton

Many pelagic animal species in the marine environment and in lakes migrate to deeper water layers before sunrise and return around sunset. The amplitude of these diel vertical migrations (DVM) varies from several hundreds of metres in the oceans to approx. 5–20 m in lakes. DVM can be studied from a proximate and an ultimate point of view. A proximate analysis is intended to reveal the underlying behavioural mechanism and the factors that cause the daily displacements. The ultimate analysis deals with the adaptive significance of DVM and the driving forces that were responsible for the selection of the traits essential to the behavioural mechanism. The freshwater cladoceran Daphnia is the best studied species and results can be used to model migration behaviour in general. Phototaxis in Daphnia spp., which is defined as a light-oriented swimming towards (positive phototaxis) or away (negative phototaxis) from a light source, is considered the most important mechanism basic to DVM. A distinction has been made between primary phototaxis which occurs when light intensity is constant, and secondary phototaxis which is caused by changes in light intensity. Both types of reaction are superimposed on normal swimming. This swimming of Daphnia spp. consists of alternating upwards and downwards displacements over small distances. An internal oscillator seems to be at the base of these alternations. Primary phototaxis is the result of a dominance of either the upwards or the downwards oscillator phase, and the direction depends on internal and external factors: for example, fish-mediated chemicals or kairomones induce a downwards drift. Adverse environmental factors may produce a persistent primary phototaxis. Rare clones of D. magna have been found that show also persistent positive or negative primary phototaxis and interbreeding of the two types produces intermediate progeny: thus a genetic component seems to be involved. Also secondary phototaxis is superimposed on normal swimming: a continuous increase in light intensity amplifies the downwards oscillator phase and decreases the upwards phase. A threshold must be succeeded which depends on the rate and the duration of the relative change in light intensity. The relation between both is given by the stimulus strength versus stimulus duration curve. An absolute threshold or rheobase exists, defined as the minimum rate of change causing a response if continued for an infinitely long time. DVM in a lake takes place during a period of 1-5-2 h when light changes are higher than the rheobase threshold. Accelerations in the rate of relative increase in light intensity strongly enhance downwards swimming in Daphnia spp. and this enhancement increases with increasing fish kairomone and food concentration. This phenomenon may represent a ‘decision-making mechanism’ to realize the adaptive goal of DVM: at high fish predator densities, thus high kairomone concentrations, and sufficiently high food concentrations, DVM is profitable but not so at low concentrations. Body axis orientation in Daphnia spp. is controlled with regard to light-dark boundaries or contrasts. Under water, contrasts are present at the boundaries of the illuminated circular window which results from the maximum angle of refraction at 48–9° with the normal (Snell's window). Contrasts are fixed by the compound eye and appropriate turning of the body axis orients the daphnid in an upwards or an obliquely downwards direction. A predisposition for a positively or negatively phototactic orientation seems to be the result of a disturbed balance of the two oscillators governing normal swimming. Some investigators have tried to study DVM at a laboratory scale during a 24 h cycle. To imitate nature, properties of a natural water column, such as a large temperature gradient, were compressed into a few cm. With appropriate light intensity changes, vertical distributions looking like DVM were obtained. The results can be explained by phototactic reactions and the artificial nature of the compressed environmental factors but do not compare with DVM in the field. A mechanistic model of DVM based on phototaxis is presented. Both, primary and secondary phototaxis is considered an extension of normal swimming. Using the light intensity changes of dawn and the differential enhancement of kairomones and food concentrations, amplitudes of DVM could be simulated comparable to those in a lake. The most important adaptive significance of DVM is avoidance of visual predators such as juvenile fish. However, in the absence of fish kairomones, small-scale DVMs are often present, which were probably evolved for UV-protection, and are realized by not enhanced phototaxis. In addition, the ‘decision-making mechanism’ was probably evolved as based on the enhanced phototactic reaction to accelerations in the rate of relative changes in light intensity and the presence of fish kairomones.     

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.     

An analysis of the phototactic behaviour of Daphnia magna clones and their sexual descendants

Daphnia magna clones, isolated from different natural populations, were inbred and crossbred. Some aspects of the heredity of the phototactic behaviour, quantified by an index, were analysed. Heritability in the broad sense was determined for several assemblages of clones through calculation of the clonal repeatability. Heritability sensu strictu was determined by offspring-parent regressions. Frequency distributions of a phototactic index of several inbred and crossbred families are given.The results indicate a significant contribution of additive genetic variance to the total phenotypic variance. Comparison of heritability estimates suggests an important genetic interaction component.Some implications of these results on the variability of vertical migration patterns in natural populations are discussed.