Control mechanisms of diel vertical migration: theoretical assumptions

We explore control mechanisms underlying the vertical migration of zooplankton in the water column under the predator-avoidance hypothesis. Two groups of assumptions in which the organisms are assumed to migrate vertically in order to minimize realized or effective predation pressure (type-I) and to minimize changes in realized or effective predation pressure (type-II), respectively, are investigated. Realized predation pressure is defined as the product of light intensity and relative predation abundance and the part of realized predation pressure that really affects organisms is termed as effective predation pressure. Although both types of assumptions can lead to the migration of zooplankton to avoid the mortality from predators, only the mechanisms based on type-II assumptions permit zooplankton to undergo a normal diel vertical migration (morning descent and evening ascent). The assumption of minimizing changes in realized predation pressure is based on consideration of DVM induction only by light intensity and predators. The assumption of minimizing changes in effective predation pressure takes into account, apart from light and predators also the effects of food and temperature. The latter assumption results in the same expression of migration velocity as the former one when both food and temperature are constant over water depth. A significant characteristic of the two type-II assumptions is that the relative change in light intensity plays a primary role in determining the migration velocity. The photoresponse is modified by other environmental variables: predation pressure, food and temperature. Both light and predation pressure are necessary for organisms to undertake DVM. We analyse the effect of each single variable. The modification of the phototaxis of migratory organisms depends on the vertical distribution of these variables.     

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     

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.     

From inducible defences to population dynamics: modelling refuge use and life history changes in Daphnia

We investigated the relative importance of a behavioural defence (refuge use through diel vertical migration) and a life history change (a reduced size at first reproduction) that are used by daphnids to decrease the risk of predation by visually hunting fish. We used an individual based model of a Daphnia population in a stratified lake to quantify the effects of these inducible defences on Daphnia predation-mortality and the resulting Daphnia population dynamics. Our analysis shows that diel vertical migration (DVM) confers a much stronger protection against fish predation than a reduced size at first reproduction (SFR). DVM allows daphnids to withstand a higher predation pressure in the epilimnion and it decelerates a Daphnia population decline more strongly than a reduced SFR. DVM effectively reduces the (P/B) flow of carbon from daphnids to fish.
Many theoretical studies have only considered the fitness benefits of DVM above 'staying up' in the epilimnion of a lake. Our results suggest that 'staying down' in the hypolimnion would confer an even stronger fitness benefit to Daphnia than DVM at times of peak predation risk. Daphnids that remain in the hypolimnion avoid the predation suffered by migrating daphnids around dusk and dawn. Staying down could prevent a Daphnia population decline, while DVM and a reduced SFR can only decelerate the decrease of Daphnia population densities under heavy fish predation. Staying down at high concentrations of fish infochemicals has in fact been observed within a variety of Daphnia clones and species, both in the laboratory and in stratified lakes.     

Diel vertical migration arising in a habitat selection game

Predator and prey react to each other, adjusting their behavior to maximize their fitness and optimizing their food intake while keeping their predation risk as low as possible. In a pelagic environment, prey reduce their predation mortality by adopting a diel vertical migration (DVM) strategy, avoiding their predator during their peak performance by finding refuge in deep layers during daylight hours and feeding at the surface during the night. Due to the duality of the interaction between prey and predator, we used a game theory approach to investigate whether DVM can be a suitable strategy for the predator as well as the prey. We formulated three scenarios in plankton ecology in order to address this question. A novel finding is that mixed strategies emerge as optimal over a range of the parameter space, where part of the predator or prey population adopts a DVM while the rest adopt one or other “sit and wait” strategies.     

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.     

Viewing DVM via general behaviors of zooplankton: a way bridging the success of individual and population

In this paper, we viewed the diel vertical migration (DVM) of copepod in the context of the animal's immediate behaviors of everyday concerns and constructed an instantaneous behavioral criterion effective for DVM and non-DVM behaviors. This criterion employed the function of 'venturous revenue' (VR), which is the product of the food intake and probability of the survival, to evaluate the gains and losses of the behaviors that the copepod could trade-off. The optimal behaviors are to find the optimal habitats to maximize VR. Two types of VRs are formulated and tested by the theoretical analysis and simulations. The sensed VR, monitoring the real-time changes of trade-offs and thereby determining the optimum habitat, is validated to be the effective objective function for the optimization of the behavior; whereas, the realized VR, quantifying the actual profit obtained by an optimal copepod in the sensed-VR-determined habitat, defines the life history of a specific age cohort. The achievement of a robust copepod overwintering stock through integrating the dynamics of the constituent age cohorts subjected to the instantaneous behavioral criterion for DVM clearly exemplified a possible way bridging the immediate pursuit of an individual and the end success of the population.     

Diel vertical migration of zooplankton following optimal food intake under predation

A model is developed to investigate the trade-offs between benefitsand costs involved in zooplanktonic diel vertical migration(DVM) strategies. The ‘venturous revenue’ (VR) isused as the criterion for optimal trade-offs. It is a functionof environmental factors and the age of zooplankter. Duringvertical migration, animals are assumed to check instantaneouslythe variations of environmental parameters and thereby selectthe optimal behavioral strategy to maximize the value of VR,i.e. taking up as much food as possible with a certain riskof mortality. The model is run on a diel time scale (24 h) infour possible scenarios during the animal’s life history.The results show that zooplankton can perform normal DVM balancingoptimal food intake against predation risk, with the profileof DVM largely modified by the age of zooplankter.     

Stay cool: habitat selection of a cyclopoid copepod in a north temperate oligotrophic lake

1. We studied the vertical distributions of Cyclops cf. sibiricus in Lake Toya, a north temperate oligotrophic lake. During the winter circulation period, their distribution was vertically homogeneous both day and night. During the summer stratification period, C. cf. sibiricus stayed below the thermocline. Diel vertical migration (DVM) was pronounced in advanced developmental stages, although the upper limit of the migration became deeper as the thermocline gradually descended. This seasonal change was observed throughout the 4‐year study period, implying that thermal structure is the primary determinant of C. cf. sibiricus distribution. 2. In a field experiment, C. cf. sibiricus incubated in the summer epilimnion, which most of the population never experience, developed faster and grew better than in their original habitat. We consider that trans‐thermocline DVM would not have evolved because of possible disadvantages such as the cost of migration offset the benefit observed in the field experiment. The rapid temperature change at the thermocline may act as a swimming‐cost estimator for the copepod. 3. Low food availability in deep water during the summer stratification period seemed to determine the lower limit of C. cf. sibiricus distribution, and the copepod minimised the risks of predation by fish via DVM. These results suggest that C. cf. sibiricus modified their distribution seasonally to obtain maximum benefit in terms of individual fitness.     

Food resource effects on diel movements and body size of cisco in north-temperate lakes

The movement patterns and body size of fishes are influenced by a host of physical and biological conditions, including temperature and oxygen, prey densities and foraging potential, growth optimization, and predation risk. Our objectives were to (1) investigate variability in vertical movement patterns of cisco (Coregonus artedi) in a variety of inland lakes using hydroacoustics, (2) explore the causal mechanisms influencing movements through the use of temperature/oxygen, foraging, growth, and predation risk models, and (3) examine factors that may contribute to variations in cisco body size by considering all available information. Our results show that cisco vertical movements vary substantially, with different populations performing normal diel vertical migrations (DVM), no DVM, and reverse DVM in lakes throughout Minnesota and northern Wisconsin, USA. Cisco populations with the smallest body size were found in lakes with lower zooplankton densities. These smaller fish showed movements to areas of highest foraging or growth potential during the day and night, despite moving out of preferred temperature and oxygen conditions and into areas of highest predation risk. In lakes with higher zooplankton densities, cisco grew larger and had movements more consistent with behavioral thermoregulation and predator avoidance, while remaining in areas with less than maximum foraging and growth potential. Furthermore, the composition of potential prey items present in each lake was also important. Cisco that performed reverse DVM consumed mostly copepods and cladocerans, while cisco that exhibited normal DVM or no migration consumed proportionally more macro-zooplankton species. Overall, our results show previously undocumented variation in migration patterns of a fish species, the mechanisms underlying those movements, and the potential impact on their growth potential.     

Zooplankton Migration in Three Lakes of Western New York

We examined the diel vertical migration (DVM) of zooplankton in three lakes of western New York; Lakes Conesus, Lime, and McCargo. In all three lakes, the dipteran predator Chaoborus was a more dramatic migrator than any of the cladocerans or copepods. In contrast, another invertebrate predator, Leptodora, seemed to have the least vertical dispersal. In Conesus Lake, following the evening ascent and upper-water night-positioning of Chaoborus flavicans, Daphnia pulicaria expanded its own vertical range of dispersal thereby decreasing its degree of overlap with Chaoborus. The vertical distribution of Diaptomus sicilis was mostly below that of Daphnia pulicaria, thus reducing possible competition for food resources. Poor oxygen conditions in the lower waters of Lakes Lime and McCargo restricted all zooplankton (except Chaoborus) to an epilimnetic zone where the regions of niche overlap and predation were narrowed. Such annually-induced restrictions on vertical dispersal are probably common features of numerous stratified lakes with hypolimnetic reducing conditions. No fixed pattern of vertical dispersal or migration is likely in lakes where stratification and non-stratification follow each other seasonally. Vertical migration does provide some flexibility in niche separation, but the potential for chaotic behavior in interacting predators and prey may confound simple explanations of DVM.     

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.     

Diel horizontal migration of zooplankton: costs and benefits of inhabiting the littoral

1. In some shallow lakes, Daphnia and other important pelagic consumers of phytoplankton undergo diel horizontal migration (DHM) into macrophytes or other structures in the littoral zone. Some authors have suggested that DHM reduces predation by fishes on Daphnia and other cladocerans, resulting in a lower phytoplankton biomass in shallow lakes than would occur without DHM. The costs and benefits of DHM, and its potential implications in biomanipulation, are relatively unknown, however. 2. In this review, we compare studies on diel vertical migration (DVM) to assess factors potentially influencing DHM (e.g. predators, food, light, temperature, dissolved oxygen, pH). We first provide examples of DHM and examine avoidance by Daphnia of both planktivorous (PL) fishes and predacious invertebrates. 3. We argue that DHM should be favoured when the abundance of macrophytes is high (which reduces planktivory) and the abundance of piscivores in the littoral is sufficient to reduce planktivores. Food in the littoral zone may favour DHM by daphnids, but the quality of these resources relative to pelagic phytoplankton is largely unknown. 4. We suggest that abiotic conditions, such as light, temperature, dissolved oxygen and pH, are less likely to influence DHM than DVM because weaker gradients of these conditions occur horizontally in shallow lakes relative to vertical gradients in deep lakes. 5. Because our understanding of DHM is rudimentary, we highlight potentially important research areas: studying a variety of systems, comparing temporal and spatial scales of DHM in relation to DVM, quantifying positive and negative influences of macrophytes, focusing on the role of invertebrate predation, testing the performance of cladocerans on littoral versus pelagic foods (quantity and quality), investigating the potential influence of temperature, and constructing comprehensive models that can predict the likelihood of DHM. Our ability to biomanipulate shallow lakes to create or maintain the desired clear water state will increase as we learn more about the factors initiating and influencing DHM.     

Benthic habitat is an integral part of freshwater Mysis ecology

1. Diel vertical migration (DVM) is common in aquatic organisms. The trade-off between reduced predation risk in deeper, darker waters during the day and increased foraging opportunities closer to the surface at night is a leading hypothesis for DVM behaviour.
2. Diel vertical migration behaviour has dominated research and assessment frameworks for Mysis, an omnivorous mid-trophic level macroinvertebrate that exhibits strong DVM between benthic and pelagic habitats and plays key roles in many deep lake ecosystems. However, some historical literature and more recent evidence indicate that mysids also remain on the bottom at night, counter to expectations of DVM.
3. We surveyed the freshwater Mysis literature using Web of Science (WoS; 1945–2019) to quantify the frequency of studies on demographics, diets, and feeding experiments that considered, assessed, or included Mysis that did not migrate vertically but remained in benthic habitats. We supplemented our WoS survey with literature searches for relevant papers published prior to 1945, journal articles and theses not listed in WoS, and additional references known to the authors but missing from WoS (e.g. only 47% of the papers used to evaluate in situ diets were identified by WoS).
4. Results from the survey suggest that relatively little attention has been paid to the benthic components of Mysis ecology. Moreover, the literature suggests that reliance on Mysis sampling protocols using pelagic gear at night provides an incomplete picture of Mysis populations and their role in ecosystem structure and function.
5. We summarise current knowledge of Mysis DVM and provide an expanded framework that more fully considers the role of benthic habitat. Acknowledging benthic habitat as an integral part of Mysis ecology will enable research to better understand the role of Mysis in food web processes.

     

Daphnia magna trade‐off safety from UV radiation for food

Research on diel vertical migration (DVM) is generally conducted at the population level, whereas few studies have focused on how individual animals behaviorally respond to threats when also having access to foraging opportunities. We utilized a 3D tracking platform to record the swimming behavior of Daphnia magna exposed to ultraviolet radiation (UVR) in the presence or absence of a food patch. We analyzed the vertical position of individuals before and during UVR exposure and found that the presence of food reduced the average swimming depth during both sections of the trial. Since UVR is a strong driver of zooplankton behavior, our results highlight that biotic factors, such as food patches, have profound effects on both the amplitude and the frequency of avoidance behavior. In a broader context, the trade‐off between threats and food adds to our understanding of the strength and variance of behavioral responses to threats, including DVM.     

Two hundred years of zooplankton vertical migration research

Vertical migration is a geographically and taxonomically widespread behaviour among zooplankton that spans across diel and seasonal timescales. The shorter-term diel vertical migration (DVM) has a periodicity of up to 1 day and was first described by the French naturalist Georges Cuvier in 1817. In 1888, the German marine biologist Carl Chun described the longer-term seasonal vertical migration (SVM), which has a periodicity of ca. 1 year. The proximate control and adaptive significance of DVM have been extensively studied and are well understood. DVM is generally a behaviour controlled by ambient irradiance, which allows herbivorous zooplankton to feed in food-rich shallower waters during the night when light-dependent (visual) predation risk is minimal and take refuge in deeper, darker waters during daytime. However, DVMs of herbivorous zooplankton are followed by their predators, producing complex predator–prey patterns that may be traced across multiple trophic levels. In contrast to DVM, SVM research is relatively young and its causes and consequences are less well understood. During periods of seasonal environmental deterioration, SVM allows zooplankton to evacuate shallower waters seasonally and take refuge in deeper waters often in a state of dormancy. Both DVM and SVM play a significant role in the vertical transport of organic carbon to deeper waters (biological carbon sequestration), and hence in the buffering of global climate change. Although many animal migrations are expected to change under future climate scenarios, little is known about the potential implications of global climate change on zooplankton vertical migrations and its impact on the biological carbon sequestration process. Further, the combined influence of DVM and SVM in determining zooplankton fitness and maintenance of their horizontal (geographic) distributions is not well understood. The contrasting spatial (deep versus shallow) and temporal (diel versus seasonal) scales over which these two migrations occur lead to challenges in studying them at higher spatial, temporal and biological resolution and coverage. Extending the largely population-based vertical migration knowledge base to individual-based studies will be an important way forward. While tracking individual zooplankton in their natural habitats remains a major challenge, conducting trophic-scale, high-resolution, year-round studies that utilise emerging field sampling and observation techniques, molecular genetic tools and computational hardware and software will be the best solution to improve our understanding of zooplankton vertical migrations.     

Diel vertical migration of the copepod <Emphasis Type="Italic">Thermocyclops inversus</Emphasis> (Kiefer, 1936) in a tropical reservoir: the role of oxygen and the spatial overlap with <Emphasis Type="Italic">Chaoborus</Emphasis>

The effect of water transparency, dissolved oxygen concentration and the invertebrate predator Chaoborus brasiliensis on the day–night vertical distribution of the copepod cyclopoid Thermocyclops inversus was investigated in a shallow tropical reservoir, Nado Reservoir, Belo Horizonte, Brazil. Diel cycles were carried out over a period of 12 consecutive months, between October 1999 and September 2000. The different developmental stages of T. inversus exhibited diel vertical migration (DVM) and displayed a clear ontogenetic trend, with the amplitude of DVM increasing with the age of the organism, and ranging from 0.4 m to 0.8 m for nauplii, 0.4 m to 1.2 m for copepodite, and 1.1 m to 2.1 m for adults. We observed that seasonal changes in dissolved oxygen and C. brasiliensis directly influenced the vertical distribution of the copepod population in this reservoir. Furthermore, it was showed that the diurnal vertical migration is an important predator avoidance behavior since it diminished the spatial overlap between prey and its potential predator. This finding supports the hypothesis that the vertical migration is a defense mechanism against predation. Thus, T. inversus is able to remain in the anoxic layers during day light hours, and at night they move upwards avoiding hypolimnetic waters to escape from predation by Chaoborus.     

Diel vertical migration of freshwater fishes – proximate triggers,ultimate causes and research perspectives

1. Diel vertical migrations (DVM) are typical for many cold‐water fish species such as Pacific salmons (Oncorhynchus spp.) and coregonids (Coregonus spp.) inhabiting deep lakes. A comprehensive recent overview of DVM in freshwater fish has not been available, however. 2. The main proximate trigger of DVM in freshwater fish is the diel change in light intensity, with declining illumination at dusk triggering the ascent and the increase at dawn triggering the descent. Additional proximate cues are hydrostatic pressure and water temperature, which may guide fish into particular water layers at night. 3. Ultimate causes of DVM encompass bioenergetics efficiency, feeding opportunities and predator avoidance. None of these factors alone can explain the DVM in all cases. Multi‐factorial hypotheses, such as the ‘antipredation window’ combined with the thermal niche hypothesis, are more likely to explain DVM. It is suggested that planktivorous fish move within a layer sufficiently well illuminated to capture zooplankton, but too dark for predators to feed upon the migrating fish. In complete darkness, fish seek layers with a temperature that optimises bioenergetics efficiency. The strength of each factor may differ from lake to lake, and hence system‐specific individual analyses are needed. 4. Mechanistic details that are still poorly explored are the costs of buoyancy regulation and migration, the critical light thresholds for feeding of planktivorous and piscivorous fish, and predator assessment by (and size‐dependent predation risk of) the prey fish. 5. A comprehensive understanding of the adaptive value of DVM can be attained only if the behaviour of individual fish within migrating populations is explicitly taken into account. Size, condition and reproductive value differ between individuals, suggesting that migrating populations should split into migrants and non‐migrants for whom the balance between mortality risk and growth rate can differ. There is increasing evidence for this type of partial DVM within populations. 6. Whereas patterns of DVM are well documented, the evolution of DVM is still only poorly understood. Because experimental approaches at realistic natural scales remain difficult, a combination of comprehensive data sets with modelling is likely to resolve the relative importance of different proximate and ultimate causes behind DVM in fish.     

Temperature selection in Brook Charr: lab experiments,field studies,and matching the Fry curve

Zooplankton diel vertical migration (DVM) is an ecologically important process, affecting nutrient transport and trophic interactions. Available measurements of zooplankton displacement velocity during the DVM in the field are rare; therefore, it is not known which factors are key in driving this velocity. We measured the velocity of the migrating layer at sunset (upward bulk velocity) and sunrise (downwards velocity) in summer 2015 and 2016 in a lake using the backscatter strength (VBS) from an acoustic Doppler current profiler. We collected time series of temperature, relative change in light intensity chlorophyll-a concentration and zooplankton concentration. Our data show that upward velocities increased during the summer and were not enhanced by food, light intensity or by VBS, which is a proxy for zooplankton concentration and size. Upward velocities were strongly correlated with the water temperature in the migrating layer, suggesting that temperature could be a key factor controlling swimming activity. Downward velocities were constant, likely because Daphnia passively sink at sunrise, as suggested by our model of Daphnia sinking rate. Zooplankton migrations mediate trophic interactions and web food structure in pelagic ecosystems. An understanding of the potential environmental determinants of this behaviour is therefore essential to our knowledge of ecosystem functioning.

     

Partial diel vertical migrations in pelagic fish

1. Field studies on diel vertical migration (DVM) usually report uniform behaviour with population-wide ascents and descents during crepuscular periods. This contrasts partial seasonal migrations of many animal populations, where individuals choose either the resident or the migrant strategy depending on population density, feeding opportunity and predation risk in the resident and migrant habitats. 2. We tested whether DVM of freshwater zooplanktivorous fish (Coregonus spp.) resembles partial migrations. Twenty-eight hydroacoustic surveys were performed in the deep Lake Stechlin (Germany) between 2000 and 2010, with samplings encompassing all months between March and December. Zooplankton samples were simultaneously taken in epilimnetic and hypolimnetic layers. Fish obtained from depth-stratified samplings by a midwater trawl were used to test for individual differences between residents and migrants. 3. We show for the first time that DVMs of freshwater fish resemble patterns of partial migrations often found in seasonal environments. Across all samplings, 7-33% of fish did not ascend at dusk, but exhibited the resident strategy. The proportion or residents increased at low zooplankton feeding rates in the daytime habitat and during months when the temperature difference between daytime and night-time habitats was minor. 4. Slightly larger size and higher caloric density of migrants over residents in one of the coexisting Coregonus species suggested that individual differences contributed to the migration strategy performed. However, these results were based on one sample only, and extrapolation to the entire data set is not possible. 5. Our results are indirect evidence that the balance between migrants and residents may primarily depend on the trade-off between feeding gains and metabolic and predation costs of migration. However, the results also suggest that the global fitness consequences for the resident and migrant strategies may not be identical, rendering the importance of individual traits in the 'decision to migrate' likely.