Clonal integration facilitates invasiveness of the alien aquatic plant Myriophyllum aquaticum L. under heterogeneous water availability

The avoidance of visually feeding fish has long been considered as the primary driver of diel vertical migration of zooplankton. The diurnal vertical distribution of Cyclops gr. abyssorum, Arctodiaptomus alpinus, and Daphnia gr. longispina from 13 alpine lakes with fish (Salvelinus fontinalis) and without, was compared in order to understand whether fish in transparent lakes reduce the presence of large zooplankton from the irradiated zone. We used the light level at each sampling depth and the size of each specimen as proxies of predation risk, and we tested two predictions: (P1) the relative abundance of zooplankton in the well-lit surface waters vs. the darker waters will be greater in fishless lakes; (P2) the size of zooplankton in the well-lit surface waters vs. the deeper, darker waters will be greater in fishless lakes. We did not find any evidence of the validity of P1, but we confirmed P2 for Arctodiaptomus alpinus. These results support with new field data the Transparency Regulator Hypothesis, which argues that in transparent lakes, fish predation is less important for the vertical distribution of zooplankton than ultraviolet radiation, and further suggest that zooplankton size rather than vertical distribution may be more effective against visual predators in transparent lakes.     

Is Ambient Light during the High Arctic Polar Night Sufficient to Act as a Visual Cue for Zooplankton?

The light regime is an ecologically important factor in pelagic habitats, influencing a range of biological processes. However, the availability and importance of light to these processes in high Arctic zooplankton communities during periods of ''complete'' darkness (polar night) are poorly studied. Here we characterized the ambient light regime throughout the diel cycle during the high Arctic polar night, and ask whether visual systems of Arctic zooplankton can detect the low levels of irradiance available at this time. To this end, light measurements with a purpose-built irradiance sensor and coupled all-sky digital photographs were used to characterize diel skylight irradiance patterns over 24 hours at 79°N in January 2014 and 2015. Subsequent skylight spectral irradiance and in-water optical property measurements were used to model the underwater light field as a function of depth, which was then weighted by the electrophysiologically determined visual spectral sensitivity of a dominant high Arctic zooplankter, Thysanoessa inermis. Irradiance in air ranged between 1–1.5 x 10^-5 μmol photons m^-2 s^-1 (400–700 nm) in clear weather conditions at noon and with the moon below the horizon, hence values reflect only solar illumination. Radiative transfer modelling generated underwater light fields with peak transmission at blue-green wavelengths, with a 465 nm transmission maximum in shallow water shifting to 485 nm with depth. To the eye of a zooplankter, light from the surface to 75 m exhibits a maximum at 485 nm, with longer wavelengths (>600 nm) being of little visual significance. Our data are the first quantitative characterisation, including absolute intensities, spectral composition and photoperiod of biologically relevant solar ambient light in the high Arctic during the polar night, and indicate that some species of Arctic zooplankton are able to detect and utilize ambient light down to 20–30m depth during the Arctic polar night.     

Photophysiological cycles in Arctic krill are entrained by weak midday twilight during the Polar Night

Light plays a fundamental role in the ecology of organisms in nearly all habitats on Earth and is central for processes such as vision and the entrainment of the circadian clock. The poles represent extreme light regimes with an annual light cycle including periods of Midnight Sun and Polar Night. The Arctic Ocean extends to the North Pole, and marine light extremes reach their maximum extent in this habitat. During the Polar Night, traditional definitions of day and night and seasonal photoperiod become irrelevant since there are only “twilight” periods defined by the sun’s elevation below the horizon at midday; we term this “midday twilight.” Here, we characterize light across a latitudinal gradient (76.5° N to 81° N) during Polar Night in January. Our light measurements demonstrate that the classical solar diel light cycle dominant at lower latitudes is modulated during Arctic Polar Night by lunar and auroral components. We therefore question whether this particular ambient light environment is relevant to behavioral and visual processes. We reveal from acoustic field observations that the zooplankton community is undergoing diel vertical migration (DVM) behavior. Furthermore, using electroretinogram (ERG) recording under constant darkness, we show that the main migratory species, Arctic krill (Thysanoessa inermis) show endogenous increases in visual sensitivity during the subjective night. This change in sensitivity is comparable to that under exogenous dim light acclimations, although differences in speed of vision suggest separate mechanisms. We conclude that the extremely weak midday twilight experienced by krill at high latitudes during the darkest parts of the year has physiological and ecological relevance.

This study shows that ambient light cycles set an internal rhythm that controls visual sensitivity of Arctic krill during the Polar Night – the darkest part of the year, when the sun remains below the horizon all day. This demonstrates that biologically relevant photoperiods can be achieved during this time of "midday twilight".     

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.     

Relaxed circadian rhythm in zooplankton along a latitudinal gradient

To test whether aquatic invertebrates are able to adjust their diel migratory cycle to different day length and presence of predators, we performed standardized enclosure experiments in shallow lakes at four different latitudes from southern Spain, with strong night–day cycles, to Finland where daylight is almost continuous during summer. We show here that nearly continuous daylight at high latitudes causes a relaxation in diel migratory behaviour in zooplankton irrespective of predation risk. At lower latitudes, however, similar conditions lead to pronounced diel rhythms in migration. Hence, zooplankton may show local behavioural adaptations in their circadian rhythm. They are also able to make risk assessments as to whether diel migration is beneficial or not, manifested in a lack of diel migration at near constant daylight, irrespective of predator presence. Our results provide an additional explanation to previous knowledge regarding diel migrations among aquatic invertebrates by showing that both physical (light) and biological (predation) factors may affect the migratory behaviour.     

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 variation of phytoplankton functional groups in a subtropical reservoir in southern Brazil during an autumnal stratification period

A knowledge of diel variation and the vertical distribution of phytoplankton communities may contribute to a better understanding of the driving factors of key species. Applying functional-group classification provides important information on the causes of species selection in the pelagic community. The diel variation of phytoplankton functional groups was analysed during an autumnal stratification period with the aim of understanding their changes in the vertical position related to light, mixing regime and grazing pressure. Phytoplankton and zooplankton communities were sampled every 4 h during a 24-h period in a vertical profile in a subtropical meso-eutrophic reservoir. Strong stratification during a 24-h cycle and a mixed clear epilimnion with partial atelomixis marked the autumn season in the Faxinal reservoir, southern Brazil. The highest phytoplankton densities and biomass were found during the second part of the day, a general pattern reported in the literature, and may be explained by zooplankton dynamics. During the 24-h cycle, phytoplankton functional groups lacking a self-regulating capacity and those able to regulate their vertical position were vertically segregated in the lake. The diel behaviour of both groups was driven by the mixing regime (including atelomixis), light and zooplankton grazing pressure.     

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.     

Orientation and navigation in coastal and estuarine zooplankton

The axiom that zooplankton species exhibit upwards migration behaviour at night is challenged by recent findings. Such behaviour is not universal, may vary during ontogeny, and is occasionally reversed. Moreover, in some estuarine and coastal zooplankton species vertical migration rhythms are of tidal, not diel, periodicity. There is evidence for several species that vertical migrations are endogenously controlled, occurring in constant conditions in the laboratory, suggesting that they have arisen under considerable selection pressure. They appear to play a significant role in orientation and navigation of coastal and estuarine zooplankton, predicated on the selective advantage of closure of life cycles. Vertical migrations between water masses moving in different directions at tidal, diel or longer timescales permit dispersal and recruitment, or retention, of planktonic larvae and adults in favourable ecological locations. Exogenous factors serving as cues for, or directly controlling, vertical migration rhythms include light, hydrostatic pressure, salinity, temperature, water movements and gravity responses, besides biological factors such as the timing of larval release, duration of larval life and predator/prey interactions. Behavioural characteristics should be quantified and factored into dispersal models which assume that zooplankton adults and larvae are advected as passive particles.     

Diel vertical distribution of zooplankton in Lake Naivasha,Kenya

Diurnal and diel vertical distribution of limnetic zooplankton species in relation to temperature and dissolved oxygen profiles was examined at a central station in Lake Naivasha. During calm days thermal stratification developed gradually from late morning to reach maximum formation at mid-day. Dissolved oxygen concentrations showed similar vertical profiles to temperature. These stratifications were, however, short lived and were broken up in late afternoons by the wind induced poly-holomictic nature of the lake. During the day most zooplankters aggregate at the top 3–4 metre zone of the water column coincident with maximum photosynthetic activity. The pattern of diel vertical distribution of zooplankton in Lake Naivasha is undefinedly even. The absence of significant diel changes in the distribution of the limnetic zooplankton may be related to the absence of permanent physico-chemical boundaries and lack of predation pressure in the open water.     

The aggregation of polar cod (Boreogadus saida) in the deep Atlantic layer of ice-covered Amundsen Gulf (Beaufort Sea) in winter

During the Circumpolar Flaw Lead System Study (CFL, 2007–2008), large aggregations of polar cod were detected in winter in the Amundsen Gulf (Western Canadian Arctic) using the EK60 echosounder of the CCGS Amundsen research icebreaker. Biomass estimated over 10 months reached a maximum of 0.732 kg m⁻² in February. Aggregations were encountered only in the presence of an ice cover from December to April. The vertical extent of the aggregations was dictated by temperature and zooplankton prey distribution. In winter, polar cod generally occupied the relatively warm deep Atlantic Layer (>0°C), but a fraction of the densest aggregations occasionally followed zooplankton prey up into the cold Pacific Halocline (−1.6 to 0°C). The diel vertical migration of polar cod was precisely synchronized with the seasonally increasing photoperiod. Throughout winter, polar cod aggregations migrated to progressively deeper regions (from 220 to 550 m bottom depths) in response to increasing light intensity, presumably to avoid predation by visual predators such as the ringed seal. Comparing Amundsen Gulf and Franklin Bay indicates that the entrapment of polar cod in embayments during winter is an important mechanism to provide marine mammal predators with dense concentrations of their main prey within their diving range.     

Diel variation in predator abundance, predation risk and prey distribution in shallow-water estuarine habitats

Predation by visual predators is often affected by light conditions and may therefore exhibit strong diel variation. The dominant predators on grass shrimp, Palaemonetes pugio, are finfish predators that are thought to locate their prey by visual cues. We examined the response of grass shrimp to diel variation in predation risk in the nearshore shallow waters of the Chesapeake Bay. We used diel shoreline seines to assess the relative abundance of predators. We assessed the relative risk of predation with shrimp tethered at refuge (30 cm) and nonrefuge (60 cm) depths. To measure grass shrimp response to predation risk, we used dipnets to monitor habitat use. Four predominantly visual predators dominated the shoreline seine catches, Fundulus heteroclitus, Micropogonias undulatus, Morone americana and Morone saxatilis. Total predator abundance had a diel component, with dramatic nighttime decreases in total abundance, whereas guild composition and relative abundance remained unchanged. Relative predation risk for tethered shrimp exhibited significant time by habitat interaction. During the day, depth negatively affected survivorship of tethered shrimp while at night overall survivorship increased and there was no effect of depth. Shrimp habitats use reflected diel predation risks. Abundances in the near shore were highest during the day with decreased abundances at night. Together, the seine and tethering data highlight the importance for a refuge (e.g., shallow water) from predation during the daytime and a relaxation of predation pressure at night.     

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.     

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

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

Modeling patterns of zooplankton diel vertical migration

Realized predation pressure, defined as the product of predationpressure and light intensity, expresses the mortality pressuredue to visual predation. The part of realized predation pressurewhich is sensed by organisms is here considered to be relatedto food level and temperature. This partly realized predationpressure is referred to as sensed predation pressure. We proposea possible control mechanism of diel vertical migration (DVM):organisms move vertically following the minimum change in sensedpredation pressure. To investigate this assumption, we presenta math ematical model of DVM. We assume that when predatorsare present, the food level is above a minimal level, and temperatureis higher than the tolerance of organisms to growth, prey organismsundertake DVM following the minimum change in sensed predationpressure. We examine how patterns of migration may be affectedby changes in water clarity, predation pressure, food leveland temperature. This work supports the assumption that minimizingchanges in sensed predation pressure can explain the wide variationin the vertical profile of zooplankton.     

Vertical distribution and pigmentation of Antarctic zooplankton determined by a blend of UV radiation, predation and food availability

Selection pressure induced by simultaneously occurring environmental threats is a major evolutionary driver for organisms in terrestrial, as well as in aquatic ecosystems. For example, protection against ultraviolet radiation (UVR) and predation include both morphological and behavioral components. Here we address those selective pressures on zooplankton by performing a latitudinal monitoring, combined with mechanistic experiments in the Antarctic Southern Ocean, where the UVR-threat is extremely high. We assessed vertical distributions of zooplankton along the Antarctic coast showing that animals were most abundant at 20–80 m and tended to avoid the surface at sites with clear water. UVR-threat disappeared at between 9 and 15 m at sites with low and high water transparency, respectively. Light levels were, however, sufficient for visual fish predation down to approximately 19 and 37 m, respectively. The few zooplankton that were present in surface waters had high levels of non-pigmented UVR-protective compounds (mycosporine-like amino acids) compared to deeper dwelling zooplankton. Overall they had low levels of red pigmented UVR-protective compounds (carotenoids), suggesting high predation on pigmented individuals. In a complementary laboratory study we showed that levels of UVR-protective compounds increased considerably when zooplankton were exposed to UVR in the absence of predator cues. The recently developed transparency-regulator hypothesis predicts that UVR avoidance is an important driver to diel vertical migration in transparent waters, such as in Antarctica. We, however, conclude that copepods resided well below the level where UVR had diminished to very low levels and that predator avoidance or food availability are more likely drivers of zooplankton vertical depth distribution in transparent marine systems.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

Size-structured risk assessments govern Daphnia migration

One of the more fascinating phenomena in nature is animal mass migrations and in oceans and freshwaters, diel variations in depth distribution of zooplankton are a phenomenon that has intrigued scientists for more than a century. In our study, we show that zooplankton are able to assess the threat level of ultraviolet radiation and adjust their depth distribution to this level at a very fine tuned scale. Moreover, predation risk induces a size-structured depth separation, such that small individuals, which we show are less vulnerable to predation than larger, make a risk assessment and continue feeding in surface waters during day, offering a competitive release from down-migrating larger animals. Hence, we mechanistically show that such simple organisms as invertebrate zooplankton are able to make individual, size-specific decisions regarding how to compromise between threats from both predators and UV radiation, and adjust their diel migratory patterns accordingly.     

Diel vertical migration of Arctic zooplankton during the polar night

High-latitude environments show extreme seasonal variation in physical and biological variables. The classic paradigm of Arctic marine ecosystems holds that most biological processes slow down or cease during the polar night. One key process that is generally assumed to cease during winter is diel vertical migration (DVM) of zooplankton. DVM constitutes the largest synchronized movement of biomass on the planet, and is of paramount importance for marine ecosystem function and carbon cycling. Here we present acoustic data that demonstrate a synchronized DVM behaviour of zooplankton that continues throughout the Arctic winter, in both open and ice-covered waters. We argue that even during the polar night, DVM is regulated by diel variations in solar and lunar illumination, which are at intensities far below the threshold of human perception. We also demonstrate that winter DVM is stronger in open waters compared with ice-covered waters. This suggests that the biologically mediated vertical flux of carbon will increase if there is a continued retreat of the Arctic winter sea ice cover.     

Diel vertical movements of zooplankton in lake La Cruz (Cuenca, Spain)

The study of diel vertical movements of zooplankton in a smallmeromictic karstic lake (lake La Cruz), presenting highly stratifiedwaters, was performed using two different methodologies: (i)samples were taken along the vertical profile and were comparedat different hours in a diel cycle; (ii) some plankton trapswere located at different depths, covering different periodsof time, to catch organisms going upwards and downwards. Themain subject of this study has been the vertical movements affectingrotifers since they were dominant in the zooplankton of thislake, but we have also included the results obtained for otherzooplankton groups. The results indicate an almost general movementin epilimnetic waters corresponding to the ‘normal’pattern of migration (ascent at dusk and descent at dawn); inmetalimnetic and hypolimnetic waters, vertical movements weregreatly reduced and were more independent of the diel cycle.The results also confirm that the dense populations at theselevels of the oxic–anoxic boundary are made up of activeanimals. Among the causes accounting for our results, the cyclesof activity (mainly feeding) connected with the diel cycle couldbe important in explaining vertical movements of filter-feederrotifers. In larger zooplankters, visual predation might bethe main reason for such behavior.