Effects of UV-B radiation on near-surface zooplankton of puget sound

Summary An increase in incident solar ultraviolet irradiation, resulting from possible deterioration of the stratospheric ozone layer, would have important biological effects. Though the oceans are relatively opaque to UV radiation, compared to visible light, increases in incident UV may affect organisms living within the first few meters of the sea surface.Shrimp larvae, crab larvae, and euphausids were exposed to various low levels of simulated solar UV radiation (UV-B, 290–315 nm) under laboratory conditions. Comparisons between solar and artificial spectra were based on spectroradiometric measurements converted to erythemally effective irradiance. These zooplankton tolerated UV-B irradiance levels up to threshold levels with no significant reduction in survival or developmental rates compared to control organisms. Beyond the threshold levels, activity, development, and survival rapidly declined. The apparent UV thresholds are near present incident UV levels.Observed survival threshold levels for each experimental group were superimposed on seasonal solar incident UV levels at the experimental site. These threshold levels appeared to be exceeded by median ambient UV levels late in the season of surface occurrence of each species. UV increases resulting from ozone depletion may significantly shorten this season. Although the apparent impact would be lessened by the decrease in UV with depth, irreversible detrimental effects would probably occur before reported survival thresholds were exceeded.Contribution No. 1107 from the Department of Oceanography, University of Washington, Seattle, WA 98195, USA     

Dose/dose-rate responses of shrimp larvae to UV-B radiation

Summary Previous work indicated dose-rate thresholds in the effects of UV-B on the near-surface larvae of three shrimp species. Additional observations suggest that the total dose response varies with dose-rate. Below 0.002 Wm^-2 _[DNA] irradiance no significant effect is noted in activity, development, or survival. Beyond that dose-rate threshold, shrimp larvae are significantly affected if the total dose exceeds about 85 Jm^-2 _[DNA]. Predictions cannot be made without both the dose-rate and the dose.These dose/dose-rate thresholds are compared to four-year mean dose/dose-rate solar UV-B irradiances at the experimental site, measured at the surface and calculated for 1 m depth. The probability that the shrimp larvae would receive lethal irradiance is low for the first half of the season of surface occurrence, even with a 44% increase in damaging UV radiation.Contribution No. 1183 from the Department of Oceanography, University of Washington, Seattle, WA 98195, USA     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Effects of UV-B irradiated algae on zooplankton grazing

We tested the effects of UV-B stressed algae on grazing rates of zooplankton. Four algal species (Chlamydomonas reinhardtii, Cryptomonas sp., Scenedesmus obliquus and Microcystis aeruginosa) were used as food and fed to three zooplankton species (Daphnia galeata, Bosmina longirostris and Brachionus calyciflorus), representing different taxonomic groups. The phytoplankton species were cultured under PAR conditions, and under PAR supplemented with UV-B radiation at two intensities (0.3 W m^–2 and 0.7 W m^–2, 6 hours per day). Ingestion and incorporation experiments were performed at two food levels (0.1 and 1.0 mg C l^–1) using radiotracer techniques. The effect of food concentration on ingestion and incorporation rate was significant for all three zooplankton species, but the effect of UV-B radiation was more complex. The reactions of the zooplankton species to UV-B stressed algae were different. UV-B stressed algae did not affect Daphnia grazing rates. For Bosmina the rates increased when feeding on UV-B stressed Microcystis and decreased when feeding on UV-B stressed Chlamydomonas, compared with non-stressed algae. Brachionus grazing rates were increased when feeding on UV-B stressed Cryptomonas and UV-B stressed Scenedesmus, and decreased when feeding on UV-B stressed Microcystis, compared with non-stressed algae. These results suggest that on a short time scale UV-B radiation may result in increased grazing rates of zooplankton, but also in decreased grazing rates. Long term effects of UV-B radiation on phytoplankton and zooplankton communities are therefore difficult to predict.     

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

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

Erratum to: How do UV radiation, temperature, and zooplankton influence the dynamics of alpine phytoplankton communities?

Plankton in mountain lakes are confronted with generally higher levels of incident ultraviolet radiation (UVR), lower temperatures, and shorter growing seasons than their lower elevation counterparts. The direct inhibitory effects of high UVR and low temperatures on montane phytoplankton are widely recognized. Yet little is known about the indirect effects of these two abiotic factors on phytoplankton, and more specifically whether they alter zooplankton grazing rates which may in turn influence phytoplankton. Here, we report the results of field microcosm experiments that examine the impact of temperature and UVR on phytoplankton growth rates and zooplankton grazing rates (by adult female calanoid copepods). We also examine consequent changes in the absolute and relative abundance of the four dominant phytoplankton species present in the source lake (Asterionella formosa, Dinobryon sp., Discostella stelligera, and Fragilaria crotonensis). All four species exhibited higher growth rates at higher temperatures and three of the four species (all except Dinobryon) exhibited lower growth rates in the presence of UVR versus when shielded from UVR. The in situ grazing rates of zooplankton had significant effects on all species except Asterionella. Lower temperatures significantly reduced grazing rates on Fragilaria and Discostella, but not Dinobryon. While UVR had no effect on zooplankton grazing on any of the four species, there was a significant interaction effect of temperature and UVR on zooplankton grazing on Dinobryon. Discostella and Dinobryon increased in abundance relative to the other species in the presence of UVR. Colder temperatures, the presence of zooplankton, and UVR all had consistently negative effects on rates of increase in overall phytoplankton biomass. These results demonstrate the importance of indirect as well as direct effects of climate forcing by UVR and temperature on phytoplankton community composition in mountain lakes, and suggest that warmer climates and higher UVR levels may favor certain species over others.     

Penetration of UV-B radiation in foliage: evidence that the epidermis behaves as a non-uniform filter

In some plants, particularly herbaceous species, a considerable proportion of incident ultraviolet-B radiation (UV-B, 280-320 nm) penetrates into the leaf mesophyll where it is potentially damaging to nucleic acids and the photosyn-thetic machinery. We used optical techniques to look at the spatial variation in UV-B penetration through the epidermis of foliage of two herbaceous species (Chenopodium album and Smilacina stellata)and a conifer (Picea pun-gens). Measurements of UV-B penetration in intact foliage with a fibre-optic microprobe revealed that 300 nm radiation reached 161±36μm (mean±SD) into leaves of C. album, 154±40μm in S. stellata and 17±2μm in P. pungens, with epidermal transmittance being 39±14%, 55±19% and 0%, respectively. A thin polymer film was developed which fluoresced blue when irradiated by UV-B. Fresh epidermal leaf peels were placed over the film and irradiated with UV-B, and microscopic examination of the film from below allowed us to determine the spatial pattern of UV-B penetration through the epidermis. In herbaceous species, film fluorescence below cell walls, but not epidermal and guard cell protoplasts indicated that UV-B transmittance was much greater through anticlinal cell wall regions than protoplasts. Ultraviolet-B transmittance through large areas of epidermal cells could be induced by plasmolysis. Epidermal transmittance was also relatively high through stomal pores (and what appear to be nuclei in Smilacina), but relatively low through stomatal guard cells. Results from the fluorescing film technique were substantiated by direct measurements of UV-B transmittance through epidermal peels with a fibre-optic microprobe run paradermally along the bottom or inner side of irradiated peels. In Smilacina, we estimate that UV-B epidermal transmittance was up to 90% through anticlinal cell wall regions, but <10% through protoplast areas. In contrast to herbaceous species, we did not detect any UV-B transmittance through the epidermis of P. pungens with either the fluorescing film or the fibre-optic microprobe technique. The epidermis appears to be a much more spatially uniform UV-B filter in conifers than in these herbaceous species.     

Reduction in omega-3 fatty acids by UV-B irradiation in microalgae

UV-B irradiation reduced the levels of omega-3 fatty acid, eicosapentaenoic acid (EPA, 20:53) and docosahexaenoic acid (DHA, 22:63), in microalgae; the degree of reduction varied with species.Chaetoceros calcitrans andSkeletonema costatum were high UV-B tolerant species, followed byPhaeodactylum tricornutum, Chroomonas salina, Pavlova lutheri, andThalassiosira pseudonana.Isochrysis galbana (T.ISO) andProrocentrum micans were UV- B sensitive. Cells in logarithmic phase were most sensitive to UV- B irradiation. Nitrate-, phosphate-, or sulphate-starved cells were more UV-B sensitive than non-starved cells grown in a complete basal medium. A relatively short exposure to high UV-B was more damaging than a longer exposure to lower irradiance. Visible light intensity levels had a profound impact on the sensitivity of microalgal cultures to UV-B, with high levels decreasing UV-B dependent damage. Addition of polyamines (putrescine, spermidine or spermine) or an amino acid (cysteine) to the culture medium minimized the reduction of omega-3 fatty acid content in microalgae caused by UV-B irradiation.Author for correspondence     

Influence of trophic status and conductivity on zooplankton composition in lakes and ponds of Torres del Paine National Park (Chile)

The Torres del Paine National Park is located in the southern Chilean Patagonia. This park has numerous and heterogeneous lakes and ponds with different trophic status and zooplankton composition. The aim of this study was to analyze the influence of trophic status and conductivity on zooplankton assemblages in lakes and ponds within the Torres del Paine National Park. The water bodies described in the present study were previously classified in three groups. The first group consisted of large, deep and oligotrophic lakes with fish populations, low zooplankton species diversity and high predominance of calanoid copepods of small body size. The second group contained small mesotrophic lakes with fish populations and relatively high predominance of small sized daphnids. The third group consisted of fishless ponds of different trophic status, wide conductivity gradient and with zooplankton species of relatively large body size. The results show that Daphnids abundance was directly related to chlorophyll-a concentration and inversely associated with conductivity. Calanoids abundance was also directly associated with conductivity.     

The influence of enhanced UV-B radiation on the spring geophyte Pulmonaria officinalis

Pulmonaria officinalis is an understorey spring geophyte, which starts its vegetative period before full foliation of the tree storey. During its early growth phase it is exposed to full solar radiation, therefore the enhanced UV-B radiation could present a threat to this species. An outdoor experiment in which potted plants were exposed to below ambient, ambient, and above ambient (corresponding to 17% ozone reduction) UV-B radiation, was conducted in order to evaluate the radiation effects. The amount of photosynthetic pigments and photochemical efficiency of PSII were not affected, but the amount of UV-B absorbing compounds was lower in plants grown under reduced UV-B. This change was measurable after only fourteen days in reproductive shoots, while in the vegetative shoots, it was not detectable until after three months. The leaves of P. officinalis are variegated and the light green spots became less transparent to PAR under enhanced UV-B. The results reveal that under simulated 17% ozone depletion the harmful effects of UV-B on the measured parameters were negligible.     

Planktivory in the changing Lake Huron zooplankton community: Bythotrephes consumption exceeds that of Mysis and fish

1. Oligotrophic lakes are generally dominated by calanoid copepods because of their competitive advantage over cladocerans at low prey densities. Planktivory also can alter zooplankton community structure. We sought to understand the role of planktivory in driving recent changes to the zooplankton community of Lake Huron, a large oligotrophic lake on the border of Canada and the United States. We tested the hypothesis that excessive predation by fish (rainbow smelt Osmerus mordax, bloater Coregonus hoyi) and invertebrates (Mysis relicta, Bythotrephes longimanus) had driven observed declines in cladoceran and cyclopoid copepod biomass between 2002 and 2007. 2. We used a field sampling and bioenergetics modelling approach to generate estimates of daily consumption by planktivores at two 91‐m depth sites in northern Lake Huron, U.S.A., for each month, May–October 2007. Daily consumption was compared to daily zooplankton production. 3. Bythotrephes was the dominant planktivore and estimated to have eaten 78% of all zooplankton consumed. Bythotrephes consumption exceeded total zooplankton production between July and October. Mysis consumed 19% of all the zooplankton consumed and exceeded zooplankton production in October. Consumption by fish was relatively unimportant – eating only 3% of all zooplankton consumed. 4. Because Bythotrephes was so important, we explored other consumption estimation methods that predict lower Bythotrephes consumption. Under this scenario, Mysis was the most important planktivore, and Bythotrephes consumption exceeded zooplankton production only in August. 5. Our results provide no support for the hypothesis that excessive fish consumption directly contributed to the decline of cladocerans and cyclopoid copepods in Lake Huron. Rather, they highlight the importance of invertebrate planktivores in structuring zooplankton communities, especially for those foods webs that have both Bythotrephes and Mysis. Together, these species occupy the epi‐, meta‐ and hypolimnion, leaving limited refuge for zooplankton prey.     

Zooplankton dynamics in a gypsum karst lake and interrelation with the abiotic environment

Zooplankton species composition and abundance variation was studied in Lake Amvrakia, which is a deep, temperate, gypsum karst lake situated in the western Greece. The two year survey of zooplankton revealed 33 species (23 rotifers, five cladocerans, four copepods and one mollusc larva). The mean integrated abundance of the total zooplankton ranged between 83.6 and 442.7 ind. L⁻¹, with the higher density to be recorded in the surface 0–20 m layer. Small numbers of specimens of almost all species were found also in the hypoxic or anoxic hypolimnion. Copepods and especially the calanoid Eudiaptomus drieschi dominated the zooplankton community throughout the sampling period, followed by Dreissena polymorpha larvae, rotifers and cladocerans. Seasonal succession among the cladocerans and the most abundant rotifer species was observed. The concentration of chlorophyll-a was the most important factor for the variation of total zooplankton, as well as for the rotifers’ community. Dissolved oxygen affected copepods and cladocerans, water level correlated mainly with the molluscs larvae of D. polymorpha, while temperature influenced the variation of several rotifers, the cladoceran Diaphanosoma orghidani and the mollusc larvae. Negative correlation of conductivity with the cladoceran Daphnia cucullata and the copepods E. drieschi and Macrocyclops albidus was found. The differences in species composition found in Lake Amvrakia in comparison to the nearby lakes are probably ought to the geographical isolation and perhaps to its particular chemistry (e.g., elevated conductivity).     

Trace Metals in Mesozooplankton of the North Sea: Concentrations in Different Taxa and Preliminary Results on Bioaccumulation in Copepod Collectives (Calanus finmarchicus/C. helgolandicus)

Trace metals (Cd, Cu, Zp, Pb, Ni) were determined in different zooplankton taxa to assess their role in the biogeochemical cycle in the North Sea. Results of semi-static bioaccumulation experiments on board ship using collectives of Calanus finmarchicus/C. helgolandicus were in agreement with a net accumulation strategy, thus meeting an inevitable precondition for their utilization as biomonitors. Cd, Cu and Zn concentrations in field samples of zooplankton displayed a pronounced variability between taxa, with Cd ranging from 0.13 mg kg-1 (d.w.) in fish larvae to 5I mg kg-1 in hyperiid amphipods. Almost no information is available about metal requirements of copepods. However, most of our and worldwide reported Cu concentrations in calanoid copepods (6-22 mg kg⁻¹) coincided with estimates of enzymatic requirements of decapod crustaceans. Cd levels in Calanus collectives increased significantly from 0.68 mg kg-1 in the German Bight to 11 mg kg⁻¹ at some stations around Scotland above 59° N, in line with literature data reported for benthic decapod crustaceans.     

The effect of ultraviolet radiation on the vertical distribution and mortality of estuarine zooplankton

The effect of UV-B radiation on the vertical distribution ofthree calanoid copepod species (Tortanus dextrilobatus, Acartiuraspp. and Acanthacartia spp.) and three larval stages of Pacificherring, Clupea pallasi (1-, 7-, and 14-day-old larvae) wasinvestigated. A series of 2 m high columnar tanks equipped withinfra-red light and video-microscopy was used to investigatethe vertical distribution of zooplankton in the presence andabsence of UV-B radiation. In the presence of UV-B radiation,T.dextrilobatus and 1-day-old C.pallasi resided about 50 cmdeeper than in the absence of UV-B, while Acartiura spp. andAcanthacartia spp. showed no (or only minimal) change in verticaldistribution. Mortality experiments were also conducted outdoorsin which each copepod species was exposed to full or reducednatural radiation levels. Only T.dextrilobatus showed an increasein mortality when exposed to full radiation. Our results showedthat T.dextrilobatus and 1-day-old C.pallasi larvae were sensitiveto UV radiation (UVR), and to reduce or eliminate UV-inducedstress, they avoided the surface of the water column when UV-Bradiation was present. Copepod species were chosen to span arange of pigmentation: T.dextrilobatus (heavily pigmented),Acartiura spp. (moderately pigmented) and Acanthacartia spp.(not pigmented). The pigmentation did not appear to play a rolein UVR tolerance of the copepods, but may be a factor determiningUV tolerance of C.pallasi.     

High Mortality of Red Sea Zooplankton under Ambient Solar Radiation

High solar radiation along with extreme transparency leads to high penetration of solar radiation in the Red Sea, potentially harmful to biota inhabiting the upper water column, including zooplankton. Here we show, based on experimental assessments of solar radiation dose-mortality curves on eight common taxa, the mortality of zooplankton in the oligotrophic waters of the Red Sea to increase steeply with ambient levels of solar radiation in the Red Sea. Responses curves linking solar radiation doses with zooplankton mortality were evaluated by exposing organisms, enclosed in quartz bottles, allowing all the wavelengths of solar radiation to penetrate, to five different levels of ambient solar radiation (100%, 21.6%, 7.2%, 3.2% and 0% of solar radiation). The maximum mortality rates under ambient solar radiation levels averaged (±standard error of the mean, SEM) 18.4±5.8% h⁻¹, five-fold greater than the average mortality in the dark for the eight taxa tested. The UV-B radiation required for mortality rates to reach ½of maximum values averaged (±SEM) 12±5.6 h⁻¹% of incident UVB radiation, equivalent to the UV-B dose at 19.2±2.7 m depth in open coastal Red Sea waters. These results confirm that Red Sea zooplankton are highly vulnerable to ambient solar radiation, as a consequence of the combination of high incident radiation and high water transparency allowing deep penetration of damaging UV-B radiation. These results provide evidence of the significance of ambient solar radiation levels as a stressor of marine zooplankton communities in tropical, oligotrophic waters. Because the oligotrophic ocean extends across 70% of the ocean surface, solar radiation can be a globally-significant stressor for the ocean ecosystem, by constraining zooplankton use of the upper levels of the water column and, therefore, the efficiency of food transfer up the food web in the oligotrophic ocean.     

P-load,phytoplankton, zooplankton and fish stock in Loosdrecht Lake and Tjeukemeer: confounding effects of predation and food availability

The fish community in the Loosdrecht lakes is dominated by bream, pikeperch and smelt and is characteristic of shallow eutrophic lakes in The Netherlands. The biomasses of the respective fish species amount to ca. 250, 25 and 10 kg ha^–1 and correspond to those in Tjeukemeer, another lake in The Netherlands. The average size of bream, however, is much smaller in the Loosdrecht lakes as a consequence of poorer feeding conditions. The zooplankton community in the Loosdrecht lakes is predominantly composed of relatively small species such as Daphnia cucullata, Bosmina coregoni and cyclopoid copepods, whereas in Tjeukemeer, Daphnia hyalina is permanently present in relatively high densities and the other species show a larger mean length. In the Loosdrecht lakes, the absence of D. hyalina and the smaller sizes of the other zooplankton species could be the consequence of a higher predation pressure, in combination with unfavourable feeding conditions for the zooplankton including the low density of green algae and the high density of filamentous cyanobacteria. A biomanipulation experiment in Lake Breukeleveen, one of the Loosdrecht lakes, indicated that feeding conditions were too unfavourable for large zooplankton to develop in spring, when the reduced fish biomass was not yet supplemented by natural recruitment and immigration.     

The Species and Size Composition of Tropical Freshwater Zooplankton with Special Reference to the Oriental Region (South East Asia)

A contrast between temperate and tropical Cladocera, Copepoda and Rotifera zooplankton fauna shows a markedly different size composition and species diversity. There are fewer species, and these are generally smaller in the tropics than in either the North or South Temperate Zones. Larger species of the genera Daphnia and Simocephalus are relatively rare in the tropics, while members of Eurycerus and Saycia are absent. Members of the families Holopedidae, Leptodoridae and Polyphemidae increase in abundance toward the temperate regions. Two small cyclopoid copepods are common in tropical zooplankton, while many of the common larger species of other regions are absent or very rare. The chief rotifer components are Keratella tropica and species of Brachionus.     

Qualitative and quantitative structure,diversity and fluctuations in abundance of zooplankton in Lake Xolotlán (Managua)

The qualitative and quantitative structure of the zooplankton community was studied in 11 localities of Lake Xolotlan (Managua). Twelve rotifers, 4 cladoceran and 5 copepods species were identified. Community diversity (Shannon-Wienerindex) is low,viz. 0,83–2.20. At all times, copepods were the most abundant group. Rotifer densities were higher in zones influenced by rivers and organic pollution. Cladocerans were permanently present in low densities. Climatic events (rainfalls and dry periods) determined population fluctuations of the main zooplankton groups.     

Quantification of invertebrate predation and herbivory in food chains of low complexity

Zooplankton grazing impact on algae, heterotrophic flagellates and bacteria, as well as invertebrate predation on herbivorous zooplankton, were investigated in two sub-Antarctic lakes with extremely simple food chains. The two species of herbivorous zooplankton present in the lakes (the copepods boeckella michaelseni and Pseudoboeckella poppei) exerted substantial grazing pressure on algae. However, the dominant algal species exhibited properties that enabled them to avoid (large size or extruding spines, e.g. Staurastrum sp., Tribonema sp.) or compensate (recruitment from the sediment, Mallomonas sp.) grazing. There are only two potential invertebrate predators on the herbivorous copepods in the two lakes: the copepod Parabroteas sarsi and the diving beetle Lancetes claussi. Vertebrate predators are entirely abscent from sub-Antarctic lakes. Based on our experiments, we estimated that the predators would remove at most about 0.4% of the herbivorous copepods per day, whereas planktivorous fish, if present in the lakes, would have removed 5–17% of the zooplankton each day. Consequently, the invertebrate predators in these high-latitude lakes had only a marginal predation impact compared to the predation pressure on zooplankton in the presence of vertebrate predators in temperate lakes. The study of these simple systems with only two quantitatively functionally important trophic links, suggests that high grazing pressure foreces the algal community towards forms with grazer resistant adaptations such as large size, recruitment from another habitat, and grazer avoidance spines. We propose that due to such adaptations, predictions from food web theory are only partly corroborated, i.e. algal biomass actually increases with increasing productivity, although the grazer community is released from predation. In more species-rich and complex systems, e.g temperate lakes with three functionally important links, such adaptations are likely to be even more important, and, consequently, the observable effects of trophic interactions from top predators on lower trophic levels even more obscured.     

Zooplankton community structure and environmental conditions in a set of interconnected ponds

We studied the zooplankton community structure in a set of 33 interconnected shallow ponds that are restricted to a relatively small area (`De Maten', Genk, Belgium, 200 ha). As the ponds share the same water source, geology and history, and as the ponds are interconnected (reducing chance effects of dispersal with colonisation), differences in zooplankton community structure can be attributed to local biotic and abiotic interactions. We studied zooplankton community, biotic (phytoplankton, macrophyte cover, fish densities, macroinvertebrate densities), abiotic (turbidity, nutrient concentrations, pH, conductivity, iron concentration) and morphometric (depth, area, perimeter) characteristics of the different ponds. Our results indicate that the ponds differ substantially in their zooplankton community structure, and that these differences are strongly related to differences in trophic structure and biotic interactions, in concordance with the theory of alternative equilibria. Ponds in the clear-water state are characterised by large Daphnia species and species associated with the littoral zone, low chlorophyll-a concentrations, low fish densities and high macroinvertebrate densities. Ponds in the turbid-water state are characterised by high abundances of rotifers, cyclopoid copepods and the opposite environmental conditions. Some ponds show an intermediate pattern, with a dominance of small Daphnia species. Our results show that interconnected ponds may differ strongly in zooplankton community composition, and that these differences are related to differences in predation intensity (top-down) and habitat diversity (macrophyte cover).