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.     

Cladoceran community responses to biomanipulation and re‐oligotrophication in Lake Vesijärvi,Finland, as inferred from remains in annually laminated sediment

1. We studied the role of zooplankton in biomanipulation and the subsequent recovery phase in the Enonselkä basin of Lake Vesijärvi, using subfossil cladocerans in annually laminated sediment. Measures to restore the Enonselkä basin included reduction in external nutrient loading and mass removal of plankti‐ and benthivorous fish. Water clarity increased and the lake changed from a eutrophic to a mesotrophic state. However, some signs of increased turbidity were observed after 5–10 years of successful recovery. 2. Annual laminae in a freeze core sample were identified and sliced, based on the seasonal succession of diatoms. Cladoceran remains and rotifer eggs were counted, and Daphnia ephippia and Eubosmina and Bosmina ephippia and carapaces were measured. Annual changes in pelagic species composition were studied with principal component analysis. Individual species abundance, size measurements and various cladoceran‐based indices or ratios (commonly used to reconstruct changes in trophic state and fish predation) were tested for change between four distinct periods: I (1985–1988) dense fish stocks, poor water quality; II (1989–1992) fish removal; III (1993–1997) low fish density, improved water quality; IV (1998–2002) slightly increased fish density and poorer water quality. 3. After the removal of fish, the mean size of Daphnia ephippia and Eubosmina crassicornis ephippia and carapaces increased significantly. In contrast, the percentage of Daphnia did not increase. When based on ephippia, the ratio Daphnia/(Daphnia + E. crassicornis) increased, but the interpretation was obscured by the tolerance of fish predation by small Daphnia and by the fact that bosminids were the preferred food of roach. Moreover, ephippial production by E. crassicornis decreased in recent years. 4. The abundance of Diaphanosoma brachyurum and Limnosida frontosa increased significantly after the fish population was reduced, while that of Ceriodaphnia and rotifers decreased. 5. The expanding littoral vegetation along with improved water clarity was clearly reflected in the concentration of littoral species in the deep sediment core. The species diversity index for the entire subfossil community also increased. 6. The period of faltering recovery was characterised by greater interannual variability and an increased percentage of rotifers. Nevertheless, the mean sizes of Daphnia ephippia and E. crassicornis ephippia and carapaces indicated a low density of fish. The deteriorating water quality was apparently related to multiple stressors in the catchment after rehabilitation, such as intensified lakeshore building, as well as to exceptional weather conditions, challenging the management methods in use.     

The consequences of a drastic fish stock reduction in the large and shallow Lake Wolderwijd,The Netherlands. Can we understand what happened?

In 1990 an experiment started in the large and shallow lake Wolderwijd (2700 ha, mean depth 1.5 m) to improve the water quality. About 75% of the fish stock was removed (425 000 kg fish). The fish was mainly composed of bream and roach. In May 600000 young pikes (3–4 cm) were introduced.In May 1991 the water became very clear (Secchi depth 1.8 m) during a spring bloom of large Daphnia. Then the grazing by zooplankton was eight times higher than the primary production of algae and the total suspended matter concentration became very low. Compared to the situation before the fish reduction, the grazing had increased only slightly, while the primary production had decreased significantly in early spring. The fish stock reduction might have contributed to the reduction in primary production by a reduced internal nutrient load. The clear water period lasted six weeks. Daphnia disappeared in July due to food limitation, the algal biomass increased and the Secchi depth became 50 cm. Daphnia did not recover during summer, due to predation that was not caused by 0 + fish but by the mysid shrimp Neomysis integer. Neomysis could develop abundantly, because of the reduced biomass of the predator perch. The production of young fish had been low because of the cold spring weather. The cold weather was probably also responsible for the slow increase in density of macrophytes. After 1991, perch probably can control Neomysis. Due to lack of spawning places and shelter for 0 + pike, pike was probably not able to control the production of 0 + fish. In a lake of this scale, it will not be easy to get more than 50% coverage of macrophytes, which seems necessary to keep the algal biomass low by nutrient competition. Therefore, we expect also in the future a decrease in transparency in the summer. Locally, especially near Characeae, the water might stay clear.     

Sub-fossils of cladocerans in the surface sediment of 135 lakes as proxies for community structure of zooplankton,fish abundance and lake temperature

To elucidate the possibilities of using zooplankton remains in the surface sediment to describe present-days community structure and population dynamics of zooplankton, fish abundance and temperature, we compared contemporary data sampled in the pelagial during summer with the sediment record from the upper 1 cm of the sediment in 135 lakes covering a latitude gradient from Greenland in the north to New Zealand in the south. The abundance of three genera Bosmina, Daphnia and Ceriodaphnia of the total pool of ephippia was significantly related to the total abundance of the same taxa in the pelagic zone. However, in most lakes the abundance of Ceriodaphnia was higher in the sediment than in the water, which may be attributed to the overall preference by this genus for the littoral habitat. Using contemporary data from 27 Danish lakes sampled fortnightly during summer for 10 years, we found substantial inter-annual variations in the abundance of Daphnia spp., Ceriodaphnia spp., B. longirostris and B. coregoni. Yet, the sediment record mimicked the medium level well for most of the lakes, which suggests that the sediment record provides an integrated picture of the pelagic cladoceran community, which otherwise can be obtained only by long-term frequent contemporary sampling for several years. The contribution of Daphnia to the sum of Daphnia and Bosmina ephippia was negatively correlated with the abundance of fish expressed as catch per night in multi-mesh sized gill nets (CPUE). Yet, region-specific differences occurred, which partly could be eliminated by including nutrient state expressed as total phosphorus (TP) in a multiple regression. The average ratio of ephippia to the sum of ephippia and carapaces of Bosmina varied 40-fold between the sampling regions and was significantly negatively related to summer mean air temperature, and for Danish lakes also, albeit weakly, to fish CPUE but not to chlorophyll a. Apparently, temperature is the most important factor determining the ratio of parthenogenetic to ephippia producing specimens of Bosmina. We conclude that the sediment record of cladocerans is a useful indicator of community structure of pelagic cladocerans and the abundance of fish and temperature.     

Relative impacts of Daphnia grazing and direct stimulation by fish on phytoplankton abundance in mesocosm communities

• 1 Planktivorous fish were hypothesised to influence the abundance of algal biomass in lakes by changing zooplankton grazing, affecting zooplankton nutrient recycling and by direct recycling of nutrients to phytoplankton. The relative roles of direct fish effects vs. zooplankton grazing were tested in mesocosm experiments by adding to natural communities large grazing zooplankton (Daphnia carinata) and small planktivorous fish (mosquitofish or juveniles of Australian golden perch).
• 2 The addition of Daphnia to natural communities reduced the numbers of all phytoplankton less than 30 µm in size, but did not affect total biomass of phytoplankton as large Volvox colonies predominated.
• 3 The addition of Daphnia also reduced the abundance of some small (Moina, Bosmina, Keratella) and large (adult Boeckella) zooplankton, suggesting competitive interactions within zooplankton.
• 4 The addition of mosquitofish to communities containing Daphnia further reduced the abundance of some small zooplankton (Moina, Keratella), but increased the numbers of Daphnia and adult Boeckella. In spite of the likely increase in grazing due to Daphnia, the abundance of total phytoplankton and dominant alga Volvox did not decline in the presence of mosquitofish but was maintained at a significantly higher level than in control.
• 5 The addition of juveniles of golden perch to communities containing Daphnia reduced the abundance of small zooplankton (Moina), increased the abundance of large zooplankton (adult Boeckella) but had no significant effect on Daphnia and total phytoplankton abundance.
• 6 The results of the present study suggest that some planktivorous fish can promote the growth of phytoplankton in a direct way, probably by recycling nutrients, and even in the presence of large grazers. However, the manifestation of the direct effect of fish can vary with fish species.

     

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

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

Effects of predator-released chemicals on some life history parameters of Daphnia pulex

The effects of chemicals released by fish and Chaoborus larvae on some life history traits in Daphnia pulex were studied in an in situ enclosure experiment. The mean size of Daphnia individuals was smaller in the presence of fish-released cues. Also the minimal size of an egg bearing female in the presence of fish exudates was smaller than in the population exposed to the chemicals released by Chaoborus larvae as well as in the control population. Fish-released chemicals caused the increase in clutch size in Daphnia. There were no statistically significant differences between the studied life history parameters of the control and Chaoborus treatments. The results are discussed in reference to recent laboratory research.     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Environmental effects of introducing whitefish,Coregonus lavaretus (L.), in Lake Ring

The impact of whitefish (Coregonus lavaretus (L.)) on the trophic structure of eutrophic lakes was studied in Lake Ring, a small eutrophic Danish lake (22.5 ha, mean depth 2.9 m) in which the natural fish fauna is dominated by pike (Esox lucius L.), perch (Perca fluviatilis L.), and eel (Anguilla anguilla (L.)), roach (Rutilus rutilus (L.)) and burbot (Lota lota (L.)) being the only other fish species present. A total of 10993 0+ whitefish were stocked in the lake from October 1989 to July 1990 and the structure of the fish, zooplankton and benthic invertebrate communities studied during the period 1989–1991. Stomach contents analysis revealed that the whitefish mainly ate Daphnia and copepods in 1990–1991, the proportion of copepods decreasing with increasing size of the fish and Daphnia being the overall most important food source. The density of Daphnia in the lake decreased from 72 ind. 1^-1 in 1989 to 9 ind. 1^-1 in 1991; concomitantly the large species Daphnia magna and D. pulex almost disappeared and the density of cyclopoid copepods increased from 72 to 101 ind. 1^-1, presumably because of improved food conditions, while that of calanoid copepods remained virtually unchanged. As a result chl-a increased from 19 to 47 µg 1^-1 and Secchi depth decreased from 2.4 m to 1.7 m, despite there being no change in total P and total N (0.6 mg P 1^-1 and 1.3 mg N 1^-1, respectively). Changes were also observed in the benthic invertebrates; Chaoborus, oligochaetes, and chironomids all decreased, whereas Pisidium increased. It is concluded that the stocking of whitefish in eutrophic lakes for commercial purposes may delay their recovery, or even lead to enhanced eutrophication.     

Seasonal Abundance and Reproduction of the Fourwing Flyingfish, Hirundichthys affinis, off Barbados

The seasonal abundance and reproductive cycle of the flyingfish Hirundichthys affinis was studied near Barbados. H. affinis shows large seasonal variation in abundance. They are common between December and June and scarce from July to November. The population size structure of H. affinis was determined by fishing with four different mesh sizes throughout one year. There appear to be two cohorts of H. affinis present in May and June, one of immature fish with a mean size of 19.8cm FL, and one of mature fish with a mean size of 21.7cm FL. From September to April there is only a single cohort of maturing fish with a mean size that increases from 20.4cm FL in September to 22.0cm FL in April. H. affinis spawns throughout the fishing-season (December–June) and shows a peak in spawning activity from March to June. Immature fish are present in Barbados waters from July to November but in very low abundance. Surface water temperature, wind speed and swell height are the only environmental factors with which catch rate of H. affinis showed significant correlation. Catch rate appears to be negatively correlated with temperature, whereas for wind speed and swell height there are optima. Three hypotheses for the seasonal variation in abundance of H. affinis were examined. The hypothesis that this species remains dispersed near Barbados during the off-season was rejected. The other hypotheses were refined. Both post-spawning mortality, resulting in an interval of low abundance between cohorts (non-overlapping generations), and migration of fish away from Barbados at the end of the fishing season, remain possible explanations for the observed seasonal variation in abundance of this species.     

Does trimethylamine induce life-history reactions in Daphnia?

Laboratory experiments were carried out to investigate life-history reactions of Daphnia to trimethylamine (TMA), a substance, which has recently been found to induce a similar phototactic reaction in Daphnia as fish kairomones. The effects of different treatments (control, fish kairomone and TMA) on the life-history traits of five clones of D. magna and one clone of D. galeata and D. hyalina, respectively, were studied. Only D. magna exhibited significant reactions to TMA and to fish kairomone. D. galeata showed no significant responses to either TMA or to fish kairomone. D. hyalina reacted to the fish treatment with two traits (size and number of eggs at maturity). The comparison of the reaction norms to TMA and fish kairomone demonstrated that the directions of some responses to both factors were similar but were different for others such as for size at maturity. TMA resulted in a larger size at maturity in D. magna, whereas fish kairomone had no significant influence on this trait. We, therefore, conclude that trimethylamine is not the primary causative chemical agent in fish induced life-history adaptations in Daphnia.     

Seasonal variation in abundance of the Indian mackerel,Rastrelliger kanagurta Cuvier (Pisces: Scombridae) along the Zanzibar coast of East Africa

Purse-seine catches of the mackerel, Rastrelliger kanagurta Cuvier along the west coast of Zanzibar revealed seasonal patterns of abundance associated with monsoon winds. Low catches were obtained during the south-east monsoons while the north east monsoons were characterised by high catches. The largest sizes of mackerel were caught between June and August with a mixture of large and smaller ones between September and March. Mackerel in spawning condition were common between June and September with immature ones between September and January. The variations in mackerel abundance, size composition, and maturity of the fish are discussed in relation to seasonal changes, fish migrations, recruitment and food supply.     

Effect of vertebrate predation on the spatio-temporal distribution of cladocerans in a temperature eutrophic lake

We analysed the spatio-temporal distribution of zooplankton along a profile of 10 stations from the shore to the pelagic zone from April to September 1988, the period when the larvae and juveniles Rutilus rutilus, the most abundant species in the Lake, are in the littoral zone. The digestive tracts of the young roach were analysed. They fed essentially on rotifers and on cladocerans. For comparison, zooplankton was also analysed at one littoral area without fish fry. There was an increase of cladoceran density from the vegetated nearshore zone to the offshore zone. Considering the density of Bosmina longirostris, Daphnia longispina, Chydorus sphaericus and Ceriodaphnia quadrangula, we observed a different distribution pattern in the course of the year. In the nearshore zone, the relative abundance of small species, Bosmina and Chydorus, was much higher than that of the larger Daphnia. From April to September, predation pressure mainly affected the smallest species: in contrast to the inshore station without fish fry, the density of Bosmina decreased in May in the littoral with fish. Chydorus was concentrated in the littoral between February and April, then grew into the pelagic zone, where predation pressure obviously was low during the warm season. The number of Daphnia, which was eaten by the fish fry at any time, remained low in the nearshore zone, which suggests that the presence of fish may cause Daphnia to avoid this zone. Ceriodaphnia which was not affected by this predation, was scarce in the nearshore zone during mid-summer. The low density of the cladocerans in the nearshore zone is likely associated with vertebrate predation by roach fry and juveniles, the result of such a process being either a depletion in density of the prey, or an avoidance behaviour.     

Is the midsummer decline of Daphnia really induced by age-0 fish predation? Comparison of fish consumption and MDaphnia mortality and life history parameters in a biomanipulated reservoir

We analysed the temporal pattern of a Daphnia galeata populationand the development of the age-0 Ash community in a long-termbiomanipulated lake (Bautzen reservoir, Germany) during springand early summer of two successive years. In Bautzen reservoir,the age-0 Rsh-Daphnia interaction is a key process within thefood web due to the low abundance of adult zooplanktivorousfish. Daily consumption of daphnids by age-0 fish, as estimatedby a bioenergetics model, was compared to daily mortality ratesof daphnids. In addition, we estimated life history parametersof Daphnia that may indicate predation impact by fish. A midsummerdecline of daphnids occurred only in July of 1996, whereas in1995 the Daphnia biomass remained >4 mg wet weight 1^–1for the entire summer. The percentage of total Daphnia mortalitydue to fish predation before the onset of the midsummer declinewas –2% day^–1. Temporal patterns of individual size,clutch size and size at maturity also indicated that the 1996midsummer decline of daphnids was not the exclusive consequenceof age-0 fish predation. Instead, low reproductive capacityof daphnids also contributed significantly to the decline. Consequently,year-to-year variation of the Daphnia dynamics may be determinedby a fine-tuned ‘timing’ between the period of reducedfecundity and the time of the strongest predation impact byage-0 fish.     

Population dynamics and production of Daphnia hyalina Leydig and Daphnia cucullata Sars in Tjeukemeer

As part of a research programme on the food chains in Tjeukemeer, the Daphnia hyalina and Daphnia cucullata populations were studied for three successive years. To analyse the factors regulating the production of these two species, their population parameters (density, size distribution, fecundity) and population dynamics (birth rate, mortality rate) were studied and related to environmental factors. Since Daphnia in Tjeukemeer shows continuous recruitment, the population dynamics model INSTAR was developed and used to integrate field data with laboratory data on development rates and length-weight relationships. The dynamics of the Daphnia species are mainly regulated by temperature and fish predation, the latter affects both birth rate and mortality. Total annual Daphnia production was 3.1–6.9 g org. dry wt M^–2, and annual P/B ratio ranged from 25 to 40 for D. cucullata and from 45 to 49 for D. hyalina.     

Chaoborus and fish-mediated influences on Daphnia longispina population structure,dynamics and life history strategies

Summary This study examined the long term effects of predation by larvae of the midge Chaoborus and simulated fish predation on experimental Daphnia longispina populations. Chaoborus predation, relative to fish predation, led to populations composed of larger individuals as a whole, larger egg-bearing individuals, and a larger primiparous instar. Daphnia retained helmets beyond the first instar in response to the presence of Chaoborus. Both types of predation, relative to predator-free controls, reduced prey population size and rates of increase, but increased population death rates. The reduction in population size due to predation led to increased resource availability for individuals remaining in the populations and increased individual fecundity in the predation treatments. The differences noted between the Chaoborus, fish, and control treatments increased with predation intensity.     

Size-dependent habitat choice in Daphnia galeata Sars and size-structured interactions among zooplankton in a subarctic lake (lake Lombola, Norway)

Seasonal changes in vertical distribution of Daphnia galeata and other zooplankters were monitored in lake Lombola (69° 07&prime; N). Depth-habitat use, availability of edible algae and zooplankton densities were recorded to examine seasonal changes in intensity of competition between Daphnia and the other herbivores in the lake. Early in July, the exephippial generation of Daphnia aggregated near the surface, independently of body-size. In late July, when fish planktivory was expected to increase, the daphnids moved down during the day. In August, as intraspecific competition for food intensified, small and large Daphnia partitioned the water column, with larger individuals staying deeper. In September, Daphnia became dominated by large individuals, edible phytoplankton reached the seasonal minimum, and the vertical distribution of Daphnia gradually stretched out towards the surface. The observations on food availability and zooplankton densities suggest that interspecific competition intensified by the end of July. Species and stages that were most exposed to exploitative and interference competition by Daphnia were those staying deeper, because their vertical distribution overlapped more with the larger, competitively superior daphnids. These susceptible competitors included Keratella cochlearis and Synchaeta, among the rotifers, and nauplii and early copepodite stages of Cyclops scutifer. Depth-habitat use is discussed in relation to copepod development, zooplankton dynamics and predator-mediated coexistence.     

Seasonal abundance,breeding and sex-structure of populations of Tilapia nilotica (L.) and Labeo niloticus (F.) in the Jebel Aulia Dam area of the White Nile

Seasonal abundance, maturation and sex-structure were studied in populations of T. nilotica and L. niloticus in the Jebel Aulia reservoir and in the White Nile below the dam. L. niloticus showed a single annual breeding cycle from mid-July to December and was most abundant in October in the lake, and in early November in the river. T. nilotica showed two distinct annual seasons of abundance: summer (April to June) and autumn (November to October). Spawning activity was highest in summer (April to June) and autumn (late July to September). The sex-ratio varied inversely with age (size); males were predominant in populations of older fish, perhaps because of their greater endurance to adverse conditions as evidenced by their better survival in hypoxic water.     

Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

Spatial avoidance of littoral and pelagic invertebrate predators by <Emphasis Type="Italic">Daphnia</Emphasis>

Studies on spatial avoidance behaviour of predators by prey often ignored the fact that prey typically face multiple predators which themselves interact and show a spatial pattern in abundance and predation rates (PRs). In a series of laboratory experiments, we investigated predation risk (PRI) and horizontal migration of the cladoceran Daphnia magna between open water and vegetation in response to two important invertebrate predators with a contrasting spatial distribution: pelagic Choaborus and vegetation-associated Ischnura. As expected, PRI by Chaoborus was higher in open water due to higher numbers and higher PRs of Chaoborus, while for Ischnura, PRI was highest in the vegetation due to higher densities, despite lower PRs of Ischnura. In accordance with this, Daphnia moved into the vegetation in the presence of the pelagic Chaoborus alone. In the presence of Ischnura alone, however, Daphnia showed no response. We hypothesize this may be the result of a constitutive behaviour of Daphnia to avoid pelagic fish, which impedes a response to the open water. In the combined predator treatment, Daphnia migrated to the open water zone. The increased risk of predation in the vegetation, due to a facilitating effect of Chaoborus on Ischnura PRs is believed to have caused this migration of the Daphnia. This response of Daphnia declined through time and Daphnia moved toward the vegetation. A decline in the activity of the Ischnura larvae through time may have switched the risk balance in favour of the vegetation environment.