Dynamics and trophic roles of heterotrophic protists in the plankton of a freshwater tidal estuary

Freshwater tidal estuaries comprise the most upstream reaches of estuaries and are often characterised by the presence of dense bacterial and algal populations which provide a large food source for bacterivorous and algivorous protists. In 1996, the protistan community in the freshwater tidal reaches of the Schelde estuary was monitored to evaluate whether these high food levels are reflected in a similarly high heterotrophic protistan biomass. Protistan distribution patterns were compared to those of metazoan zooplankton to evaluate the possible role of top-down regulation of protists by metazoans. Apart from the algivorous sarcodine Asterocaelum, which reached high densities in summer, heterotrophic protistan biomass was dominated by ciliates and, second in importance, heterotrophic nanoflagellates (HNAN). HNAN abundance was low (annual average 2490 cells ml^–1) and did not display large seasonal variation. It is hypothesised that HNAN were top-down controlled by oligotrich ciliates throughout the year and by rotifers in summer. Ciliate abundance was generally relatively high (annual average 65 cells ml^–1) and peaked in winter (maximum 450 cells ml^–1). The decline of ciliate populations in summer was ascribed to grazing by rotifers, which developed dense populations in that season. In winter, ciliate populations were probably regulated `internally' by carnivorous ciliates (haptorids and Suctoria). Our observations suggest that, in this type of productive ecosystems, the microbial food web is mainly top-down controlled rather than regulated by food availability.     

On the quantitative characteristics of the pelagic ecosystem of Dalnee lake (Kamchatka)

The parameters of biomass and productivity of the main components of the pelagic ecosystem have been estimated in lake Dalnee (Kamchatka) during the first phase of the period of vegetation in July 1969. The water column in the lake during this period was stratified. The layer-character of vertical distribution of bacteria, phytoplankton, protozoa, rotifers and young stages of crustaceans was elucidated. The adult copepods migrate during the day. The nutrition of the mass species of zooplankton was studied using the C¹⁴-method. The data on the spectrum of feeding, on rations and on optimal food concentrations were ascertained. All these data, together with the observations in the lake, were used for the construction of a scheme of energy flow. The scheme shows that the ecosystem of the lake receives the third part of its energy from the land as allochtonous organic matter via microbial biosynthesis. The main part of energy accessible to the animals of the second trophical level is used by protozoa, and of a third part by the predatory rotiferAsplanchna.     

Relating the ontogeny of functional morphology and prey selection with larval mortality in Amphiprion frenatus

Survival during the pelagic larval phase of marine fish is highly variable and is subject to numerous factors. A sharp decline in the number of surviving larvae usually occurs during the transition from endogenous to exogenous feeding known as the first feeding stage in fish larvae. The present study was designed to evaluate the link between functional morphology and prey selection in an attempt to understand how the relationship influences mortality of a marine fish larva, Amphiprion frenatus, through ontogeny. Larvae were reared from hatch to 14 days post hatch (DPH) with one of four diets [rotifers and newly hatched Artemia sp. nauplii (RA); rotifers and wild plankton (RP); rotifers, wild plankton, and newly hatched Artemia nauplii (RPA); wild plankton and newly hatched Artemia nauplii (PA)]. Survival did not differ among diets. Larvae from all diets experienced mass mortality from 1 to 5 DPH followed by decreased mortality from 6 to 14 DPH; individuals fed RA were the exception, exhibiting continuous mortality from 6 to 14 DPH. Larvae consumed progressively larger prey with growth and age, likely due to age related increase in gape. During the mass mortality event, larvae selected small prey items and exhibited few ossified elements. Cessation of mass mortality coincided with consumption of large prey and ossification of key elements of the feeding apparatus. Mass mortality did not appear to be solely influenced by inability to establish first feeding. We hypothesize the interaction of reduced feeding capacities (i.e., complexity of the feeding apparatus) and larval physiology such as digestion or absorption efficiency contributed to the mortality event during the first feeding period. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Filtering and feeding rates of cyclopoid copepods feeding on phytoplankton

The algal biomass ingested by omnivorous cyclopoid copepods (Cyclops kolensis and C. vicinus) was measured by two methods in the hypertrophic Heiligensee in Berlin (West Germany). The clearance and ingestion rates inferred from measurements of natural populations of ¹⁴C labelled phytoplankton were compared with those obtained from chlorophyll a determinations using the presence/absence method (observed chlorophyll a content of natural lake phytoplankton with and without addition of cyclopoids). Both methods gave similar results. Nevertheless, the radio tracer method is preferred, mainly because the short feeding duration excludes high variations in both the food composition and food concentration that limit the presence/absence method.     

Relative Effect of Food Supplementation and Natural Resources on Female Red Deer Distribution in a Mediterranean Ecosystem

ABSTRACT Supplementary feeding is a widespread game management practice in several red deer (Cervus elaphus) populations, with important potential consequences on the biology of this species. In Mediterranean ecosystems food supplementation occurs in the rutting period, when it may change mating system characteristics. We studied the role of food supplementation relative to natural resources in the spatial distribution, aggregation, and mean harem size of females in Iberian red deer (Cervus elaphus hispanicus) during the rut. We studied 30 red deer populations of southwestern Spain, 63% of which experienced supplementary feeding. Using multivariate spatial analyses we found that food supplementation affected distribution of females in 95% of the populations in which it occurred. Green meadows present during the mating season acted as an important natural resource influencing female distribution. Additionally, the level of female aggregation and mean harem size were significantly higher in those populations in which food supplementation determined female distribution than in populations in which female distribution did not depend on supplementary feeding. Because female aggregation and mean harem size are key elements in sexual selection, supplementary feeding may constitute an important anthropogenic element with potential evolutionary implications for populations of Iberian red deer.     

Diel pattern of pelagic distribution and feeding in planktivorous fish

Summary The spatial distribution of juvenile roach (Rutilus rutilus), rudd (Scardinius erythrophtalamus), bream (Abramis brama) and bleak (Alburnus alburnus) was registered by echosounding during two years in small Bavarian lakes. The gut contents of the fish were analysed in order to reconstruct their rhythmicity of food intake. Fish were found in the pelagic zone only during the night, feeding before midnight with maximal rate. The vertical position of the fish was dependent on the water transparency following a light intensity in the order of 10^-3 Lux. The diel migration between littoral and pelagic zones was a mainly horizontal one with a vertical component. Both the speed of increase of fish density in the open water and of the vertical movement was correlated to the speed of change of light intensity. Shoals broke up into single fish during the offshore migration. The span of presence in the pelagic zone was identical with the duration of the dark period. During winter no diel migration nor periodical feeding took place.     

Food consumption of five deep‐sea fishes in the Balearic Basin (western Mediterranean Sea): are there daily feeding rhythms in fishes living below 1000 m?

Predation and food consumption of five deep‐sea fish species living below 1000 m depth in the western Mediterranean Sea were analysed to identify the feeding patterns and food requirements of a deep‐sea fish assemblage. A feeding rhythm was observed for Risso's smooth‐head Alepocephalus rostratus, Mediterranean grenadier Coryphaenoides mediterraeus and Mediterranean codling Lepidion lepidion. Differences in the patterns of the prey consumed suggest that feeding rhythms at such depths are linked with prey availability. The diets of those predators with feeding rhythms are based principally on active‐swimmer prey, including pelagic prey known to perform vertical migrations. The diets of Günther's grenadier Coryphaenoides guentheri and smallmouth spiny eel Polyacanthonotus rissoanus, which did not show any rhythm in their feeding patterns, are based mainly on benthic prey. Food consumption estimates were low (<1% of body wet mass day^?1). Pelagic feeding species showing diel feeding rhythms consumed more food than benthic feeding species with no feeding rhythms.     

Effect of two microalgae concentrations on body size and egg size of the rotifer <Emphasis Type="Italic">Brachionus calyciflorus</Emphasis>

The rotifer, Brachionus calyciflorus, was grown with two algae species (Chlorella sp. and Scenedesmus obliquus) at different concentrations (0.1, 1 and 10 × 10⁶ cells ml⁻¹). The body size (lorica biovolume) of individual rotifer and their egg size were measured when the populations were roughly in the exponential phase of population growth. The body size of the rotifers differed significantly (P < 0.05) among the two algae species used, however this effect was not observed for egg size. The body size of rotifers fed on higher densities of Chlorella sp. (10 × 10⁶ cells ml⁻¹) was significantly larger than for those fed on lower and medium densities (0.1 and 1 × 10⁶ cells ml⁻¹). Body size and egg size of rotifers fed with different amounts of Scenedesmus did not differ significantly. The egg size was significantly larger at higher food level of Chlorella. A significantly positive correlation was observed between the adult rotifer body size and their egg size.     

δ15N of zooplankton species in subarctic lakes in northern Sweden: effects of diet and trophic fractionation

1. To assess the use of stable nitrogen isotopes (δ¹⁵N) for reconstructing trophic relationships in planktonic food webs, crustacean zooplankton species and pelagic dissolved and particulate matter were analysed in 14 subarctic lakes in northern Sweden. The lakes are situated along an altitudinal gradient and show a substantial variation in nutrient content and energy mobilization by bacterioplankton and phytoplankton. 2. The δ¹⁵N of dissolved and particulate matter was comparatively low, suggesting efficient N recycling and low losses of depleted N from the pelagic zone of these unproductive lakes. 3. Copepods had a systematically higher δ¹⁵N than cladocerans, with an average difference of 3.1–4.9‰ within lakes, implying different trophic positions of the two groups. Comparisons of nitrogen pools and energy fluxes suggest that the low cladoceran δ¹⁵N was a result of feeding on bacteria. 4. The difference in δ¹⁵N between copepods and cladocerans declined with decreasing bacterioplankton production among lakes, due either to increasing trophic isotope fractionation or decreasing relative importance of bacteria in the diet of cladocerans.     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

Temporal and areal feeding behavior of the butterflyfish,Chaetodon trifascialis,at Johnston Atoll

Synopsis The chevron butterflyfish,Chaetodon trifascialis, is found throughout the Indo-Pacific. It is a territorial, diurnal, corallivore found in close association withAcropora spp. corals. The feeding behavior of 33 individuals was studied over six seasons in three habitats.Chaetodon trifascialis spent one third of its active time feeding. However, there was much individual variation. Fish had significantly higher feeding rates during the early afternoon, and there were no significant differences in the feeding rates between the seasons. Feeding rates were significantly different between the three habitats. TheMontipora-rich habitat had the highest feeding rates (x = 10.74 bites min^-1 ± 0.87, all corals combined) and theAcropora-Montipora mixed habitat had the lowest feeding rates (x = 4.58 bites min^-1 ± 0.63, all corals combined). Females fed significantly more than males. WhileC. trifascialis had been thought to only eatAcropora spp. corals, it occasionally fed onMontipora spp. andPocillopora sp. corals whenAcropora spp. were scarce.Chaetodon trifascialis exhibited patterns predicted by foraging theory of an energy maximizer. Territory sizes were inversely related to food density and feeding rates were inversely related to intruder rates. This is a promising system for future testing of foraging strategy models.     

Rotifers as food in aquaculture

The rotifer Brachionus plicatilis (O.F. Muller) can be mass cultivated in large quantities and is an important live feed in aquaculture. This rotifer is commonly offered to larvae during the first 7–30 days of exogenous feeding. Variation in prey density affects larval fish feeding rates, rations, activity, evacuation time, growth rates and growth efficiencies. B. plicatilis can be supplied at the food concentrations required for meeting larval metabolic demands and yielding high survival rates. Live food may enhance the digestive processes of larval predators. A large range of genetically distinct B. plicatilis strains with a wide range of body size permit larval rearing of many fish species. Larvae are first fed on a small strain of rotifers, and as larvae increase in size, a larger strain of rotifers is introduced. Rotifers are regarded as living food capsules for transferring nutrients to fish larvae. These nutrients include highly unsaturated fatty acids (mainly 20: 5 n–3 and 22: 6 n–3) essential for survival of marine fish larvae. In addition, rotifers treated with antibiotics may promote higher survival rates. The possibility of preserving live rotifers at low temperatures or through their resting eggs has been investigated.     

Anostracans and microcrustaceans as potential food sources of waterbirds on sodic pans of the Hungarian plain

Hungarian sodic water bodies have a rich macro- and microcrustacean fauna due to the lack of fish populations. The crustacean population is very abundant, for this reason these wetlands provide good feeding resources for waterbirds. The density of macro- and microcrustacean populations together with feeding waterbirds was investigated in March, April, and May of 2002, on two characteristic sodic pans, “Kelemen-szék” and “Zab-szék”. The following dabbling-filtering waterfowls and pelagic forager wader species were counted: northern pintail (Anas acuta), northern shoveler (Anas clypeata), garganey (Anas querquedula), common teal (Anas crecca), avocet (Recurvirostra avosetta), spotted redshank (Tringa erythropus), greenshank (Tringa nebularia), and marsh sandpiper (Tringa stagnatilis). The dominant macrocrustacean species was the [Anostraca – Branchinectidae] natronophile Branchinecta orientalis, and its density was significantly higher in Zab-szék than in Kelemen-szék. The microcrustacean zooplankton community was also different in the pans, [Cladocera] Daphnia magna density was significantly higher in Kelemen-szék than in Zab-szék, but the density of the [Copepoda] natronophile Arctodiaptomus spinosus, was higher in Zab-szék than in Kelemen-szék. The density of the investigated waterbird species was also significantly higher in Zab-szék than in Kelemen-szék during spring. We can conclude that the macrocrustacean B. orientalis is one of the most important potential food resources for migrating pelagic foraging waders in spring on characteristic Hungarian sodic pans. However, the most abundant available food item for waterbirds are copepod microcrustacean zooplankton, which have a biomass that is larger by approximately one order of magnitude than the macrocrustacean zooplankton biomass. Considering the lack of submerged water vegetation, we suggest that planktonic microcrustaceans are an important food resource for dabbling-filtering ducks because they can utilise the small crustacean biomass more effectively than the less abundant and rapidly moving macrocrustacean B. orientalis.     

Characterization of the bacterial flora of mass cultivated Brachionus plicatilis

Bacterial density and composition in association of mass cultivated rotifers (Brachionus plicatilis, SINTEF-strain) was investigated, during experimental conditions identical to the procedures used for preparing rotifers as live food for marine cold water fish larvae. These procedures include cultivation, enrichment with squid meal and acclimation to low temperature by storage of the rotifer culture at 6 °C. Large variations were observed in the number of rotifer associated (1.8–7.6 · 10³ colony forming units per rotifer^–1) and free-living (0.6–25 10⁷ cells·ml^–1) bacteria. An increase of 50–150% in the bacterial number was normally observed after feeding the rotifer with squid meal, but after three days of acclimation at 6 °C, the bacterial numbers decreased to the initial level.After enrichment of the cultures with squid meal, the similarity in the composition of the bacterial flora between the rotifers and water was reduced. However, acclimation of the culture at 6 °C resulted in better agreement of the rotifer associated flora and that in water. Enrichment of the cultures induced a shift in the bacterial composition from Cytophaga/Flavobacterium dominance to Pseudomonas/Alcaligenes dominance. The bacterial flora of the rotifer cultures are dominated by presumably opportunistic species after enrichment, which may have detrimental effects when rotifers are fed as live food to marine fish larvae.     

Evaluation of bacterivory of Rotifera based on measurements of in situ ingestion of fluorescent particles, including some comparisons with Cladocera

Bacterivory of pelagic rotifers and cladocerans in eutrophicLake Loosdrecht (The Netherlands) was determined by microscopicobservation of in situ tracer particle uptake. Ingestion ratesof rotifer species using 0.51 µm microspheres or fluorescentlylabelled bacteria as tracers differed, with one exception. Theingestion rates depended on both the species and the tracertype. For cladocerans, fluorescently labelled bacteria seemedto underestimate grazing, presumably due to rapid digestionof tracer cells. Comparing results obtained with 0.51 µmmicrospheres, rotifers were much more important grazers on bacteriathan cladocerans in the study period (April-September). Basedon microspheres, the rotifer populations with the highest uptakeof bacteria were Filinia longiseta (May-July) andAnuraeopsisfissa (June-September). According to the uptake of fluorescentlylabelled bacteria, Conochilus unicornis had the highest uptakein June and A.fissa in July.     

Food sources and lipid retention of zooplankton in subarctic ponds

1. Subarctic ponds are seasonal aquatic habitats subject to short summers but often have surprisingly numerous planktonic consumers relative to phytoplankton productivity. Because subarctic ponds have low pelagic productivity but a high biomass of benthic algae, we hypothesised that benthic mats provide a complementary and important food source for the zooplankton. To test this, we used a combination of fatty acid and stable isotope analyses to evaluate the nutritional content of benthic and pelagic food and their contributions to the diets of crustacean zooplankton in 10 Finnish subarctic ponds. 2. Benthic mats and seston differed significantly in total lipids, with seston (62.5 μg mg^?1) having approximately eight times higher total lipid concentrations than benthic mats (7.0 μg mg^?1). Moreover, the two potential food sources differed in their lipid quality, with benthic organic matter completely lacking some nutritionally important polyunsaturated fatty acids (PUFA), most notably docosahexaenoic acid and arachidonic acid. 3. Zooplankton had higher PUFA concentrations (27–67 μg mg^?1) than either of the food sources (mean benthic mats: 1.2 μg mg^?1; mean seston: 9.9 μg mg^?1), indicating that zooplankton metabolically regulate their accumulation of PUFA. In addition, when each pond was evaluated independently, the zooplankton was consistently more ¹³C‐depleted (δ¹³C ?20 to ?33‰) than seston (?23 to ?29‰) or benthic (?15 to ?27‰) food sources. In three ponds, a subset of the zooplankton (Eudiaptomus graciloides, Bosmina sp., Daphnia sp. and Branchinecta paludosa) showed evidence of feeding on both benthic and planktonic resources, whereas in most (seven out of 10) ponds the zooplankton appeared to feed primarily on plankton. 4. Our results indicate that pelagic primary production was consistently the principal food resource of most metazoans. While benthic mats were highly productive, they did not appear to be a major food source for zooplankton. The pond zooplankton, faced by strong seasonal food limitation, acquires particular dietary elements selectively.     

Evidence of coprophagy in freshwater zooplankton

1. Vertical transport of nutrients in sedimenting faecal material is greatly reduced by coprophageous organisms. Unfortunately, nearly all work on faecal production, sedimentation and coprophagy has dealt with copepods in marine ecosystems. Here, we report the first evidence of coprophagy in freshwater zooplankton from oligotrophic and eutrophic lakes. We used ¹⁴C‐labelled algae and faecal material to estimate the rates of algal clearance and coprophagy. 2. Measured feeding rates per individual on faecal material were similar (Daphnia pulex, D. rosea, Leptodiaptomus tyrelli) or even higher (D. lumholtzi) than filtering rates on phytoplankton. This finding does not necessarily implicate active selection of faeces over algae because: (i) we did not use the same food concentrations for faeces and algae, and (ii) grazers of slightly different sizes were used in each test. 3. Weight‐specific clearance rates of L. tyrelli and Holopedium gibberum on faecal matter (0.084–0.089 mL μg^?1 h^?1) were higher than in the daphniids (0.026 mL μg^?1 h^?1). 4. The data indicate that coprophagy in freshwater ecosystems is an important mechanism of nutrient recycling, and this process should be taken into account when studying nutrient fluxes within lakes and reservoirs.     

Creating conditions for changes in prey community structure by Chaoborus spp. in a lake in Sweden

In Lake G»rdsjön (Southwest Sweden), liming as an experimental improvement of living conditions for pelagic algae, resulted in a significant increase of algal biomass and a reduction of mean cell size. The algal development was beneficial for small sized filter feeding zooplankton, particularly rotifers, which showed a significant increase. The increase in abundance of small sized zooplankton created better food conditions for the smaller instars, and thus a much better overall survival of Chaoborus larvae. The resulting, 6–7 times larger population of Chaoborus larvae significantly changed the structure of the crustacean zooplankton community. Bosmina coregoni, the fastest swimmer of the crustacean species suffered most and was strongly reduced by the increased predation from Chaoborus. The share of cladocerans decreased, while copepods increased in importance.     

Combined effects of food level and inoculation density on competition between Brachionus patulus (Rotifera) and the cladocerans Ceriodaphnia dubia and Moina macrocopa

Competition among cladocerans and rotifers is of considerable interest not only due to their close similarity in life history strategies, but also due to the considerable overlap they exhibit in their feeding habits. In tropical waterbodies, several genera of cladocerans, including Ceriodaphnia and Moina occur, simultaneously with rotifers. We tested over a period of 3 weeks the combined effects of food (0.5×10⁶ and 1.5×10⁶ cells ml^–1 of Chlorella) level and rotifer density on the competition between B. patulus and C. dubia and M. macrocopa using population growth experiments. For each cladoceran species we used 30 test jars of 50 ml capacity. The initial density of cladocerans was 0.2 ind ml^–1, while for B. patulus it was either 1 ind ml^–1 or 5 ind ml^–1. Neither the maximal population density nor the rate of population increase (r) of C. dubia was significantly affected by B. patulus. However, for M. macrocopa, both these variables were negatively affected by the rotifers. The combined effects of low food level and high initial density of B. patulus resulted in a 50% reduction in the peak population density of M. macrocopa. The population growth of B. patulus was negatively influenced by the presence of C. dubia and M. macrocopa. The results of the competition experiments conducted in the present study between cladocerans and rotifers suggest the existence of a more complex and delicate interaction than is generally thought.     

Chlorophyll budget in a productive tropical pond: algal production,sedimentation, and grazing by microzooplankton and rotifers

Chlorophyll a and pheopigment standing stocks and fluxes were used during a two weeks colonization experiment in a productive tropical pond (Layo, Côte d'Ivoire) in order to establish a chlorophyll budget. The experiment started from an azoïc state (the pond was dried, limed and progressively filled with ground water). Algal production was the only input to the phytoplanktonic system, while grazing and algal sedimentation were the main outputs. Chlorophyll a reflected the algal biomass, and degradation pigments were considered as an index of grazing by zooplankton (here, protozoans and rotifers). An estimation of the input through the algal growth rate was performed for the two main biological events observed during the study. The first algal bloom, with a large picoplankton participation, was mainly regulated by microzooplankton (increase of the peak) and rotifers (decrease of the peak). The second bloom (exclusively nanoplankton) was regulated by rotifers (increase) and by sedimentation of living cells (decrease). This last process was related to a sudden exhaustion of ammonia in the water column. Because of the time-lag between algal proliferation and zooplanktonic bloom, the phytoplanktonic biomass was able to be adjusted according to the availability of nutrients. This self-regulation took the form of sinking of active algal cells, resulting in a transient reduction of the food available for rotifers. This process had drastic consequences in these shallow waters, since a major part of the phytoplankton produced was removed from the pelagic system. For an optimal exploitation of the natural resources of an aquaculture pond, a study of the equilibrium nutrients-phytoplankton-zooplankton would provide a basis for artificial intervention, with a view to limit the impact of this mode of natural regulation.