Daphnia population growth but not moulting is a substantial phosphorus drain for phytoplankton

SUMMARY 1. Negative effects of zooplankton on the availability of phosphorus (P) for phytoplankton as a result of the retention of nutrients in zooplankton biomass and the sedimentation of exoskeletal remains after moulting, have been recently proposed. 2. In a mesocosm study, the relative importance of these mechanisms was tested for the freshwater cladoceran Daphnia hyalina×galeata. A total of 13 mesocosm bags was suspended in a mesotrophic German lake during summer 2000 and fertilised with inorganic P in order to obtain a total nitrogen to total P ratio closer to the Redfield ratio. D. hyalina×galeata was then added at a logarithmically scaled density gradient of up to 40 ind. L^?1. Zooplankton densities, dissolved inorganic, particulate organic (seston <100 μm), as well as total nutrient concentrations were monitored. Additionally, nutrient concentrations of sediment water removed from the bottom of the mesocosm bags via a manual pump were determined. 3. Seston carbon (C), seston P and total P were significantly negatively correlated with Daphnia densities. The amount of particulate P (～5–6 μg P L^?1) sequestered from the seston compartment by Daphnia corresponded roughly to the increase of zooplankton biomass (population growth). Soluble reactive phosphorous (SRP) was at all times high (～25–35 μg P L^?1) and possibly unavailable to phytoplankton as a result of P adsorption to calcite during a calcite precipitation event (whiting). P concentrations determined in sediment water were generally <60 μg P m^?2 and thus never exceeded 1% of the total amount of P bound in particulate matter of the overlying water column. 4. Seston C : P ratios followed a polynomial second‐order function: At Daphnia densities <40 ind. L^?1 a positive linear relationship was evident, which is explained by the stronger reduction of P compared with C in seston, and transfer of seston P to zooplankton. Highest seston C : P ratios of ～300 : 1 were observed at Daphnia densities of ～30–50 ind. L^?1, which is in agreement with proposed threshold values limiting Daphnia reproductive growth. At Daphnia densities >40–50 ind. L^?1 C : P ratios were decreased because of the strong reduction of seston C at close to constantly low seston P‐values of ～3–4 μg P L^?1. 5. At least for Daphnia, it may be concluded that – unlike population growth – the sedimentation of faecal pellets and carapaces after moulting seem negligible processes in pelagic phosphorus dynamics.     

Effects of zooplankton on the sinking flux of organic carbon in Lake Biwa

To clarify the roles of zooplankton in the sedimentation of seston from the epilimnion, the sinking flux of particulate carbon was measured along with primary production rate and zooplankton biomass from July 1996 to October 1997 at a pelagic site in the north basin of Lake Biwa. During the study period, the flux varied seasonally from 66 to 510 mg C m⁻² day⁻¹ and was low in summer when zooplankton, composed mainly of Eodiaptomus japonicus and Daphnia galeata, were abundant. Simple correlation analysis revealed that the sinking flux correlated neither with the primary production rate nor with the amount of sestonic carbon above the sediment trap. However, the particle elimination rate, estimated as the difference between the primary production rate and the sinking flux, correlated positively with the zooplankton biomass. These results suggest that zooplankton play a substantial role in decreasing the sinking flux in Lake Biwa. Received: March 6, 2000 / Accepted: October 7, 2000     

Dynamics of a Boreal Lake Ecosystem during a Long-Term Manipulation of Top Predators

We assessed the long-term (16 years) effects of introducing piscivores (northern pike) into a small, boreal lake (Lake 221, Experimental Lakes Area) containing abundant populations of two planktivorous fish species. After the introduction, pearl dace were extirpated and yellow perch abundance was greatly reduced. Daphnia species shifted from D. galeata mendota to larger bodied Daphnia catawba, but the total zooplankton biomass did not increase, nor did the biomass of large grazers such as Daphnia. Phytoplankton biomass decreased after the northern pike introduction, but increased when northern pike were partially removed from the lake. Phosphorus (P) excretion by fish was ∼0.18 mg P m⁻² d⁻¹ before pike addition, declined rapidly to approximately 0.03–0.10 as planktivorous perch and dace populations were reduced by pike, and increased back to premanipulation levels after the pike were partially removed and the perch population recovered. When perch were abundant, P excretion by fish supported about 30% of the P demand by primary producers, decreasing to 6–14% when pike were abundant. Changes in phytoplankton abundance in Lake 221 appear to be driven by changes in P cycling by yellow perch, whose abundance was controlled by the addition and removal of pike. These results confirm the role of nutrient cycling in mediating trophic cascades and are consistent with previous enclosure experiments conducted in the same lake.     

AUTOLYSIS KINETICS OF THE MARINE DIATOM DITYLUM BRIGHTWELLII (BACILLARIOPHYCEAE) UNDER NITROGEN AND PHOSPHORUS LIMITATION AND STARVATION1

Autolysis kinetics in axenic cultures of the diatom Ditylum brightwellii (West) Grunow were studied under nutrient limitation in continuous cultures and under nutrient starvation in batch-mode cultures obtained by switching off nutrient supply in the continuous cultures. Under N limitation, the specific algal autolysis rates (δ, day^?1) were found constant at 0.014 ± 0.002 day^?1over a broad range of specific dilution rates (D, day^?1) (0.09–0.56 day^?1), implying an intrinsic death factor independent of the physiologzc state of the algal cells. Under P limitation, 8 was inversely related to D and ranged between 0.067 and 0.005 day^?1 at D = 0.17–0.44 day^?1. Under conditions of nutrient stamation, the degree of algal nutrient deficiency prior to stamation affected autolysis rates (δ_b, day^?1) and subsequently survival of the algal cultures. Nitrogen-starved D. brightwellii showed highest δ_b (maximum, 0.10 day^?1) when precultured at the higher growth rates. Phosphorus stamation led to highest δ_b (maximum, 0.21 day^?1) in the cultures preconditioned at the lower steady state growth rates. The lower death rates for D. brightwellii under limitation and starvation of N compared to P suggest that D. brightwellii was better equipped to handle N than P deficiency. The present results showed that cell lysis induced by nutrient stress was a significant cause of mortality in D. brightwellii and provided more insight into the field distribution of this neritic diatom.     

The evaluation of the role of pelagic invertebrate versus vertebrate predators on the seasonal dynamics of filtering Cladocera in a shallow, eutrophic reservoir

An evaluation of the role of invertebrate (Leptodora kindtii) versus vertebrate (juvenile fish) predators in the dynamics of filtering zooplankton (Daphnia sp.) in the pelagic zone of the Sulejow Reservoir was conducted during summer from 1994 to 1996. Leptodora kindtii appeared in high densities (mid-July: 2–6 ind. l^?1) in the pelagic zone. Its predation impact on the dominant filtering zooplankter population Daphnia cucullata, expressed by consumption, was significant and accounted for a reduction of 10 to 51% of Daphnia biomass. Although D. cucullata was a preferred food of juvenile pikeperch in June, due to low biomass, these fish were able to eliminate daily only 0.1–5.4% of Daphnia biomass in the pelagic zone. Thus, the role of juvenile fish in the midsummer decline of D. cucullata density in the pelagic zone seems to be much smaller than that of Leptodora kindtii.     

Secondary production of crustacean zooplankton and biomass of major rotifer species in Lake Bosumtwi/Bosomtwe,Ghana, West Africa

Studies have shown a strong linkage between zooplankton and fisheries' potential in tropical lakes. High zooplankton production provides the basis for fish production, but knowledge of zooplankton production dynamics in African lakes is extremely limited. Crustacean zooplankton production and the biomass of dominant rotifers in Lake Bosumtwi were assessed over a 2‐year period. The crustaceans comprised an endemic and extremely abundant cyclopoid copepod, Mesocyclops bosumtwii and the cladoceran Moina micrura. Mean standing stock of the crustaceans was 429 mg dw m^?3, whilst annual production averaged 2.1 g dw m^?3 y^?1. Production doubled from 1.4 g dw m^?3 y^?1 in 2005 to 2.8 g dw m^?3 y^?1 in 2006. Copepods accounted for 98.5% of crustacean production. The biomass of the dominant rotifers Brachionus calyciflorus and Hexarthra intermedia was less than 1% of total zooplankton biomass. Daily turnover rate and turnover time of the crustaceans was 0.19 day^?1 and 6.2 days respectively. Crustacean production yielded no statistical relationship with phytoplankton biomass. Production was well within the range of tropical lakes. Peak crustacean production synchronized maximum rainfall, lake mixing and phytoplankton production. Most importantly, no one year's set of dynamics can be used to characterize zooplankton production in the lake.     

The role of phosphorus release by roach [Rutilus rutilus (L.)] in the water quality changes of a biomanipulated lake

SUMMARY 1. Following fish removal, the water quality in biomanipulated lakes often improves concomitant with decreased phosphorus (P) levels. Because the decrease in P concentrations derives most probably either directly or indirectly from fish, which are the main target of biomanipulation, this study examined the P release of 0+, 1+ and 2+ roach [Rutilus rutilus (L.)] and changes in the P release during summer in a shallow eutrophic lake in Finland. 2. The P release was separated into P derived from benthic and littoral food items and into recycled P derived from feeding on zooplankton, to estimate the contribution of net P additions to the water column by the fish to the increase in P concentrations of the lake water (75–110 mg P m^?3) in summer 1991–96. 3. Individual P release of roach by both egestion and excretion was estimated with a bioenergetics model. The size of the roach population was estimated with a depletion method and the proportions of different age groups from catch samples, using a programme separating mixtures of normal distributions. The sensitivity of the release estimates to variation in the growth data was estimated with the jackknife technique. 4. The biomass‐specific P release by 0+ roach (0.36–0.54 mg P g^?1 day^?1) was higher than that by older roach (0.07–0.16 mg P g^?1 day^?1) throughout the summer. The P release by the whole roach population deriving from benthic and littoral food items (0.7–2.7 mg m^?3 during July to August, representing a net addition to the water column) was 5–19 times lower in 1991–96 than the recycled P release deriving from zooplankton (8.9–25.7 mg m^?3), and too low to explain the increase in the P concentration of the lake water during the summer. Because the biomass‐specific P release and roach diet composition vary with fish age, it is important to consider the age structure of fish populations to obtain correct estimates of P release and net additions to the water column. 5. The removal of roach by fishing diminished the roach stock greatly, but the fish‐mediated P release to the water column changed little. This effect was because of the high compensation capacity of the roach population, leading to high recruitment of young fish with higher biomass‐specific P release rates. 6. External loading is very low during summer months and therefore it cannot explain the increase in the P concentration of water during that time. Internal loading from the sediment might be as high as 10.2 mg P m^?2 day^?1, i.e. 50 times higher than the maximum net P addition by the total roach population.     

Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments

1. Invasion of tropical zooplankton into temperate lakes provides an interesting opportunity to explore habitat segregation in a thermal gradient. 2. We explored differing vertical positioning of native and exotic Daphnia (Daphnia mendotae and Daphnia lumholtzi) in a large indoor mesocosm system (Plön plankton towers) during 2 month‐long experiments. The two towers were manipulated to provide gradients of both temperature (15–29 °C) and algal food (0.05–0.58 mg C L^?1) and a day–night cycle. 3. Both juvenile and adult D. lumholtzi showed a ‘typical’ vertical migration pattern, with higher densities in the epilimnion at night than during the day. They avoided the food‐poor middle layer. In contrast, D. mendotae adults showed little tendency to migrate into the epilimnion at night, remaining in the cooler hypolimnion while juveniles migrated. The vertical distribution of D. mendotae appeared unaffected by the presence of D. lumholtzi. 4. The strong migration behaviour of D. lumholtzi in the absence of fish cues suggests that this behaviour may be a constitutive trait. Habitat partitioning of the two species is probably the result of different thermal tolerances, with D. mendotae constrained to remaining in deeper water by high temperatures in the epilimnion and the tropical D. lumholtzi able to use the warm epilimnion at night.     

How<Emphasis Type="Italic"> Daphnia</Emphasis> copes with excess carbon in its food

Animals that maintain near homeostatic elemental ratios may get rid of excess ingested elements from their food in different ways. C regulation was studied in juveniles of Daphnia magna feeding on two Selenastrum capricornutum cultures contrasting in P content (400 and 80 C:P atomic ratios). Both cultures were labelled with ¹⁴C in order to measure Daphnia ingestion and assimilation rates. No significant difference in ingestion rates was observed between P-low and P-rich food, whereas the net assimilation of ¹⁴C was higher in the treatment with P-rich algae. Some Daphnia were also homogeneously labelled over 5 days on radioactive algae to estimate respiration rates and excretion rates of dissolved organic C (DOC). The respiration rate for Daphnia fed with high C:P algae (38.7% of body C day^-1) was significantly higher than for those feeding on low C:P algae (25.3% of body C day^-1). The DOC excretion rate was also higher when animals were fed on P-low algae (13.4% of body C day^-1) than on P-rich algae (5.7% of body C day^-1) . When corrected for respiratory losses, total assimilation of C did not differ significantly between treatments (around 60% of body C day^-1). Judging from these experiments, D. magna can maintain its stoichiometric balance when feeding on unbalanced diets (high C:P) primarily by disposing of excess dietary C via respiration and excretion of DOC.     

Répartition verticale du phytoplancton,des bactéries et du zooplancton dans un milieu stratifié en Baie de Biétri (Lagune Ebrié, Cote d'Ivoire). Relations trophiques

During the period of September–November 1981, a stratified condition was observed in Biétri Bay, a part of Ebrié Lagoon (Ivory Coast, Africa). The epilimnion was oxidised and in the hypolimnion, high concentrations of hydrogen sulfide were noted (200–400 mg · l^?1). At the surface of the hypolimnion, a large population of phototrophic bacteria developed, constituting a brown layer composed of the purple and green bacteria Rhodopseudomonas sp., Chromatium gracile, Chlorobium vibrioforme, C. phaeobacteroides and Pelodictyon sp. These bacteria can be considered as particulate organic matter producers. Their production is estimated at about 1 530 mg C · m^?2 · day^?1. In the epilimnion, the algal production is 2 200 mg C · m^?2 · day^?1. Therefore, phototrophic bacterial production represented 41% of the total photosynthetic production. In the epilimnion, the zooplankton community (composed of copepods, rotifers and some cyclopids) was particularly concentrated near the chemocline where only low concentration of dissolved oxygen was available. Analysis of gut contents of the copepod Acartia clausi, which is the dominant species of the zooplankton, shows the importance of phototrophic bacteria and especially Rhodospirillaceae and Chromatium in its diet. This copepod seems to prefer phototrophic bacteria to both heterotrophic bacteria and phytoplancton. The vertical migration of the zooplankton community is partly conditioned by the search for nutrients i.e. phototrophic bacteria. Thus, bacteria contribute to the first trophic level of the food chain in this lagoon. Since they derive part of their energy from organic matter formed by primary producers, they cannot be considered as pure primary producers. However, in respect to the food chain they form food material for secondary producers (zooplankton).     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Goose-mediated nutrient enrichment and planktonic grazer control in arctic freshwater ponds

A dramatic increase in the breeding population of geese has occurred over the past few decades at Svalbard. This may strongly impact the fragile ecosystems of the Arctic tundra because many of the ultra-oligotrophic freshwater systems experience enrichment from goose feces. We surveyed 21 shallow tundra ponds along a gradient of nutrient enrichment based on exposure to geese. Concentrations of total phosphorus (P) and dissolved inorganic nitrogen (DIN) in the tundra ponds ranged from 2–76 to 2–23 μg l⁻¹ respectively, yet there was no significant increase in phytoplankton biomass (measured as chlorophyll a; range: 0.6–7.3 μg l⁻¹) along the nutrient gradient. This lack of response may be the result of the trophic structure of these ecosystems, which consists of only a two-trophic level food chain with high biomasses of the efficient zooplankton grazer Daphnia in the absence of fish and scarcity of invertebrate predators. Our results indicate that this may cause a highly efficient grazing control of phytoplankton in all ponds, supported by the fact that large fractions of the nutrient pools were bound in zooplankton biomass. The median percentage of Daphnia–N and Daphnia–P content to particulate (sestonic) N and P was 338 and 3009%, respectively, which is extremely high compared to temperate lakes. Our data suggest that Daphnia in shallow arctic ponds is heavily subsidized by major inputs of energy from other food sources (bacteria, benthic biofilm), which may be crucial to the persistence of strong top–down control of pelagic algae by Daphnia.     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Influence of diet shifts in underyearling fish on phosphorus recycling in a hypertrophic biomanipulated reservoir

1. In addition to effects of direct predation by planktivorous fish, nutrient recycling by fish may also contribute to structuring foodwebs in lakes. There is little evidence, however, about whether underyearling fish undergoing several ontogenetic diet shifts may have a comparable bottom-up impact. 2. This study examined seasonal patterns of phosphorus (P) concentration and external load, phytoplankton, zooplankton and benthos, and diet shifts in three underyearling fish [perch (Perca fluviatilis), roach (Rutilus rutilus) and ruffe (Gymnocephalus cernuus)] in the shallow, hypertrophic biomanipulated Bautzen reservoir, Germany. Phosphorus metabolism of fish was calculated by a balanced bioenergetics model on the basis of fish diet, growth and water temperature. 3. The fish showed several shifts from planktivory to other food sources during the sampling period from May to September. These shifts were probably caused by the seasonal succession of the zooplankton community, mainly the midsummer decline of Daphnia galeata. 4. The diet shifts in fish also had consequences for the amount of P consumed and released. During periods of dominant zooplanktivory, the excretion of P did not exceed the removal of P stored in pelagic prey. By contrast, if benthivory dominated, fish subsidized the pelagic P pool by excreting more P from benthic prey than had been removed from the pelagic area. This occurred predominantly in perch and ruffe during periods of low zooplankton biomass, whereas the roach ate more algae and therefore excreted less P of benthic origin. 5. Phosphorus release by underyearling fish was estimated at a maximum of 0.1 mg m^–3 JY day^–1. This value was negligible compared with both the external load of P to Bautzen reservoir and the concentration of P in the pelagic area during summer. It is therefore concluded that both the predominance of underyearling zooplanktivorous fish and the high Daphnia biomass during certain periods of the year in the Bautzen reservoir may be the reason that nutrient release by the fish structured the foodweb only marginally. 6. This study suggests that biomanipulation has altered both top-down and bottom-up impacts of fish in Bautzen reservoir. The highest efficiency of foodweb manipulations may be obtained after reduction of the external P loading below a certain threshold. In turn, if external restoration of eutrophied lakes is not accompanied by changes in fish community, then the combined forces of strong zooplanktivory and high P recycling of dense stocks of zooplanktivorous and benthivorous fish may hold the water in a eutrophic-like stage, even if external load has been significantly reduced.     

Strong interactions between stoichiometric constraints and algal defenses: evidence from population dynamics of Daphnia and algae in phosphorus-limited microcosms

The dynamic interactions among nutrients, algae and grazers were tested in a 2 × 3 factorial microcosm experiment that manipulated grazers (Daphnia present or absent) and algal composition (single species cultures and mixtures of an undefended and a digestion-resistant green alga). The experiment was run for 25 days in 10-L carboys under mesotrophic conditions that quickly led to strong phosphorus limitation of algal growth (TP ? 0.5 μM, N:P 40:1). Four-day Daphnia juvenile growth assays tested for Daphnia P-limitation and nutrient-dependent or grazer-induced algal defenses. The maximal algal growth rate of undefended Ankistrodesmus (mean ± SE for three replicate microcosms; 0.92 ± 0.02 day^?1) was higher than for defended Oocystis (0.62 ± 0.03 day^?1), but by day 6, algal growth was strongly P-limited in all six treatments (molar C:P ratio >900). The P-deficient algae were poor quality resources in all three algal treatments. However, Daphnia population growth, reproduction, and survival were much lower in the digestion-resistant treatment even though growth assays provided evidence for Daphnia P-limitation in only the undefended and mixed treatments. Growth assays provided little or no support for simple threshold element ratio (TER) models that fail to consider algae defenses that result in viable gut passage. Our results show that strong P-limitation of algal growth enhances the defenses of a digestion-resistant alga, favoring high abundance of well-defended algae and energy limitation of zooplankton growth.     

Benthic resources are the key to Daphnia middendorffiana survival in a high arctic pond

1. Shallow arctic lakes and ponds have simple and short food webs, but large uncertainties remain about benthic–pelagic links in these systems. We tested whether organic matter of benthic origin supports zooplankton biomass in a pond in NE Greenland, using stable isotope analysis of carbon and nitrogen in the pond itself and in a ¹³C‐enrichment enclosure experiment. In the latter, we manipulated the carbon isotope signature of benthic algae to enhance its isotopic discrimination from other potential food sources for zooplankton. 2. The cladoceran Daphnia middendorffiana responded to the ¹³C‐enrichment of benthic mats with progressively increasing δ¹³C values, suggesting benthic feeding. Stable isotope analysis also pointed towards a negligible contribution of terrestrial carbon to the diet of D. middendorffiana. This agreed with the apparent dominance of autochthonous dissolved organic matter in the pond revealed by analysis of coloured dissolved organic matter. 3. Daily net production by phytoplankton in the pond (18 mg C m^?2 day^?1) could satisfy only up to half of the calculated minimum energy requirements of D. middendorffiana (35 mg C m^?2 day^?1), whereas benthic primary production alone (145 mg C m^?2 day^?1) was more than sufficient. 4. Our findings highlight benthic primary production as a major dietary source for D. middendorffiana in this system and suggest that benthic organic matter may play a key role in sustaining pelagic secondary production in such nutrient‐limited high arctic ponds.     

High rates of net primary production and turnover of floating grasses on the Amazon floodplain: implications for aquatic respiration and regional CO2 flux

We investigated whether rates of net primary production (NPP) and biomass turnover of floating grasses in a central Amazon floodplain lake (Lake Calado) are consistent with published evidence that CO₂ emissions from Amazon rivers and floodplains are largely supplied by carbon from C4 plants. Ground‐based measurements of species composition, plant growth rates, plant densities, and areal biomass were combined with low altitude videography to estimate community NPP and compare expected versus observed biomass at monthly intervals during the aquatic growth phase (January–August). Principal species at the site were Oryza perennis (a C3 grass), Echinochloa polystachya, and Paspalum repens (both C4 grasses). Monthly mean daily NPP of the mixed species community varied from 50 to 96 g dry mass m^?2 day^?1, with a seasonal average (±1SD) of 64±12 g dry mass m^?2 day^?1. Mean daily NPP (±1SE) for P. repens and E. polystachya was 77±3 and 34±2 g dry mass m^?2 day^?1, respectively. Monthly loss rates of combined above‐ and below‐water biomass ranged from 31% to 75%, and averaged 49%. Organic carbon losses from aquatic grasses ranged from 30 to 34 g C m^?2 day^?1 from February to August. A regional extrapolation indicated that respiration of this carbon potentially accounts for about half (46%) of annual CO₂ emissions from surface waters in the central Amazon, or about 44% of gaseous carbon emissions, if methane flux is included.     

Contributions of Different Organic Carbon Sources to <Emphasis Type="Italic">Daphnia</Emphasis> in the Pelagic Foodweb of a Small Polyhumic Lake: Results from Mesocosm DI<Superscript>13</Superscript>C-Additions

Abstract Freshwater ecosystems derive organic carbon from both allochthonous and autochthonous sources. We studied the relative contributions of different carbon sources to zooplankton in a small, polyhumic, steeply stratified lake, using six replicate surface-to-sediment enclosures established during summer and autumn 2004. We added ¹³C-enriched bicarbonate to the epilimnion of half the enclosures for three weeks during each season and monitored carbon stable isotope ratios of DIC, DOC, POC and Daphnia, along with physical, chemical and biological variables. During summer, ¹³C-enriched DIC (δ¹³C up to 44 ± 7.2‰) was soon taken up by phytoplankton (δ¹³C up to −5.1 ± 13.6‰) and was transmitted to Daphnia (δ¹³C up to −1.7 ± 7.2‰), demonstrating consumption of phytoplankton. In contrast, during autumn, ¹³C-enriched DIC (δ¹³C up to 56.3 ± 9.8‰) was not transmitted to Daphnia, whose δ¹³C became progressively lower (δ¹³C down to −45.6 ± 3.3‰) concomitant with decreasing methane concentration. Outputs from a model suggested phytoplankton contributed 64–84% of Daphnia diet during summer, whereas a calculated pelagic carbon mass balance indicated only 30–40% could have come from phytoplankton. Although autumn primary production was negligible, zooplankton biomass persisted at the summer level. The model suggested methanotrophic bacteria contributed 64–87% of Daphnia diet during autumn, although the calculated carbon mass balance indicated a contribution of 37–112%. Thus methanotrophic bacteria could supply virtually all the carbon requirement of Daphnia during autumn in this lake. The strongly ¹³C-depleted Daphnia values, together with the outputs from the models and the calculated carbon mass balance showed that methanotrophic bacteria can be a greater carbon source for Daphnia in lakes than previously suspected.     

Prey preference and utilization by Mesostoma lingua (Turbellaria,Rhabdocoela) at a low arctic site

The rhabdocoel Mesostoma lingua is a common inhabitant of rock bluff ponds in the area of Churchill, Manitoba. Unlike most flatworms it has the ability to prey actively on zooplankton. Among the zooplankton, Mesostoma prefers Daphnia pulex, Simocephalus vetulus, and Diaptomus tyrrelli. Larger calanoid copepods are difficult to capture and take much longer to consume. Growth in Mesostoma is maintained when individuals consume 0.5 Daphnia day^–1 and is maximal at 4 Daphnia day^–1. In comparison, Mesostoma maintained growth only when offered the copepod Diaptomus victoriaensis at the rate of 8–10 individuals-day^–1. Mesostoma may be an important agent in structuring pond zooplankton communities in the arctic, producing a shift in dominance away from cladocerans such as Daphnia toward larger calanoid copepods.     

Effects of benthivorous bream (Abramis brama) and carp (Cyprinus carpio) on sediment resuspension and concentrations of nutrients and chlorophyll a

1. The effect of benthivorous bream and carp on sediment resuspension and the concentrations of nutrients and chlorophyll a were studied in sixteen experimental ponds (mean depth 1m, mean area 0.1 ha, sandy clay/clay sediment), stocked with bream or carp at densities varying from 0 to 500 kg ha^?1. Planktivorous perch (Perca fluviatilis L.) were added to some ponds to suppress zooplankton. 2. Suspended sediment concentrations increased linearly with biomass of benthivorous fish. Bream caused an increase of 46 g sediment m^?2 day^?1 per 100kg bream ha^?1 and a reduction of 0.38m^?1 in reciprocal Secchi disc depth, corresponding to an increase in the extinction coefficient of 0.34m^?1. 3. No relationship was found between size of fish and amount of resuspension, but the effect of bream was twice as great as that of carp. Benthivorous feeding was reduced in May because alternative food (zooplankton) was available. 4. Assuming a linear relationship, chlorophyll a level increased by 9.0 μgI^?1, total P by 0.03mgl^?1 and Kjeldahl-N by 0.48mgl^?1 per 100kg bream ha^?1. Silicate, chlorophyll a, total P and total N were all positively correlated with fish biomass, but orthophosphate showed no correlation.