Diet segregation between two sympatric 'small'Barbus spp: an experimental study of mechanisms

Gut contents of two co‐occurring species of 'small' diploid barbs (<10  L _F cm) in Lake Tana revealed that zooplankton is the major diet component for B. tanapelagius (75% based on volume), but less prominent in B. humilis (40%). Functional response experiments in the laboratory were conducted to elucidate the mechanisms causing this difference. The type of functional response by the two 'small' barbs under different microcrustacean zooplankton densities (10, 20, 40, 60 and 80 ind.l⁻¹) was examined. The functional response of B. tanapelagius to increasing prey densities corroborates with Holling Type II model, whereas B. humilis exhibits a Type III functional response. Predation rate is higher for B. tanapelagius at low zooplankton density (<40 ind.l⁻¹) and equals the level of B. humilis at higher densities (>40 ind.l⁻¹). This suggests that at lower zooplankton densities B. humilis is a less efficient forager on zooplankton prey items than B. tanapelagius . In Lake Tana average zooplankton density is relatively low (<35 ind.l⁻¹). Under these food conditions, B. humilis is forced to feed on other food items (e.g. benthic invertebrates), whereas B. tanapelagius primarily feeds on zooplankton. The feeding potentials of the two 'small' barbs, as deduced from their morphology explain their different performances and their segregation in space and food resources.     

Population dynamics of Bythotrephes cederstroemii in south-east Lake Michigan 1995–1998

1. Population characteristics (density, size, reproductive patterns) of the predatory cladoceran Bythotrephes cederstroemii in south-east Lake Michigan were monitored at an offshore station (110 m) in 1995–98 and at a nearshore station (45 m) in 1997–98.
2. The mean density of B. cederstroemii at the offshore station was generally highest in July–September (145–914 m⁻²) and at the nearshore station in October–November (168–1625 m⁻²). In 1995 and 1998, density was also high at the offshore station in November (211–284 m⁻²). Fish predation may limit B. cederstroemii in nearshore regions in the summer. The maximum annual densities of B. cederstroemii for 1995–98 were generally similar to those reported from the late 1980s, when the species arrived in Lake Michigan.
3. Body size increased rapidly each year to a maximum in August. Thereafter, body size declined and converged for stage-2 and 3 individuals, suggesting food scarcity or size-selective fish predation was affecting large individuals.
4. Most reproduction occurred asexually (90%), and by stage 2 or 3 females (99%). Asexual brood size was highest when B. cederstroemii first appeared each year, and decreased in August, when larger neonates were produced. There appeared to be differences in reproductive mode for stage 2 and 3 females, with a higher percentage of stage 2 females reproducing sexually.     

The effect of an emergent macrophyte (Typha angustifolia) on sediment resuspension in a shallow north temperate lake

1. The effects of emergent macrophytes on water turbidity and sediment resuspension in the shallow Kirkkojärvi basin of Lake Hiidenvesi were studied with sediment traps, and concomitant sediment and water samples. The study was conducted during May–August in three different zones of a stand of the emergent Typha angustifolia .
2. Within the stand (5 m from the edge), both the concentration of suspended solids and the rate of sediment resuspension were significantly lower than at the edge and outside the stand (5 m from the edge). The differences between the zones increased towards the end of summer together with the growing stem density. During the study period (82 days), 2210 g dw m⁻² of sediment was resuspended in the outer zone. At the edge and in the inner zone, the corresponding numbers were 1414 and 858 g dw m⁻², respectively.
3. With the resuspended sediment, 39.4 mg P  m⁻² day⁻¹ was brought to the water column outside the stand, 22.4 mg P  m⁻² day⁻¹ at the edge and 13.4 mg P  m⁻² day⁻¹ within the stand.
4. In early summer, the concentration of suspended solids had a highly significant positive effect on soluble reactive phosphorus (SRP) concentration in the water, whereas in late summer no effect was found. During the study period, phosphorus retention by emergent macrophyte stands corresponded to 3–5% of the present annual external phosphorus loading of the Kirkkojärvi basin.     

Phytoplankton production and growth rate in Lake Tanganyika: evidence of a decline in primary productivity in recent decades

1. This study focused on phytoplankton production in Lake Tanganyika. We provide new estimates of daily and annual primary production, as well as growth rates of phytoplankton, and we compare them with values published in former studies.
2. Chlorophyll- a (chl- a ) in the mixed layer ranged from 5 to 120 mg chl- a  m⁻² and varied significantly between rainy and dry seasons. Particulate organic carbon concentrations were significantly higher in the south basin (with 196 and 166 mg C m⁻³ in the dry and the rainy season, respectively) than in the north basin (112 and 109 mg C m⁻³, respectively).
3. Carbon : phosphorus (C : P) ratios varied according to season. Phosphorus limitation seemed to occur more frequently than nitrogen limitation, especially during the rainy season. Severe P deficiencies were rare.
4. Measured particulate daily primary production ranged from 110 to 1410 mg C m⁻² day⁻¹; seasonal contrasts were well marked in the north basin, but less in the south basin, where primary production peaks occurred also in the rainy season. Estimates of annual primary production, based on daily primary production calculated from chl- a and water transparency, gave values lower than those reported in previous studies. Picophytoplankton accounted on average for 56% of total particulate production in the south basin during the wet season of 2003.
5. Phytoplankton growth rates, calculated from primary production, ranged from 0.055 to 0.282 day⁻¹; these are lower than previously published values for Lake Tanganyika.     

Reaction distance of Bythotrephes longimanus, encounter rate and index of prey risk for Harp Lake, Ontario

SUMMARY 1. Some predatory zooplankton, such as the large cladoceran Bythotrephes longimanus (Leydig), have a large compound eye, hence vision may play an important role in prey detection and encounter. Our objectives were to quantify, under different light regimes, the reaction distance of Bythotrephes to an assortment of zooplankton prey and to model encounter rate with prey from Harp Lake, Ontario. Reaction distance to prey increased at higher light intensity.
2. Results from the encounter model show that small, slow-moving prey faced the greatest risk from Bythotrephes and most encounters occurred in the upper 10 m of the water column throughout the 24-h period. The model was highly sensitive to ambient light. Encounter rate and prey risk were two to three orders of magnitude more sensitive to swimming velocity of the predator than to that of the prey.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Sublittoral abundance and food consumption of Baltic gobies

Two small demersal fishes, the sand goby Pomatoschistus minutus and the common goby Pomatoschistus microps , were quantified on soft bottoms at 20–40 m depth in the Baltic Sea, using a camera placed above the bottom. The largest numbers of gobies were seen following the settlement of young in late summer and autumn. Most recorded fishes were sand gobies. An annual average of 4·7 individuals m⁻²(0·24 g dry mass m⁻²) was recorded in 1983–1985 and 2·5 individuals m⁻²(0·13 g m⁻²) in 1997–1998. Using these densities, the annual goby food consumption was estimated to 100 kJ m⁻² in 1983–1985 and 50 kJ m⁻² in 1997–1998, corresponding to most of the annual macrobenthos production available to the gobies. The resulting goby production, assumed equal to 25% of the food consumed, must have been an important food source for the larger fishes occasionally recorded in the photographs.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

Effects of artificial ultraviolet-B radiation on experimental aquatic microcosms

1. The effects of prolonged ultraviolet-B (UVB) radiation on freshwater communities were studied in indoor microcosms (600 L) with artificial light sources, simulating a clear, shallow, mesotrophic aquatic ecosystem. A range of six intensities (in duplicate) of UVB radiation, ranging from 0 (control) to 9.56 kJ m⁻² day⁻¹ at the water surface, was applied for 8 weeks. The UVB radiation levels, attenuation, shading and scattering were comparable to those in Dutch shallow freshwater systems. Physical, chemical and biological variables were monitored weekly.
2. The UVB treatment did not affect the abundance, species composition or biovolume of the phytoplankton or zooplankton communities, nor did it affect the periphyton or the macroinvertebrate community. A few species showed a significant response on some of the sampling dates, but there was no negative UVB effect at the community level. Overall, the ecosystems in the microcosms were not affected by the UVB treatment.
3. In a bio-assay, a laboratory clone of Daphnia pulex , not subjected to UVB radiation, was fed with seston from the microcosms. Daphnia pulex feeding on seston from the control microcosms grew faster, had better survival and better reproduction than D. pulex feeding on seston from the UVB treated microcosms. The phytoplankton–zooplankton interaction may have been influenced by the UVB treatment.
4. The dissolved oxygen content (DOC) concentrations in the microcosms were around 5 mg L^−1. The DOC levels in Dutch systems rarely fall below 10 mg L^−1. This might provide sufficient protection against the detrimental effects of increased UVB radiation.     

Long-term effects of successive Ca(OH)2 and CaCO3 treatments on the water quality of two eutrophic hardwater lakes

1. Whole-lake experiments were conducted in two hardwater lakes (Halfmoon and Figure Eight) in Alberta, Canada, to investigate the effectiveness of repeated lime (slaked lime: Ca(OH)₂ and/or calcite: CaCO₃) treatments (5–78 mg L^–1) for up to 7 years.
2. Randomized intervention analysis of intersystem differences between the experimental and three reference lakes demonstrated a decline in euphotic total phosphorus and chlorophyll a concentrations in the experimental lakes after repeated lime treatments.
3. After the second lime application to Halfmoon Lake, mean winter total phosphorus release rates (TPRR) decreased to < 1 mg m^–2 day^–1 compared with 3.6 mg m^–2 day^–1 during the winter after initial treatment. In the final year of lime application, mean summer TPRR decreased to 4.5 mg m^–2 day^–1 compared with 7.6 mg m^–2 day^–1 in the pre-treatment year.
4. Mean macrophyte biomass declined and species composition was altered at 1 and 2 m depths in Figure Eight Lake during lime application. Over the first 6 years of treatment, macrophyte biomass at 2 m declined by 95% compared with concentrations recorded during the initial treatment year. In the last year of the study, macrophyte biomass at 2 m reached initial treatment concentrations, which coincided with the greatest water transparency. Over the treatment period, macrophyte species shifted from floating to rooted plants.
5. Multiple lime applications can improve water quality in eutrophic hardwater lakes for periods of up to 7 years.     

ESTIMATION OF PREY DENSITIES REQUIRED BY WESTERN NORTH ATLANTIC RIGHT WHALES

The apparent lack of significant recovery of western North Atlantic right whale populations may be related to the availability of prey in exploitable densities. We have attempted to estimate the concentration of zooplankton required by right whales to obtain a net energetic benefit over the long term, using estimated values for body weight, metabolic rate, assimilation efficiency, time spent feeding, mouth size and swimming speed. The estimated range of required energy densities is 7.57 to 2,394 kcal m⁻³ (3.17 × 10⁴ to 1.00 × 10⁷ joule m⁻³). These values are from one to more than three orders of magnitude greater than the densest concentration sampled in the vicinity of right whale aggregations in the Great South Channel. Right whales must seek out and exploit extremely dense patches of prey organisms in order to feed efficiently. The presence of such dense patches of zooplankton is likely to be a primary characteristic of spring, summer and fall right whale habitats.     

An introduced invertebrate predator (Bythotrephes) reduces zooplankton species richness

Rarely do ecologists have the data needed to assess the impacts of invading species on biodiversity, i.e. pre- and post-invasion census information from both invaded and control sites. Using a 21-year time series, we demonstrate that the invasion of Harp Lake, Ontario, Canada, by the Eurasian spiny water flea, Bythotrephes longimanus , a zooplanktivore, was accompanied by a rapid and long-lasting reduction in the average species richness of crustacean zooplankton, particularly of cladoceran taxa. No such reduction was observed in seven nearby un-invaded lakes over the same two decades. If the Harp Lake results are typical, we predict a widespread reduction in crustacean zooplankton richness on the Canadian Shield for three reasons. Shield lakes provide the invader with good habitat. Its dispersal rates and colonization success are high. Zooplankton richness in Harp Lake is now unusually low for a Shield Lake of its size and acidity.     

The effect of turbidity and illumination on the reaction distance and search time of the marine planktivore Gobiusculus flavescens

Both reduced illumination and increased turbidity caused a significant reduction in reaction distance of Gobiusculus flavescens . The longest reaction distance, 18.9 cm for larger prey (Calanus finmarchicus) , occurred at a light level of 80 μmol m ⁻² s ⁻¹ compared to 12.9 cm for a smaller prey (Acartia clausi) at 8 μmol m⁻² s⁻¹. Above a light saturation level of 10 μmol m⁻² s⁻¹, additional light had little influence on reaction distance. In the turbidity experiments, the longest reaction distances were measured at turbidity levels of 10–20 JTU. Prey size influenced reaction distance at all tested light levels. Search time was influenced by prey size only at low illumination. With increasing turbidity, reaction distance to a group of prey was longer than to one prey.     

Differing effects of suspended sediments on the performance of native and exotic Daphnia

1. Although large cladocerans are usually uncommon in large rivers, Daphnia lumholtzi (an exotic species in North America) is widespread and occasionally abundant. We collected zooplankton on the Illinois River (Illinois, U.S.A.) in 1995 and 1996 and found that the peak density of D. lumholtzi (25 L⁻¹) typically exceeded that of all native species combined. Maximum density occurred during warm periods (up to 27 °C) when concentrations of inorganic suspended sediments were high (>50 mg L⁻¹).
2. Using a life table experiment, we examined the effects of variation in suspended sediment (0 and 80 mg L⁻¹) and food (10⁴ and 10⁵ Ankistrodesmus cells mL⁻¹) on fitness of D. lumholtzi and the native Daphnia parvula. Daphnia lumholtzi had greater survivorship than D. parvula in most treatments and higher life-time fertility than D. parvula in all treatments. Both species achieved their fastest intrinsic rates of growth in treatments with high food, but their responses to suspended solids differed. The growth rate of D. lumholtzi in high food was slightly increased by higher turbidity, whereas that of D. parvula was depressed.
3. Results suggest that the ability of D. lumholtzi to tolerate suspended solids is an important factor contributing to its success in invading North American rivers.     

Functional response of juvenile pink and chum salmon: effects of consumer size and two types of zooplankton prey

Feeding rate experiments were conducted for pink salmon Oncorhynchus gorbuscha fry [mean fork length ( L _F) 39 mm], juveniles (103–104 mm L _F) and juvenile chum salmon Oncorhynchus keta (106–107 mm L _F). Fishes were presented with small copepod ( Tisbi sp.) or larger mysid shrimp ( Mysidopsis bahia ) prey at varying densities ranging from 1 to 235 prey l⁻¹ in feeding rate experiments conducted at water temperatures ranging from 10·5 to 12·0° C under high light levels and low turbidity conditions. Juvenile pink and chum salmon demonstrated a type II functional response to mysid and copepod prey. Mysid prey was readily selected by both species whereas the smaller bodied copepod prey was not. When offered copepods, pink salmon fry fed at a higher maximum consumption rate (2·5 copepods min⁻¹) than larger juvenile pink salmon (0·4 copepods min⁻¹), whereas larger juvenile chum salmon exhibited the highest feeding rate (3·8 copepods min⁻¹). When feeding on mysids, the maximum feeding rate for larger juvenile pink (12·3 mysids min⁻¹) and chum (11·5 mysids min⁻¹) salmon were similar in magnitude, and higher than feeding rates on copepods. Functional response models parameterized for specific sizes of juvenile salmon and zooplankton prey provide an important tool for linking feeding rates to ambient foraging conditions in marine environments, and can enable mechanistic predictions for how feeding and growth should respond to spatial-temporal variability in biological and physical conditions during early marine life stages.     

Plant palatability and disturbance level in aquatic habitats: an experimental approach using the snail Lymnaea stagnalis (L.)

1. The palatability of aquatic macrophytes to the snail Lymnaea stagnalis was investigated in the laboratory. Eight species of macrophyte were selected from habitats that differed in either flood disturbance regime or nutrient status.
2. In a non-choice test, single macrophyte species were offered to individual snails. The average amount of plant dry mass consumed per Lymnaea dry mass ranged from 3.6 ± 1.4 (±SE) to 63.6 ± 13.9 mg g^–1 day^–1 across plant species. In a choice test, all eight plant species were presented simultaneously to sets of five snails. The average total consumption was 66.1 ± 3.8 mg g^–1 day^–1 and the maximum average consumption for a single plant was 26.2 ± 3.6 mg g^–1 day^–1.
3. In both tests, the amount consumed by snails differed significantly between the plant species. The species growing in undisturbed habitats were the least consumed. Habitat nutrient status was unrelated to plant palatability.
4. These results suggest that macrophyte species growing in habitats that are rarely disturbed by floods allocate a greater proportion of their resources to resisting herbivory.     

Mercury in tigerfish (Hydrocynus vittatus, Castelnau), green happy (Sargochromis codringtonii, Boulenger) and kapenta (Limnothrissa miodon, Boulenger) from Lake Kariba, Zimbabwe

Concentrations of total mercury were determined in Hydrocynus vittatus (Castelnau), Sargochromis codringtonii (Boulenger), and Limnothrissa miodon (Boulenger) from two localities in Lake Kariba, Zimbabwe.
The mean concentrations of total mercury in H. vittatus from Basin 5 and Basin 2 were 0.08 mg kg⁻¹ and 0.094 mg kg⁻¹, respectively. In S. codringtonii, mean concentrations were 0.004 mg kg⁻¹ and 0.026 mg kg⁻¹ for Basins 5 and 2, respectively. No mercury was detected in L. miodon from Basin 2 while samples from Basin 5 had a mean concentration of 0.069 mg kg⁻¹ (wet weight). Total mercury concentrations were also determined on a dry weight basis.
Within each sampling area, total mercury concentrations were significantly different among species ( P  < 0.05). For H. vittatus and S. codringtonii, total mercury concentrations (in the same species) were not significantly different between the two localities ( P  < 0.05).
The factors causing the observed differences in total mercury between similar species from different localities and among different species in the same locality (sampling area) are discussed. From the observed low levels of mercury in all three species, it was concluded that the mercury constituted 'background levels'. These levels are below the maximum concentrations permissible in human fish foods.     

Swimming performance, oxygen consumption and excess post-exercise oxygen consumption in adult transgenic and ocean-ranched coho salmon

Routine oxygen consumption ( M o ₂) was 35% higher in 1 day starved and 21% higher in 4 day starved adult transgenic coho salmon Oncorhynchus kisutch relative to end of migration ocean-ranched coho salmon. Critical swimming speed ( U _crit) and M o ₂ at U _crit ( M o _2max) were significantly lower in 4 day starved transgenic coho salmon (1·25 BL s⁻¹; 8·79 mg O₂ kg⁻¹ min⁻¹) compared to ocean-ranched coho salmon (1·60 BL s⁻¹; 9·87 mg O₂ kg⁻¹ min⁻¹). Transgenic fish swam energetically less efficiently than ocean-ranched fish, as indicated by a poorer swimming economy at U _crit ( M o _2max     ). Although M o _2max was lower in transgenic coho salmon, the excess post-exercise oxygen consumption (EPOC) measured during the first 20 min of recovery was significantly larger in transgenic coho salmon (44·1 mg O₂ kg⁻¹) compared with ocean-ranched coho salmon (34·2 mg O₂ kg⁻¹), which had a faster rate of recovery.     

Spatial and temporal patterns of microcrustacean assemblage structure and secondary production in a wetland ecosystem

1. In contrast to extensive studies of zooplankton in lakes, the role of microcrustaceans in wetlands is not well studied. In this study, spatial and temporal patterns of microcrustacean assemblage structure and secondary production were quantified over a 2-year period in a southeastern U.S.A. wetland.
2. Thirty-two species, including 19 cladocerans, 10 copepods and three ostracods, generated different temporal patterns of density and production between vegetated ( Nymphaea ) and non-vegetated (open-water) zones reflecting species-specific differences in life histories.
3. Summer assemblages were dominated by small, planktonic filter-feeders, typified by high annual production/biomass ( P / B ) and daily production. In contrast, winter assemblages were dominated by larger, epibenthic detritivores with low P / B and high biomass. Seasonal shifts in the relative importance of planktonic species in the warmer months to benthic and epiphytic species in the cooler months suggest that energy flow pathways through microcrustaceans may vary seasonally.
4. Total annual production was higher during both years in the Nymphaea zone (13.0 g and 13.6 g DM m⁻² year⁻¹) than the open-water (8.2 and 6.3 g DM m⁻² year⁻¹), and was similar between years for the entire wetland pond (12.3 and 12.2 g DM m⁻² year⁻¹).
5. Although wetland ecosystems have been the subject of considerable ecological research in the past 20 years, our study is one of the few to demonstrate a highly diverse and relatively productive microcrustacean assemblage. Such comprehensive production studies can be used to quantify the ecological importance of microcrustaceans in freshwater wetland ecosystems.