Temporal and spatial change of the investment in carnivory of the tropical Utricularia foliosa

Investment by bladderwort (Utricularia foliosa L.) in carnivory, in terms of biochemical composition (carbohydrates per bladder), elemental composition (carbon and nitrogen per bladder), and morphology of the bladders (length, depth, size of the trap door, and size of antennae), was estimated in seven plants located in Yahuarcaca creek (Colombian Amazon) five times from March to May 2005. The aims were to determine whether investment in carnivory varies temporally (over the growing season of the plant) and/or spatially, and if this potential change in carnivory investment varies according to nutrient conditions. The main differences in the investment in carnivory (changes in bladder number and bladder size, and changes in the size of the antennae) were among locations and there were not important differences over the growing season of the plant. Nitrogen and not phosphorus, was the element that stimulated the investment in carnivory. In addition to changes in bladder number and bladder size, we observed a new strategy to enhance prey capture under nitrogen limitation: changes in the size of the antennae. The size of the antennae was approximately 1.3 higher in those plants located in sites with low NO₃⁻. However, we did not observed changes in the carbon/nitrogen ratio of the bladders or in the relationship between bladder length with bladder depth or size of the trap door. The amount of carbohydrates per bladder was also 1.8 higher in those plants located in sites with low NO₃⁻ (0.13 μM) than those with higher NO₃⁻ concentration (0.39 μM). However, the amount of carbohydrates in the bladder was related with the abundance of periphyton and, hence, it is not possible to conclude that carbohydrate production was a strategy of the plant to enhance the capture of prey. Therefore, our findings do not support the carbohydrate mucilage lure speculations.     

Predation by the tropical plant Utricularia foliosa

1. We examined the prey captured by individual plants of the tropical carnivorous plant Utricularia foliosa, located in different areas along a creek in the Colombian Amazon and the zooplankton and macroinvertebrate communities associated with the plants. The aims were: (i) to determine whether bladders of different sizes within each plant catch different numbers of prey or exploit different size ranges and types of prey, (ii) if the quantity and composition of prey captured varies temporally and/or spatially and (iii) if the plant has evolved effective mechanisms of attracting prey. 2. Utricularia foliosa captured the most abundant species of macroinvertebrates associated with the plant. Larger bladders captured more, larger and more diverse prey. However, benefits of the extra prey caught by large bladders were not offset by the greater cost of producing bladders larger than approximately 1650 μm. 3. The number of prey captured was higher in those plants with more carbohydrates per bladder and with a higher ratio of antenna size/bladder length. The antennae enhance capture success by offering the prey a favourable substratum that exploits their natural locomotor and feeding behaviour. However, although carbohydrates may lure prey, carbohydrate production was not a strategy of the plant to enhance the capture of prey, because the amount of carbohydrates in the bladder was related to the abundance of periphyton.     

A wide window of migration phenology captures inter‐annual variability of favourable conditions: Results of a whole‐lake experiment with juvenile Pacific salmon

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Periphyton and phytoplankton associated with the tropical carnivorous plant Utricularia foliosa

The abundance and taxonomic composition of periphyton attached to the bladders and phytoplankton associated with the aquatic carnivorous plant Utricularia foliosa were quantified, to determine whether periphyton associated with U. foliosa would enhances predation success. Bladder size, periphyton abundance and periphyton richness together explained 76% of the variation observed in the number of prey captured by the bladders. The abundance and richness of periphyton followed the same pattern as phytoplankton, i.e., both increased as dissolved inorganic phosphate concentration rose. This nutrient concentration explained 84 and 74% of the variation observed in richness and abundance of periphyton, respectively. This suggests that abundance and richness of the periphyton associated with U. foliosa depend mostly on environmental conditions rather than on facilitation mechanisms displayed by the plant. In conclusion, if periphyton affects U. folisosa negatively due to the competition for light or nutrients, the plant is “fated to get along with the enemy” but apparently without the capacity to manipulate this “enemy” to its own advantage.     

Impact of a Protective Limestone Treatment on the Water Chemistry and Zooplankton Community of Thrush Lake, Minnesota

A limestone slurry was sprayed on the surface of Thrush Lake, a small headwater lake in northeastern Minnesota, to test a treatment designed to protect acid-sensitive waters from anthropogenic acidification. The 6-year study, consisting of pretreatment, transition, and post-treatment phases, was part of the four-state Acid Precipitation Mitigation Program directed by the U.S. Fish and Wildlife Service. Measured water-chemistry parameters, including acid-neutralizing capacity, pH, dissolved calcium, and dissolved inoroganic carbon, increased following treatment, although local climatic conditions influenced the magnitude and duration of the chemical changes. Physical changes to the lake, other than an increase in conductivity and a short-term alteration of water clarity subsequent to treatment, were not documented. The composition of the zooplankton community was altered, with the proportion of rotifers increasing after treatment. Individual zooplankton species showed a variety of changes in abundance that were associated with treatment over both seasonal and multi-year intervals. For example, Holopedium gibbemm was absent from lake samples immediately following treatment and recovered within a season, whereas Diaptomus minutus and Keratella taurocephala populations were reduced after treatment and had not recovered by the end of the study. Alternately, Asplanchna priodonta increased in abundance after treatment. These observed abundance patterns were generally consistent with previzous acidification or base-addition studies. In contrast, the changes in community composition of zooplankton did not consistently fit patterns developed from regional studies across water-chemistry gradients. These differences emphasize the importance of biotic as well as abiotic factors in controlling zooplankton community composition.     

Contrasting Zooplankton Assemblages in Two Oxbow Lakes with Low Transparencies and Narrow Emergent Macrophyte Belts (Krapina River,Croatia)

The aim of this study was to examine the combined effect of water transparency and narrow macrophyte belts on zooplankton assemblages in two oxbow lakes (Krapina River, Croatia). Samples were collected in open water and among helophytes in the littoral zone from April until September 2008. Rotifers were the most abundant group of zooplankton in both lakes, and dominated in the Krapina oxbow lake 1 (KO1). Lake KO1 had significantly lower transparency, lower percentage macrophyte cover and higher chlorophyll a concentration than Krapina oxbow lake 2 (KO2). In lake KO1, variation in the horizontal distribution of cladocerans and rotifers in terms of their abundance seemed to be determined by competition between Bosmina longirostris and Keratella cochlearis, initiated by oscillation in transparency and detritus availability. In lake KO2, with higher transparency and higher percentage macrophyte cover, the abundance of small‐ and large‐bodied cladocerans increased in the littoral zone simultaneously with higher transparency, suggesting fish predation. Results of this study indicated that small differences in transparencies between the two lakes caused significant differences in horizontal distribution of the zooplankton assemblage. Even narrow helophyte belts offered a refuge to zooplankton, although lower transparencies reduced the effectiveness of macrophytes as a refuge from predators. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Zooplankton densities in a Hydrilla infested lake

The number of individuals and species of zooplankton were sampled concurrently with Hydrilla biomass and water quality for one year in a small, eutrophic central Florida lake. Throughout the study, rotifer species and individuals dominated the zooplankton. The abundance of the zooplankton tended to remain high when Hydrilla biomass was at its seasonal low during late winter and early spring. When hydrilla growth increased in the late spring and summer months causing a decrease in total alkalinity, specific conductivity, water color, turbidity, orthophosphate and chlorophyll a concentrations; the abundance of the zooplankton declined. During this time, there was a shift from limnetic to littoral species, principally rotifers. Hydrilla growth did not affect the mean number of cladoceran or copepod species, but may have led to an increase in rotifer species.     

Long-Term Changes in the Diet of Gymnogobius isaza from Lake Biwa,Japan: Effects of Body Size and Environmental Prey Availability

Body size and environmental prey availability are both key factors determining feeding habits of gape-limited fish predators. However, our understanding of their interactive or relative effects is still limited. In this study, we performed quantitative dietary analysis of different body sizes of goby (Gymnogobius isaza) specimens collected from Lake Biwa between 1962 and 2004. First, we report that the diet was composed mainly of zooplankton (cladocerans and copepods) before the 1980s, and thereafter, shifted to zoobenthos (gammarids). This foraging shift coincided with, and thus can be linked to, known historical events in the lake at that time: decrease in zooplankton abundance with the alleviation of eutrophication, increase in fish body size resulting from fish population collapse, and increase in gammarid abundance due to reduced fish predation pressure. Supporting this view, our data analyses revealed how the long-term changes in the diet composition would be co-mediated by changes in fish body size and environmental prey availability. Specifically, while zoobenthos abundance strongly affected the fish diet composition, larger (smaller) fish preferred zoobenthos (zooplankton). Furthermore, the body size effects were stronger than those of prey availability. These results provide the best long-term evidence that fish feeding habits vary over decades with its body size and prey community due to anthropogenic disturbances.     

Beneficial and detrimental interactive effects of dissolved organic matter and ultraviolet radiation on Zooplankton in a transparent lake

While changes in dissolved organic matter (DOM) concentrations are expected to affect zooplankton species through attenuation of potentially damaging ultraviolet (UV) radiation, generation of potentially beneficial or harmful photoproducts, pH alteration, and microbial food web stimulation, the combined effects of such changes on zooplankton community structure have not been studied previously. Our purpose was to determine how an increase in allochthonous DOM and associated changes in pH in an initially transparent lake may affect zooplankton community structure, and how exposure to solar UV may alter these DOM and pH effects. We ran microcosm experiments manipulating UV, DOM, and pH near the surface of Lake Giles in northeastern Pennsylvania. We found that when DOM was added in the presence of ambient UV, Daphnia and copepod UV-mortality was reduced by approximately three and two times compared to UV exposure without extra DOM. When DOM was added in the absence of UV, adult Daphnia and copepods were reduced compared to no DOM addition in the absence of UV. Daphnia and cyclopoid egg production and rotifer abundance were generally higher in the presence of DOM, regardless of UV treatment. The lower abundance yet high egg production in the presence of DOM and absence of UV may be explained by higher abundance of egg-bearing adults compared to non-egg-bearers. We conclude that allochthonous DOM benefits some zooplankton in a high-UV environment, but may be detrimental under low-UV conditions. Overall, Daphnia abundance and egg production were higher than that of calanoid copepods in the DOM additions, indicating that in some lakes an increase in allochthonous DOM may lead to a zooplankton community shift favoring Daphnia over calanoid copepods.     

Pelagic C:N:P Stoichiometry in a Eutrophied Lake: Responses to a Whole-Lake Food-Web Manipulation

Changes in the ecological stoichiometry of C, N, and P in the pelagic zone are reported from a whole-lake manipulation of the food web of Lake 227, an experimentally eutrophied lake at the Experimental Lakes Area, Canada. Addition of northern pike eliminated populations of planktivorous minnows by the third year (1995) after pike introduction, and in the fourth year after pike addition (1996), a massive increase in the abundance of the large-bodied cladoceran Daphnia pulicaria occurred. Accompanying this increase in Daphnia abundance, zooplankton community N:P declined, seston concentration and C:P ratio decreased, and dissolved N and P pools increased. During peak abundance, zooplankton biomass comprised a significant proportion of total epilimnetic phosphorus (greater than 30%). During the period of increased Daphnia abundance, concentrations of dissolved inorganic nitrogen (TIN) increased more strongly than dissolved phosphorus (TDP), and thus TIN:TDP ratios were elevated. Sedimentation data indicated that increased grazing led to greatly reduced residence times of C, N, and especially P in the water column during 1996. Finally, previously dominant N-fixing cyanobacteria were absent during 1996. Our results show that strong effects of food-web structure can occur in eutrophic lakes and that stoichiometric mechanisms play a potentially important role in generating these effects.     

Environmental factors influencing zooplankton species composition of lakes in north-central Ontario,Canada

To assess the relative importance of lake chemistry, morphometry and zoogeography on limnetic zooplankton, we collected zooplankton, water, and morphometric data from 132 headwater Canadian Shield lakes in 6 regions across north-central Ontario. A subset of these lakes (n = 52) were fished with gill nets. We clustered lakes based on their zooplankton species composition (presence/absence). Discriminant analysis was employed to determine how well lake characteristics could predict zooplankton community types. Correct classification of zooplankton communities for three models ranged from 72 to 91%. Lake size, lake location, and buffering capacity were ranked as the most important factors separating lake groups. Fish abundance (CPUE) was not significant in distinguishing between zooplankton communities. Though the range of lake sizes was limited (1–110 ha), larger lakes tended to support more species. Lake location (zoogeography) also influenced species composition patterns. Although Algoma lakes tended to be larger (\-x = 18.0 ha, other lakes \-x = 2.5 ha), they supported relatively depauperate zooplankton communities. Buffering capacity was ranked third in the discriminant analysis models, but pH and alkalinity were not significantly different between lake groups.     

An Analysis of the Biotic Community in a Kumaun Himalayan Lake,Nainital (U. P.), India

A comprehensive study of phyto- and zooplankton and macrozoobenthic components in Lake Nainital showed that species richness was high for plankton and low for macrozoobenthos. The algal biomass was dominated by greens (54 %) and blue-greens (31 %), the zooplankton population by copepods (84 %), and the macrozoobenthic community by a Tubifex-Chironomus association constituting≥95 % of the annual number of the macrobenthic invertebrates. Respiration (807.5g C m⁻² year⁻¹) surpassed gross production (630.5 g C m⁻² year⁻¹). The mean annual ratio between phyto- and zooplankton biomass is 3.3 and between phytoplankton and herbivores it is 4.6. If biomass is treated as a measure of crude production, the relationship among the three trophic levels suggests that herbivory is inefficient while carnivory is efficient, because part of the primary production remains unutilized by dominant herbivorous zooplankters, whereas Mesocyclops leuckarti, the sole carnivore, feeds efficiently on rotifers and juveniles of other copepods. The low diversity of different biotic components and the P/R ratio of less than 1 perhaps suggest that the lake is passing through the stage of heterotrophic succession.     

Determination of zooplankton dietary shift following a zebra mussel invasion, as indicated by stable isotope analysis

1. Freshwaters with established zebra mussel populations typically exhibit reduced chlorophyll a concentrations, but the subsequent impacts upon zooplankton are varied. We hypothesised that in an invaded system with less phytoplankton but available allochthonous subsidy, zooplankton may utilise greater proportions of allochthonous matter and that this could be traced by analysis of stable carbon and nitrogen isotopes. 2. We used archived zooplankton samples which had been consistently preserved and which spanned the invasion period of an Irish lake, Lough Erne. Increasing reliance upon allochthonous resources would be reflected in an increase in zooplankton δ¹³C away from phytoplankton which is relatively ¹³C‐depleted in humic‐stained L. Erne. 3. Analysis of a series of monthly samples (1992–96, 1999–2003) revealed significant ¹³C‐enrichment of mixed zooplankton, Eudiaptomus gracilis and Mysis relicta post‐zebra mussel invasion; δ¹³C values approached ?27‰ typical of terrestrial organic matter during spring and autumn. Changes in zooplankton elemental composition also suggested a switch to a lower quality diet. However, analysis of zooplankton δ¹³C from an annual, single‐point (June) time series spanning 28 years (1977–2004) suggested that when phytoplankton was sufficiently abundant, zooplankton used this resource and their δ¹³C remained relatively constant around ?32‰. Post‐invasion enrichment of mysid δ¹⁵N may reflect a shift towards carnivory, but planktonic prey abundance was reduced and a subsequent loss of body condition could result in the same isotopic changes. 4. Our results indicate that in L. Erne, when phytoplankton was reduced by zebra mussel filtering, zooplankton assimilated more from allochthonous matter, and potentially sustained a higher population than would otherwise be possible. Thus, zebra mussel impact on foodweb structure and function is likely to be different in lakes subject to varying subsidy levels.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Nutrient availability and the carnivorous habit in Utricularia vulgaris

1. Carnivory in plants is thought to enhance growth through an increased supply of nutrients, although there are considerable costs involved. It has been assumed that the relative investment of biomass in traps is inversely proportional to the availability of nutrients from non-carnivorous sources. Our aim was to test the effect of increasing nutrient concentration on investment in carnivory by Utricularia vulgaris .
2. Plants were grown under controlled conditions and nitrogen and phosphorus added at three loadings in a crossed design. Investment in carnivory was assessed as the proportion of (i) leaf biomass and (ii) leaf area comprising traps.
3. There was no effect of nutrient additions on plant growth or periphyton abundance. Investment in carnivory declined with increasing phosphorus loading. There was no effect of nitrogen, despite this being the nutrient commonly thought to be sought by carnivorous plants. Analysis of previously published data also indicated a decline in investment with increasing P availability.
4. Investment in carnivory in U. vulgaris is inversely proportional to the availability of phosphorus from non-carnivorous sources.     

Recurrent response patterns of a zooplankton community to whole-lake fish manipulation

1. In a series of whole-lake manipulations conducted from 1984 to 1991, planktivorous fishes were alternately removed and restocked in a small mesotrophic lake, resulting in dramatic changes in the zooplankton community. 2. Response patterns in the zooplankton community, which include species and size structure, and within-year community variability, were examined. Variation in the zooplankton community in unmanipulated years was much lower than that in manipulated years, regardless of direction of the manipulation (i.e. decreasing or increasing planktivory). 3. The succession of zooplankton species abundance was repeated in the second removal of planktivorous fishes. The community shifted from small-bodied cladocerans, copepods and rotifers, through an intermediate state with high abundance of Holopedium, to an assemblage dominated by large-bodied daphnids.     

Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.     

The food, growth and abundance of five co-existing cyprinids in lake basins of different morphometry and water quality

The food, growth and abundance of five co-existing cyprinid fish species in the eutrophic Lake Hiidenvesi were studied. The diet overlaps within the cyprinid community in shallow lake basins were compared with those in a deep basin, where littoral resources are less available. Roach, bleak and white bream inhabited both the shallow and the deep parts of the lake. Their growth rate was slow, probably due to the low availability of animal food, indicated by the increasing proportion of detritus and plant material in the diets towards the end of the summer. In the deep basin, roach and bleak, contrary to white bream, did not forage on the very abundant invertebrate Chaoborus flavicans, explained by the migration behaviour of C. flavicans. Blue bream, unlike other cyprinids, utilized copepods and had a relatively fast growth rate, but was mostly restricted to the shallow areas. The condition of the bream stock was weak both in terms of growth and abundance. The availability of zoobenthos was low and bream was not able to compete for zooplankton with the more specialized planktivores.     

Effects of the non-indigenous cladoceran Cercopagis pengoi on the lower food web of Lake Ontario

1. In North America, the invasive predatory cladoceran Cercopagis pengoi was first detected in Lake Ontario. We explored the impact of Cercopagis on the lower food web of Lake Ontario through assessments of historical and seasonal abundance of the crustacean zooplankton, by conducting feeding experiments on the dominant prey of the invader, and by estimating its food requirements. 2. Between 1999 and 2001, a decrease in the abundance of dominant members of the Lake Ontario zooplankton community (Daphnia retrocurva, Bosmina longirostris and Diacyclops thomasi) coincided with an increase in the abundance of Cercopagis. Daphnia retrocurva populations declined despite high fecundity in all 3 years, indicating that food limitation was not responsible. Chlorophyll a concentration generally increased, concomitant with a decline in the herbivorous cladoceran zooplankton in the lake. 3. Laboratory experiments demonstrated that Cercopagis fed on small‐bodied species including D. retrocurva and B. longirostris. 4. Consumption demand of mid‐summer populations of Cercopagis, estimated from a bioenergetic model of the confamilial Bythotrephes, was sufficient to reduce crustacean abundance, although the degree of expected suppression varied seasonally and interannually. 5. Predatory effects exerted by Cercopagis on the Lake Ontario zooplankton, while initially very pronounced, have decreased steadily as the species became established in the lake.     

Some ecological aspects of Lake Qarun,Fayoum, Egypt. Part II. Production of plankton and benthic organisms

Distribution and abundance of phyto-, zooplankton and benthic organisms in Lake Qarun were investigated during the period from January 1974 to December 1977.Average number of phytoplankton cells was 152,300 cells/L and its biomass was 0.365 g/C/m³; average number of zooplankton was 31.44 × 10³/m³ and its biomass was 194.19 mg/m³. The average number of benthic fauna was 19889/m² and its biomass was 400.22 g/m² (dry wt.). Therefore, Lake Qarun may be considered as a highly eutrophic body of water.Freshwater planktonic species, that used to inhabit the lake, such as Diaptomus salinus and the cladoceran Moina salinarum, disappeared completely when the salinity of the lake water reached 30–34 However, some Rotatoria were able to withstand the high salinity. The new composition of the zooplankton community shows that the marine zooplankton species include not only Acartia latisetosa and Cirripedia nauplii, but also other species such as Polychaeta, Obelia medusae, etc.The benthos of Lake Qarun is characterised by an intensive growth of few species. The major part (i.e. 93.54% by weight) of bottom fauna in the lake is Mollusca, mainly Cerastoderma glaucum (69·84% by weight).