Energy transformations in two tropical Oxbow lakes

The flow and pattern of energy utilization in two tropical ecosystems where polyculture of fishes was practiced were studied. The rates of organic production made by the two tropical natural aquatic systems (oxbow lakes) were studied for two annual cycles. The study of utilization of energy in two tropical lakes revealed that herbivores were more efficient in energy utilization than autotrophs. The energy available in these systems was supplied by allochthonous source. Silver carp and grass carp were found to be most efficient in conversion of energy in the grazing food chain of the lake. The efficiency of the system to fish production was found to be 0.06 (OL-I) ?0.05% (OL-II) of the total available solar radiation. Various forms of life are all accompanied by energy changes, in the studied lakes were found to form a food web.     

Phytoplankton-zooplankton relationships in the Western Pacific Ocean and adjacent seas

Regressions of biomass and daily food requirements of herbivorous zooplankton on daily primary production were calculated, using assumptions based on data collected in various sea areas of the western Pacific Ocean and adjacent seas. A regression coefficient (1.470) of calculated herbivorous biomass on observed daily primary production is significantly higher than unity (P<0.01). This indicates that the herbivorous biomass sustained by unit amount of primary production is large in the more productive high latitudes, and small in the less productive tropical sea areas. This is attributed to relatively larger food requirements per unit biomass of the tropical herbivores as compared with those found in cold waters. Despite distinct areal differences in the herbivorous biomass-primary production ratios, the calculated daily food requirement of herbivores was in direct proportion to the daily primary production, when equilibrium had been established between phytoplankton and zooplankton. Under conditions of limited food supplies, the small body size of the tropical herbivores may be advantageous both in reducing the total energy consumption per individual, and in inducing rapid growth and reproduction. Therefore, the low ratio of biomass to primary production in the tropics could beregarded as a result of possible regulation of tropical herbivores to scarce food conditions rather than as evidence of failure of adaptation to such conditions.     

Kelp as a trophic resource for marine suspension feeders: a review of isotope-based evidence

Kelp forests are enormously productive, and they and adjacent habitats support large populations of suspension feeders. What do these suspension feeders eat? Intuitively, we might expect that kelp primary production is a key form of trophic support for these animals. Indeed, a large and growing number of studies using carbon stable isotope data, typically collected over short time periods, have asserted that detritus from kelps is an important, and in some cases the main, food source for coastal benthic suspension feeders. This view has been incorporated into several textbooks and review papers covering kelp forest ecosystems, and loss of trophic support for benthic suspension feeders is now often invoked as an ecosystem consequence of top-down or other impacts on kelp forests. More direct evidence, however, suggests that these animals mainly eat phytoplankton and, in some cases, bacteria or zooplankton. Because isotope values of pure coastal phytoplankton, uncontaminated with detritus, are difficult to obtain, present studies have largely relied on single measurements from offshore environments or from the literature, which typically reflects offshore values. We review the evidence showing that phytoplankton isotope values can, and are expected to, vary widely in coastal waters and that inshore phytoplankton may often be enriched in ¹³C compared to offshore phytoplankton. This unaccounted-for variation may have systematically biased the results of such trophic studies toward finding large contributions of kelp detritus to suspension-feeder diets. We review some key stable isotope studies and put forth evidence for alternative explanations of the isotope patterns presented. Finally, we make recommendations for future isotope studies and describe several approaches for progress in this area. New techniques, particularly flow cytometry and compound-specific stable isotope analysis, provide ways to shed light on this interesting and important ecological issue.     

Aquatic and terrestrial organic matter in the diet of stream consumers: implications for mercury bioaccumulation

The relative contribution of aquatic vs. terrestrial organic matter to the diet of consumers in fluvial environments and its effects on bioaccumulation of contaminants such as mercury (Hg) remain poorly understood. We used stable isotopes of carbon and nitrogen in a gradient approach (consumer isotope ratio vs. periphyton isotope ratio) across temperate streams that range in their pH to assess consumer reliance on aquatic (periphyton) vs. terrestrial (riparian vegetation) organic matter, and whether Hg concentrations in fish and their prey were related to these energy sources. Taxa varied in their use of the two sources, with grazing mayflies (Heptageniidae), predatory stoneflies (Perlidae), one species of water strider (Metrobates hesperius), and the fish blacknose dace (Rhinichthys atratulus) showing strong connections to aquatic sources, while Aquarius remigis water striders and brook trout (Salvelinus fontinalis) showed a weak link to in-stream production. The aquatic food source for consumers, periphyton, had higher Hg concentrations in low-pH waters, and pH was a much better predictor of Hg in predatory invertebrates that relied mainly on this food source vs. those that used terrestrial C. These findings suggest that stream biota relying mainly on dietary inputs from the riparian zone will be partially insulated from the effects of water chemistry on Hg availability. This has implications for the development of a whole-system understanding of nutrient and material cycling in streams, the choice of taxa in contaminant monitoring studies, and in understanding the fate of Hg in stream food webs.     

Ecological implications of fecal pellets produced by the Antarctic krill Euphausia superba in the Antarctic Ocean

Organic analyses and electron microscopic observations on fecal pellets produced by the Antarctic krill Euphausia superba Dana showed that krill fed on choanoflagellates, the abundant heterotrophic flagellate in the Antarctic Ocean. Two new pathways of organic materials in the Antarctic ecosystem are proposed: (1) a new food chain including non-living particulate and dissolved organics, and bacteria-choanoflagellate-krill-vertebrate, which coexists with the traditional diatom-krill-vertebrate food chain; (2) non-phytoplanktonic organic materials in surface waters are transferred into choanoflagellates and are transported to deep water as fecal pellets which are still useful as nutrition for other organisms there.     

Responses to extreme seasonality in food supply: diet plasticity in Antarctic brachiopods

The ice gouged, shallow, polar seabed is a challenging place to live, but suspension feeders are particularly rich and abundant there. The extreme seasonality of food supply from phytoplankton at high latitudes, combined with very stable temperatures has not, however, reduced the range of life history adaptations. Some species extend feeding and growing periods by concentrating on smaller, lower biomass but longer duration, fractions of phytoplankton. Here we show that shallow-water Antarctic brachiopods can switch from pelagic to benthic food sources when required. Like most suspension feeders they utilise the extremely abundant summer phytoplankton blooms, but unlike many other groups brachiopods do not appear to become dormant in winter. In shallow sites around the world wind and wave action resuspend benthic material, making it available to filter feeders. Widespread ice disturbance at polar latitudes may also cause resuspension enhancing local food supply. Articulate brachiopods have blind-ended guts and probably feed little once a large bolus of food has been collected, so a resuspension event may represent a significant enhancement of yearly food availability. The key to their ability to utilise unpredictable food sources is the same trait responsible for their polar success; very low maintenance costs and a catholic diet. Utilising resuspended material could be an important factor in global brachiopod distributions. Previously difficult to explain high brachiopod densities in New Zealand, Northwest Europe and the Canadian Pacific Coast may occur because these areas are all in the worlds windiest, and hence strongest resuspension, latitudes.Communicated by J.P. Thorpe, Port Erin     

Is benthic trophic structure affected by food quality? The Dogger Bank example

The aim of this study was to evaluate the relationship between food quality and trophic structure of macrobenthic communities of the Dogger Bank. Therefore, 21 stations along different depth gradients (a transect across the central Dogger Bank and at a station cross in the north-eastern part) were sampled and analysed for macrofauna trophic structure and food supply, indicated by phytopigment composition and organic matter contents of the sediment. The study revealed differences in the trophic structure of the benthic community in areas with contrasting food supply. The trophic structure reflected differences in relative quality of organic matter. The sandy sediments on top of the bank receiving intermediate quantities of high food quality were inhabited by high abundances of interface feeders and sand-licking amphipods of the genus Bathyporeia. The southern stations characterised by a comparatively high amount of intermediate food quality were dominated by interface feeders such as Amphiura filiformis. The northern and north-eastern stations showing the highest diversity of feeding types were characterised by a comparatively low, but highly variable input of intermediate-quality organic matter consisting of both fresh and degraded organic matter. Results are discussed with a special emphasis on specific Dogger Bank features such as benthic primary production and frontal system characteristics.Communicated by O. Kinne, Oldendorf/Luhe     

Biogeographic patterns on shallow subtidal reefs in the western Indian Ocean

To resolve biogeographic limits and patterns on the east coast of Africa, the presence/absence and quantitative biomass data were collected from 55 shallow subtidal reefs along 4,800 km of coastline (5.2°–31.1°S). Multivariate analysis of distributions, trophic structure and biomass revealed two distinct marine provinces, the Tropical Indo-West Pacific and Subtropical Natal, with a transitional overlap (of ca. 120 km) located between Leven Point and Kosi Mouth on north-east South Africa. This region of overlap was one of three bioregions revealed by post hoc analyses. Biomass was unexpectedly highest in the tropics. ‘Auto-heterotrophs’ (species with autotrophic symbionts such as corals and clams), deposit feeders and grazers contributed significantly more biomass to the Tropical Indo-West Pacific Bioregion, whereas filter feeders dominated the Subtropical Natal Bioregion. ‘Auto-heterotrophs’ declined with latitude while filter feeders increased; soft corals made a defining contribution in the overlap bioregion. Possible underlying causes of these patterns include productivity, nutrient levels, riverine input, light penetration and temperature.     

Feeding ecology of benthopelagic zooplankton on an eastern tropical Pacific seamount

The gut contents of dominant deep-sea benthic boundary layer zooplankton (primarily copepods and mysids) and the vacuole contents of phaeodarians collected and preserved in situ at four depths (from 724 to 3112 m) on an eastern tropical Pacific seamount (Volcano 7; 13°23N, 102°27W) between 23 November and 4 December 1988 were analyzed using transmission electron microscopy (TEM). Suspended, and sinking plus resuspended particles, were quantitatively sampled to characterize potential food sources. The broad oxygen minimum characteristic of this region intersects the summit of the seamount and affects the feeding ecology of these organisms. Several copepods and mysids and an amphipod contained guts packed with what appeared to be gram-positive bacteria, an unusual finding. We hypothesize that the source of these bacteria-like bodies was a mat or aggregate that originated in the oxygen minimum or at its upper or lower interface. The presence of the bacteria-like bodies in 43 to 100% of the particlefeeding zooplankton that were sectioned and that had gut contents, suggests that the bacteria-like bodies are an important food source. The diverse gut and vacuole contents of other detritivores were similar among depths. Particles and microorganisms from the depths were also similar. This finding can be explained by the rapid sinking of particles and aggregates from surface waters and their relatively intact transit through the broad oxygen minimum with its reduced populations of zooplankton. The presence of algal cells in guts and vacuoles of benthic boundary-layer zooplankton suggests that these zooplankton select particles of recent surface origin for consumption. The presence of the guts filled with bacteria-like bodies shows that some deep-sea copepods and mysids that are normally generalist feeders can specialize opportunistically. The similarity of gut contents of crustaceans and vacuole contents of phaeodarians suggests that these two very different groups of particle feeders utilize similar food sources in the deep sea.     

Food web analysis in intertidal <Emphasis Type="Italic">Zostera marina</Emphasis> and <Emphasis Type="Italic">Zostera noltii</Emphasis> communities in winter and summer

The food web of two intertidal seagrass (Zostera marina and Zostera noltii) beds that may be influenced by the seasonal variation in food source abundance was studied in winter and in summer with δ¹³C and δ¹⁵N analysis. In spite of high relative variation of abundance of main primary producers at the two sites, the food web did not vary between winter and summer. The δ¹³C range of primary producers was wide. Zostera leaves, the most ¹³C-enriched source, were not consumed directly by grazers. Deposit and filter feeders have a similar δ¹³C and could use a mix of suspended and sedimented organic particulate matter, largely composed of detritus from macroalgae to seagrass. This trophic pathway allows the local incorporation of the high biomass produced by seagrasses. The wide δ¹⁵N range of predators was linked either to a large variety from omnivore to carnivore predators or to the also wide ranges of δ¹⁵N of primary consumers.     

On detritus as a food source for pelagic filter-feeders

Data on the amount of organic detritus within the particle size fraction 1 to 150 are presented for the Western Kiel Bight. Grouped into a mixed surface layer and a stratified lower layer in accordance with the main hydrographic features of the Western Baltic Sea, the rounded-off values show a seasonal variation between 100 and 600 mg m^-3 expressed as dry weight of organic matter. The overall average for both layers is about 200 mg m^-3. Organic detritus thus comprises more than 40% of total organic matter in the above size class, which is the class most easily accessible to the relatively small filter-feeders in this area. Attempting to trace the origin of organic detritus, a positive correlation to phytoplankton standing stock was found in some cases, suggesting the predominance of autochthonous detritus. Proceeding from the assumption that pelagic filter-feeders select their food mainly by size and not by taste, it is concluded that organic detritus plays an important role as a supplementary food source, being ingested together with phytoplankton and small nonmotile heterotrophs. The nutritive value of detritus is increased by the adsorption of dissolved organic matter and above all through the subsequent colonization by bacteria, which utilize the dissolved substances. Detritus particles serving as a substratum for bacteria thus form a means whereby dissolved organic substances reenter the food chain. The ingestion of detritus by filter-feeders is, therefore, thought to be instrumental in increasing the effectivity of energy transfer from the primary to the secondary food-chain level.     

Match/mismatch between the Mytilus edulis larval supply and seston quality: effect on recruitment

We considered Cushing's match/mismatch theory in a heterotrophic environment and hypothesized that settlement and recruitment success in blue mussel are higher when the food supply is rich in polyunsaturated and essential fatty acids (PUFA/EFA). To test this hypothesis, we monitored larval development as well as fatty acid composition in trophic resources during two successive reproductive seasons. The decoupling we found between the presence of competent larvae in the water column and settlement rates strongly suggests that metamorphosis is delayed until conditions are suitable. In both years, the major mussel settlement peak was synchronized with a phytoplanktonic pulse rich in EFA, consisting of a large autotrophic bloom in 2007 and a short but substantial peak of picoeukaryotes in 2008. These results suggest a "trophic settlement trigger" that indirectly affects recruitment by strongly improving the settlement rate. Despite similar larval settlement rates during both years, the lower 2007 recruitment likely resulted from a mismatch with a high lipid-quality trophic resource. The seasonal trophic conditions differed greatly between the two years, with fatty acids profiles reflecting heterotrophic plankton production in 2007 and mostly autotrophic production in 2008. In agreement with Cushing's theory, our results highlight a match/mismatch, related to the food lipid quality rather than food quantity. For the first time, we show that the recruitment in marine bivalves may be dependent on phytoplanktonic pulses characterized by high levels of PUFA.     

A directly coupled,artificial two-step food chain for long-term experiments with filter-feeders at constant food concentrations

A system is described consisting of a chemostat for culturing algae, a dilution unit and special experimental vessels, allowing long-term experiments with filter feeders at a constant concentration and quality of food. The algal suspension flows continuously from a chemostat, is mixed with sterile filtered water, and pumped directly into the experimental vessels. The flow through the vessels is intensive enough to avoid a significant change in food concentration by sedimentation or grazing. There are two possible arrangements, the first providing many duplicates with exactly the same food concentrations, the second providing a series of graduated food concentrations. Details of construction are given. The application of both arrangements is demonstrated by experiments with Daphnia pulex. Further possibilities of application are suggested.     

Sea ice microbial production supports Ross Sea benthic communities: influence of a small but stable subsidy

Diversity in guilds of primary producers enhances temporal stability in provision of organic matter to consumers. In the Antarctic ecosystem, where temporal variability in phytoplankton production is high, sea ice contains a diatom and microbial community (SIMCO) that represents a pool of organic matter that is seasonally more consistent, although of relatively small magnitude. The fate of organic material produced by SIMCO in Antarctica is largely unknown but may represent an important link between sea ice dynamics and secondary production in nearshore food webs. We used whole tissue and compound-specific stable isotope analysis of consumers to test whether the sea ice microbial community is an important source of organic matter supporting nearshore communities in the Ross Sea. We found distinct gradients in delta13C and delta15N of SIMCO corresponding to differences in inorganic carbon and nitrogen acquisition among sites with different sea ice extent and persistence. Mass balance analysis of a suite of consumers demonstrated large fluxes of SIMCO into the nearshore food web, ranging from 5% to 100% of organic matter supplied to benthic species, and 0-10% of organic matter to upper water column or pelagic inhabitants. A delta13C analysis of nine fatty acids including two key biomarkers for diatoms, eicosapentaenoic acid (EPA, 20:5omega3), and docosahexaenoic acid (DHA, 22:6omega3), confirmed these patterns. We observed clear patterns in delta13C of fatty acids that are enriched in 13C for species that acquire a large fraction of their nutrition from SIMCO. These data demonstrate the key role of SIMCO in ecosystem functioning in Antarctica and strong linkages between sea ice extent and nearshore secondary productivity. While SIMCO provides a stabilizing subsidy of organic matter, changes to sea ice coverage associated with climate change would directly affect secondary production and stability of benthic food webs in Antarctica.     

Breeding territory quality and agonistic behavior: effects of energy availability and intruder pressure in hummingbirds

Summary The relative importance of food a vailability and intruder pressure on breeding territorial behavior was studied in two experiments with male calliope hummingbirds. In the first experiment, extra food was provided in inconspicuous feeders. Territory owners who fed from the feeders increased their display rates (power-diving and hovering) when on their territories, and spent less time out of sight (mostly off their territories). Food addition by this method had no significant effect on intruder pressure, as measured by number and duration of chases. In the second experiment, intruder pressure was increased by attracting feedertrained males to feeders in territories of individuals who did not feed from feeders. In this experiment, display activity of residents did not change but chasing activity increased. These results suggest that display rates by male calliope hummingbirds depend at least in part on the amount of food available, but are not strongly affected by the number of feeding attempts by male intruders. No attempt was made to determine whether displays can deter competitors, attract mates, or do both. However, since these energetically expensive displays depend at least in part on the energy available to owners, they provide readily available indices of male and/or territory quality which could potentially be used for different purposes by different individuals.     

Evidence supporting the importance of terrestrial carbon in a large-river food web

Algal carbon has been increasingly recognized as the primary carbon source supporting large-river food webs; however, many of the studies that support this contention have focused on lotic main channels during low-flow periods. The flow variability and habitat-heterogeneity characteristic of these systems has the potential to significantly influence food web structure and must be integrated into models of large-river webs. We used stable-isotope analysis and IsoSource software to model terrestrial and algal sources of organic carbon supporting consumer taxa in the main channel and oxbow lakes of the Brazos River, Texas, USA, during a period of frequent hydrologic connectivity between these habitat types. Standardized sampling was conducted monthly to collect production sources and consumer species used in isotopic analysis. Predictability of hydrologic connections between habitat types was based on the previous 30 years of flow data. IsoSource mixing models identified terrestrial C3 macrophytes (riparian origin) as the primary carbon source supporting virtually all consumers in the main channel and most consumers in oxbow lakes. Small-bodied consumers (<100 mm) in oxbow lakes assimilated large fractions of algal carbon whereas this pattern was not apparent in the main channel. Estimates of detritivore trophic positions based on delta15N values indicated that terrestrial material was likely assimilated via invertebrates rather than directly from detritus. High flows in the river channel influenced algal standing stock, and differences in the importance of terrestrial and algal production sources among consumers in channel vs. oxbow habitats were associated with patterns of flooding. The importance of terrestrial material contradicts the findings of recent studies of large-river food webs that have emphasized the importance of algal carbon and indicates that there can be significant spatial, temporal, and taxonomic variation in carbon sources supporting consumers in large rivers.     

Why do honey bees dance?

The honey bee dance language, used to recruit nestmates to food sources, is regarded by many as one of the most intriguing communication systems in animals. What were the ecological circumstances that favoured its evolution? We examined this question by creating experimental phenotypes in which the location information of the dances was obscured. Surprisingly, in two temperate habitats, these colonies performed only insignificantly worse than colonies which were able to communicate normally. However, foraging efficiency was substantially impaired in an Asian tropical forest following this manipulation. This indicates that dance language communication about food source locations may be important in some habitats, but not in others. Combining published data and our own, we assessed the clustering of bee forage sites in a variety of habitats by evaluating the bees’ dances. We found that the indicated sites are more clustered in tropical than in temperate habitats. This supports the hypothesis that in the context of foraging, the dance language is an adaptation to the particular habitats in which the honey bees evolved. We discuss our findings in relation to spatial aggregation patterns of floral food in temperate and tropical habitats.     

Variation in the trophic structure and species composition of some invertebrate communities in polluted kelp forests in the North Sea

The absence of species from kelp holdfast habitats is used to establish changes in invertebrate community structure in polluted coastal waters along the shores of North Eastern Britain. The study shows the following changes: (1) Approximately 45% of the species complement studied in clean-water habitats are missing from polluted habitats. (2) Rare species are lost and food chains are missing. (3) Dislocations of the marine food web occur: removal of parts of the primary producer to herbivore food chain; suppression of some carnivores; removal of some omnivores and selection of certain suspension feeders. (4) No new species appear in polluted communities. (5) Recombinations of resistant species compensate for the reduction in energy flow through a damaged food web by proportionally increasing the suspension-feeding pathways. (6) The ecological equilibrium of the suspension-feeding groups which are responsible for maintaining the damaged food web are in a delicate state of balance. The further loss of 1 large food chain could result in the collapse of the remaining fragments of a polluted-water invertebrate community food web.     

Sedimentation of organic detritus in Lochs Etive and Creran,Argyll, Scotland

The rates of sedimentation of organic detritus were measured at 3 stations in two Scottish sea lochs, Loch Etive and Loch Creran, using sedimentation jars exposed at various depths. Details are given of the seasonal and depth distribution of sedimenting material, and its composition for a 1-year period. Differences in the pattern of sedimenting material collected at different depths and in the seasonal patterns of sedimentation at the different stations suggested that, in each case there were differences in the relative importance of detritus from various sources. At all stations, phytoplankton production made a relatively small contribution to the total detritus collected, either directly as dead cells, or indirectly as the faeces of zooplankton organisms. Near the head of Loch Etive there were contributions by filamentous algae and Enteromorpha sp., but a major source of detritus was terrestrial debris, mainly carried into the loch in the waters of the River Etive. In the lower basin of Loch Etive, terrestrial detritus also contributed to the total sedimenting near the surface, but at greater depths much of the material collected in the sedimentation jars probably resulted from short-term resuspension and re-deposition of bottom material, reflecting a net transport of fine sediment from the shallower to the deeper areas of the loch. Secondarily resuspended material was also a major source of material collected in the jars exposed in Loch Creran.     

Animal-sediment relations and community analysis of a Florida estuary

Five species assemblages of the intertidal infauna of Old Tampa Bay, Florida, USA are identified. Two assemblages are judged to constitute distinct communities, while a third is shown to be an interdigitation of the two communities. Dominance by on species is the prevalent pattern within the assemblages. Numbers of deposit feeders are found to be inversely correlated to that of filter feeders, and both trophic types are found to be correlated to the sediment parameters of median grain size, sorting and skewness. Three transects with three stations each were established along the south side of Courtney Campbell Causeway in Tampa Bay, Florida. A faunal sample (0.4 m²), a sediment sample, and a water sample were taken at each station in September, December, and March of 1968/1969. Sediment samples were wet-seived. Animal samples were reduced to numbers of organisms and biomass per species. Trellis diagrams and correlation tests were generated. Support is shown for the trophic group-amensalims hypothesis, however, the silt-clay fraction is apparently of lesser importance to deposit feeders in Florida sediments than in Buzzards Bay sediments. An attempt is made to relate an analysis of the optimal grain size for filter feeders to the trophic group-amensalism hypothesis. A view of communities as abstractions from continua is more realistic than communities as discrete units.