Variation in the trophic basis of production and energy flow associated with emergence of larval salamander assemblages from forest ponds

1. Larval amphibians are a dominant consumer in many freshwater systems, yet limited data on energy transfers between aquatic food resources and larvae and between metamorphosed larvae and adjacent habitats preclude an accurate assessment of their roles as links between aquatic and terrestrial food webs.
2. During 2003–04, we derived prey-specific assimilation efficiencies, analysed stomach contents, and intensively sampled ambystomatid salamander assemblages in four ponds to quantify the trophic basis of larval production. Using estimates of the contribution of each prey taxon to larval production, we constructed quantitative food webs and assessed variation in pathways of energy flow associated with emergences.
3. Overall, metamorphosed salamanders exported 3–8% of total prey production, required to account for total salamander production, to adjacent forest. Aquatic insects, zooplankton and amphibian prey were most important to energy flow associated with emergence; amounts of larval production attributed to each of these prey types shifted during development and varied among salamander taxa.
4. The majority of variation in the trophic basis of production among species was attributed to copepods (Cyclopidae) and three families of aquatic insects (Chironomidae, Chaoboridae and Culicidae). Dominant prey types contributing to the production of metamorphosed salamanders varied among ponds, representing different pathways for energy transfers between aquatic resources and forest habitats. These findings further our understanding of the ecological roles of amphibians and thus the consequences of amphibian declines and extinctions.     

Nitrogen subsidies to pelagic food webs through profundal methane-oxidising bacteria in oligotrophic fresh water

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Assimilation of Diazotrophic Nitrogen into Pelagic Food Webs

The fate of diazotrophic nitrogen (N_D) fixed by planktonic cyanobacteria in pelagic food webs remains unresolved, particularly for toxic cyanophytes that are selectively avoided by most herbivorous zooplankton. Current theory suggests that N_D fixed during cyanobacterial blooms can enter planktonic food webs contemporaneously with peak bloom biomass via direct grazing of zooplankton on cyanobacteria or via the uptake of bioavailable N_D (exuded from viable cyanobacterial cells) by palatable phytoplankton or microbial consortia. Alternatively, N_D can enter planktonic food webs post-bloom following the remineralization of bloom detritus. Although the relative contribution of these processes to planktonic nutrient cycles is unknown, we hypothesized that assimilation of bioavailable N_D (e.g., nitrate, ammonium) by palatable phytoplankton and subsequent grazing by zooplankton (either during or after the cyanobacterial bloom) would be the primary pathway by which N_D was incorporated into the planktonic food web. Instead, in situ stable isotope measurements and grazing experiments clearly documented that the assimilation of N_D by zooplankton outpaced assimilation by palatable phytoplankton during a bloom of toxic Nodularia spumigena Mertens. We identified two distinct temporal phases in the trophic transfer of N_D from N. spumigena to the plankton community. The first phase was a highly dynamic transfer of N_D to zooplankton with rates that covaried with bloom biomass while bypassing other phytoplankton taxa; a trophic transfer that we infer was routed through bloom-associated bacteria. The second phase was a slowly accelerating assimilation of the dissolved-N_D pool by phytoplankton that was decoupled from contemporaneous variability in N. spumigena concentrations. These findings provide empirical evidence that N_D can be assimilated and transferred rapidly throughout natural plankton communities and yield insights into the specific processes underlying the propagation of N_D through pelagic food webs.     

Isotopic analysis of the sources of organic carbon for zooplankton in shallow subarctic and arctic waters

Shallow high-latitude lakes and ponds are usually characterized by an oligotrophic water column overlying a biomass-rich, highly productive benthos. Their pelagic food webs often contain abundant zooplankton but the importance of benthic organic carbon versus seston as their food sources has been little explored. Our objectives were to measure the δ¹³C and δ¹⁵N isotopic signatures of pelagic and benthic particulate organic matter (POM) in shallow water bodies in northern Canada and to determine the relative transfer of this material to zooplankton and other aquatic invertebrates. Fluorescence analysis of colored dissolved organic matter (CDOM) indicated a relatively strong terrestrial carbon influence in five subarctic waterbodies whereas the CDOM in five arctic water columns contained mostly organic carbon of autochthonous origin. The isotopic signatures of planktonic POM and cohesive benthic microbial mats were distinctly different at all study sites, while non-cohesive microbial mats often overlapped in their δ¹³C signals with the planktonic POM. Zooplankton isotopic signatures indicated a potential trophic link with different fractions of planktonic POM and the non-cohesive mats whereas the cohesive mats did not appear to be used as a major carbon source. The zooplankton signals differed among species, indicating selective use of resources and niche partitioning. Most zooplankton had δ¹³C values that were intermediate between the values of putative food sources and that likely reflected selective feeding on components of the pelagic or benthic POM. The results emphasize the likely importance of benthic-pelagic coupling in tundra ecosystems, including for species that are traditionally considered pelagic and previously thought to be dependent only on phytoplankton as their food source.     

Habitat coupling in lake ecosystems

Lakes are complex ecosystems composed of distinct habitats coupled by biological, physical and chemical processes. While the ecological and evolutionary characteristics of aquatic organisms reflect habitat coupling in lakes, aquatic ecology has largely studied pelagic, benthic and riparian habitats in isolation from each other. Here, we summarize several ecological and evolutionary patterns that highlight the importance of habitat coupling and discuss their implications for understanding ecosystem processes in lakes. We pay special attention to fishes because they play particularly important roles as habitat couplers as a result of their high mobility and flexible foraging tactics that lead to inter-habitat omnivory. Habitat coupling has important consequences for nutrient cycling, predator-prey interactions, and food web structure and stability. For example, nutrient excretion by benthivorous consumers can account for a substantial fraction of inputs to pelagic nutrient cycles. Benthic resources also subsidize carnivore populations that have important predatory effects on plankton communities. These benthic subsidies stabilize population dynamics of pelagic carnivores and intensify the strength of their interactions with planktonic food webs. Furthermore, anthropogenic disturbances such as eutrophication, habitat modification, and exotic species introductions may severely alter habitat connections and, therefore, the fundamental flows of nutrients and energy in lake ecosystems.     

Chemical information transfer in freshwater plankton

The structure of aquatic ecosystems is determined by complex interactions among individual organisms at different trophic levels. Although our basic understanding of how top-down and bottom-up processes interact to determine food-web dynamics has advanced, we still lack insights into how complex interactions and feedbacks affect the dynamics and structure of food webs. It is now becoming increasingly clear that, in addition to energy transfer from one trophic level to the other, there is exchange of information between these levels facilitated by the release of infochemicals by the organisms. There is evidence from recent studies that the exchange of chemical information in freshwater ecosystems is likely to play a decisive role in shaping structure and functioning of these systems. Chemical communication among freshwater organisms mediates many aspects of both predation and interspecific competition, which play key roles in determining community structure and ecosystem functioning. For example, consumer-induced defences in phytoplankton and zooplankton include modifications in the characteristics relating to life history, behaviour, morphology and biochemistry. These inducible defences affect trophic interactions by altering predator feeding rates through changes in attack rate or handling time, or both. Also host-specific fungal parasitism in phytoplankton is probably controlled by infochemicals. The motile fungi recognise their host by host-secreted compounds. Until now models describing the functioning of ecosystems mainly considered flows of biomass and energy. Integration of new knowledge about the role of chemical communication in these models may be one of the aims of ecological informatics. In this chapter I discuss how infochemicals may affect the dynamics and structure of planktonic food webs.     

Benthos as the basis for arctic lake food webs

Plankton have traditionally been viewed as the basis for limnetic food webs, with zooplankton acting as a gateway for energy passing between phytoplanktonic primary producers and fish. Often, benthic production has been considered to be important primarily in shallow systems or as a subsidy to planktonic food web pathways. Stable isotope food web analyses of two arctic lakes (NE14 and I minus) in the Toolik Lake region of Alaska indicate that benthos are the primary source of carbon for adults of all species of benthic and pelagic fish present. We found no effect of turbidity, which may suppress benthic algae by shading, on food web structure. Even though Secchi transparency varied from 10.2 m in NE14 to 0.55–2.6 m in I minus, food webs in both lakes were based upon benthos, had four trophic levels, and culminated with omnivorous lake trout. We suggest that the importance of benthos in the food webs of these lakes is due to their extreme oligotrophy, resulting in planktonic resources that are insufficient for the support of planktivorous consumers.     

Associating particulate essential fatty acids of the ω3 family with phytoplankton species composition in a Siberian reservoir

1. We studied variation in the composition of fatty acids in the seston of a small freshwater reservoir with changes in phytoplankton composition during four growth seasons. We focused on the dynamics of the ω3 fatty acids because of their potential importance for zooplankton nutrition. 2. Total diatoms were related to the 20:5ω3 fatty acid (eicosapentaenoic, EPA) content in seston. Among two dominant diatom genera, Cyclotella was not associated with EPA content. In contrast, there was a significant correlation between Stephanodiscus and the percentage contribution and content of EPA throughout the study. Hence, freshwater diatoms can differ strongly in content of the essential EPA. 3. We considered abundant cyanobacteria as a potential source of 18:3ω3 fatty acid (linolenic, ALA) to aquatic food webs. Among four dominant cyanobacteria species, two (Anabaena flos‐aquae and Planktothrix agardhii) showed significant correlation with the ALA content of the seston, while the other two (Aphanizomenon flos‐aquae and Microcystis aeruginosa) did not. 4. Dinophyta had a relatively high level of 22:6ω3 (docosahexaenoic, DHA) for freshwater species and can be also a source of EPA to aquatic food webs. 5. Our results show that various species of diatoms as well as cyanobacteria can be of contrasting nutritional value for zooplankton because of their different content of the essential PUFAs. Diatoms, which are low in EPA, could not be considered as a valuable food, while some field populations of cyanobacteria might be valuable sources of essential ALA.     

Chemical cues, defence metabolites and the shaping of pelagic interspecific interactions

Several observations and model calculations suggest that chemically mediated interactions can structure planktonic food webs. However, only recently have improvements in chemical methods, coupled with ecological assays, led to the characterization of chemical cues that affect the behaviour and/or physiology of planktonic organisms. We are currently beginning to elucidate if or how chemical signals can directly affect the interactions between species and even shape complex community structures in aquatic systems. Here, we highlight recent research on the nature and action of chemical signals in the pelagic marine and freshwater environments, with an emphasis on kairomones and defence metabolites.     

Habitat connectivity and ecosystem productivity: implications from a simple model

The import of resources (food, nutrients) sustains biological production and food webs in resource-limited habitats. Resource export from donor habitats subsidizes production in recipient habitats, but the ecosystem-scale consequences of resource translocation are generally unknown. Here, I use a nutrient-phytoplankton-zooplankton model to show how dispersive connectivity between a shallow autotrophic habitat and a deep heterotrophic pelagic habitat can amplify overall system production in metazoan food webs. This result derives from the finite capacity of suspension feeders to capture and assimilate food particles: excess primary production in closed autotrophic habitats cannot be assimilated by consumers; however, if excess phytoplankton production is exported to food-limited heterotrophic habitats, it can be assimilated by zooplankton to support additional secondary production. Transport of regenerated nutrients from heterotrophic to autotrophic habitats sustains higher system primary production. These simulation results imply that the ecosystem-scale efficiency of nutrient transformation into metazoan biomass can be constrained by the rate of resource exchange across habitats and that it is optimized when the transport rate matches the growth rate of primary producers. Slower transport (i.e., reduced connectivity) leads to nutrient limitation of primary production in autotrophic habitats and food limitation of secondary production in heterotrophic habitats. Habitat fragmentation can therefore impose energetic constraints on the carrying capacity of aquatic ecosystems. The outcomes of ecosystem restoration through habitat creation will be determined by both functions provided by newly created aquatic habitats and the rates of hydraulic connectivity between them.     

The ecology of chytrids in aquatic ecosystems: roles in food web dynamics

Chytrids are very important components of freshwater ecosystems, but the ecological significance of this group of fungi is not well understood. This review considers some of the significant environmental factors affecting growth and population composition of chytrids in aquatic habitats. The physical factors include primarily salinity, dissolved oxygen concentration and temperature. The biological factors include the role of chytrids as saprobes and parasites and methods of dispersal of propagules throughout the ecosystem. Dispersal depends upon both zoospores for short range and whole thalli for long range dispersal. Five roles for chytrids in food-web dynamics are proposed: (1) chytrid zoospores are a good food source for zooplankton, (2) chytrids decompose particulate organic matter, (3) chytrids are parasites of aquatic plants, (4) chytrids are parasites of aquatic animals and (5) chytrids convert inorganic compounds into organic compounds. New molecular methods for analysis of chytrid diversity in aquatic environments have the potential to provide accurate quantitative data necessary for better understanding of ecological processes in aquatic ecosystems.     

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Recreational boats as a vector of secondary spread for aquatic invasive species and native crustacean zooplankton

Recreational boats in tow between lakes are a known vector of the spread of aquatic invading species (AIS), but we have no test of the hypothesis that recreational boats are also a vector of secondary spread of AIS among freshwater ecosystems via in-water transport i.e., while boating between interconnected waterways. In this study, we surveyed recreational boaters travelling into Lake Simcoe (44°25′N, 79°20′W), Ontario, Canada, on their recreational activities, boat maintenance, and travel destinations, measured the degree of vessel fouling, and sampled all standing water and attached macrophytes associated with their vessels. A total of 321 zooplankton individuals comprising 15 different species were collected from the standing water in vessels, including veligers of the invasive zebra mussel Dreissena. The volume of water collected within the vessels significantly increased the number of zooplankton transported. Zooplankton species from pelagic habitats or with planktonic life stages were collected more frequently than species that occupy littoral or benthic habitats, likely reflecting the recreational activities of boaters. Patterns of boater activities, movements and hygiene habits, suggest recreational boating in the Lake Simcoe region is contributing to the spread of native and invasive species into nearby waterways. Our study validates the widespread assumption that recreational boats are an important in-water vector for the secondary spread of both native and invasive zooplankton species. Future management strategies to reduce the spread of AIS should be aimed at increasing awareness of boater hygiene practices, particularly the frequent draining of standing water.     

Ecological functions of volatile organic compounds in aquatic systems

In terrestrial ecosystems, volatile organic compounds (VOCs) are widely acknowledged as an important group of infochemicals. They play a major role in pollinator attraction by terrestrial plants and as insect pheromones. Furthermore, they are the mediating agent of so-called 'tritrophic interactions'. When plants are attacked by herbivorous insects, volatile signal substances are emitted, which act as attractants for parasitoids that kill the herbivores, thereby protecting the plant from herbivory. Despite the generally acknowledged importance of VOCs in terrestrial chemical ecology, their functions in aquatic food webs are largely unknown. VOCs produced by algae and cyanobacteria are a major concern in water processing, since aquatic primary producers are the reason for regularly encountered taste and odour problems in drinking water. Only very recently, research in aquatic chemical ecology has started to investigate possible ecological functions for the production of VOCs by algae and cyanobacteria. Volatile aldehydes released by wounded cells of marine planktonic diatoms seem to act as defensive compounds against herbivorous copepods on the population level. Just recently, it was found that VOCs released from benthic algae and cyanobacteria can be utilised as food and/or habitat finding cues by aquatic invertebrates such as freshwater gastropods and nematodes. Here, I review concepts and recent experimental studies on the ecological functions of such VOCs in aquatic ecosystems. Understanding the factors that lead to the liberation of volatile compounds is an essential prerequisite to properly assessing their ecological functions. It appears that (similar to terrestrial plant-herbivore interactions) VOCs can also play a steering role for both attraction and defence in aquatic ecosystems.     

Survival of Campylobacter jejuni in water: effect of grazing by the freshwater crustacean Daphnia carinata (Cladocera)

Environmental studies of the human-pathogenic bacterium Campylobacter jejuni have focused on linking distributions with potential sources. However, in aquatic ecosystems, the abundance of C. jejuni may also be regulated by predation. We examine the potential for grazing by the freshwater planktonic crustacean Daphnia carinata to reduce the survival of C. jejuni. We use a system for measuring grazing and clearance rates of D. carinata on bacteria and demonstrate that D. carinata can graze C. jejuni cells at a rate of 7% individual(-1) h(-1) under simulated natural conditions in the presence of an algal food source. We show that passage of C. jejuni through the Daphnia gut and incorporation into fecal material effectively reduces survival of C. jejuni. This is the first evidence to suggest that grazing by planktonic organisms can reduce the abundance of C. jejuni in natural waters. Biomanipulation of planktonic food webs to enhance Daphnia densities offers potential for reducing microbial pathogen densities in drinking water reservoirs and recreational water bodies, thereby reducing the risk of contracting water-borne disease.     

Algal Polyunsaturated Fatty Acids: Food,Pheromones and Foul Odour

Algal polyunsaturated fatty acids (PUFAs) are known to be high‐energy, often‐essential resources to freshwater aquatic food webs. On the other hand, high PUFA cell content in some algal taxa has been linked with the production of derivatives that may act as pheromones, allelogens or toxins. It has been known for some time that these compounds function in marine chemical ecology, but recent evidence indicates that they may play similar roles in freshwater ecosystems. This paper presents field and laboratory studies of planktonic and periphytic communities from freshwater systems, and shows that these communities are rich sources of certain PUFAs and derivatives which may function in both positive and negative foodweb interactions. This also has important implications for surface water quality, as these compounds are potent sources of rancid fishy odours.     

The importance of omnivory and predator regulation of prey in freshwater fish assemblages of North America

Synopsis Food webs for 15 freshwater ecosystems in North America are reconstructed, based upon fish feeding habits in these ecosystems and established trophic categorizations for aquatic invertebrates. My own research in Goose Creek, Virginia, as well as literature data, provided the necessary information. One property of theoretical interest for food webs, namely omnivory, is examined, and the results are related to various topics more typically studied by freshwater fish ecologists, including predator-prey regulation, optimal foraging theory, and tropho-dynamics. The data indicate that omnivory is often important in freshwater ecosystems, such that fish may not always control the abundance of their prey. It is emphasized that knowledge of omnivory patterns can provide important insights into community structure and function in freshwater ecosystems.     

Zooplankton,fish and fisheries in tropical freshwaters

About 40% of all fish species occur in freshwater, although only 1% of the globe is occupied by freshwaters. The tropics harbour a high percentage of these fishes. Freshwater zooplankton on the other hand is far less diverse than its marine counterpart and the tropics do not harbour a markedly high percentage of freshwater species either. The antecedents of freshwater zooplankton appear to have come from riverine and terrestrial habitats via temporary habitats (ponds, floodplains). The present zooplankton composition has also been shaped by, among other factors, the highly efficient zooplanktivorous modern teleosts which have restricted the formerly widespread Branchiopoda mainly to fishless freshwaters. Those Branchiopoda frequently co-existing with fishes (Cladocera) have their size composition strongly influenced by fish predation. Circumstantial evidence indicates that pelagic zooplankton (Cladocera, Copepoda, Rotifera) appear to provide a relatively scarce food supply relative to the littoral region for the early stages of fishes. Also, unpredictability of zooplankton may be involved here. These factors have led to the loss of planktonic eggs and the siting of fish breeding in shallow littoral situations, where other animal foods besides zooplankton are also available, especially for later stages of juvenile fish. The Ostariophysi breed in the shallow expanses of flood waters while the Cichlidae, some of which breed like Ostariophysi, also breed in standing waters in the littoral of lakes or floodplains. In all these locations zooplankton and benthic organisms, especially insects, are available. The cichlids are, in addition, provided with parental care. Predation on young fishes is also reduced by these strategies. Young fishes may also be adapted to feed on patches of zooplankton and other food in their breeding grounds. Tropical pelagic clupeids and cyprinids may breed continuously. Some of these clupeids in rivers breed at low water.Zooplankton, supplemented by other animal food is more critical to achieving the potential fish yields in the tropics than in temperate freshwaters because fish yields in the tropics can be very high indeed. The high metabolic rate of young fishes in the tropics adds to the demand for food. Tropical freshwaters have a relatively high primary production but a low zooplankton/phytoplankton ratio. Zooplankton is kept small in size and biomass by continuous predation. Herbivorous fishes can sustain very high fish yields in the tropics but they must have a high fecundity and high survival of young produced seasonally, mainly in rivers or even continuously as in lakes and reservoirs to weather predation. Rich littoral zooplankton and benthos combined with omnivory and a higher efficiency in the use of the available animal food by newly hatched fishes may be critical factors linking fish yields to zooplankton in tropical freshwaters. The ability of herbivorous tilapias to give very high fish yields in shallow tropical lakes may also be due to their efficient use of animal food, algae and microphagy in young stages besides other favourable adaptations like opportunistic feeding on detrities and the ability to feed on and digest cyanobacteria, abundant in the tropics.