Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

Are closure terms appropriate or necessary descriptors of zooplankton loss in nutrient–phytoplankton–zooplankton type models?

Our current knowledge of plankton ecology ascribes a large proportion of zooplankton losses to zooplankton cannibalism and carnivory, rather than via the activity of higher trophic levels beyond the plankton. However, planktonic ecosystem models, such as the widely used nutrient–phytoplankton–zooplankton (NPZ) type models, typically represent all zooplankton losses by mathematically (rather than biologically) justified closure functions. Even where it is assumed that these closure functions include zooplanktonic cannibalism and carnivory, these processes are not explicitly implemented within the grazing function of the zooplankton. Here it is argued that this representation of zooplankton losses through “closure” terms within planktonic food web models is neither appropriate nor necessary. The general consequences of implementing a simple function incorporating zooplankton cannibalism and carnivory (intra-guild predation) within a planktonic food web model are compared against models implementing different types of traditional closure functions. While the modelled biomass outputs may appear similar, the fate of annual primary production and f-ratios vary widely. There appears no justification for the continued use of traditional closure term to depict zooplankton loss processes on biological or modelling arguments. To do so can seriously misrepresent the fate of primary production and thence trophic dynamics.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Which forcing factors fit? Using ecosystem models to investigate the relative influence of fishing and changes in primary productivity on the dynamics of marine ecosystems

Fishing mortality and primary production (or proxy for) were used to drive the dynamics of fish assemblages in 9 trophodynamic models of contrasting marine ecosystems. Historical trends in abundance were reconstructed by fitting model predictions to observations from stock assessments and fisheries independent survey data. The model fitting exercise derives values for otherwise unknown parameters that specify the relative strength of trophic interactions and, in some instances, a time series anomaly for changes in primary production. We measured how much better or worse were model predictions when bottom-up forcing by primary production were added to top-down forcing by fishing. Searching for cross system patterns, the relative contribution of fishing and changes in primary production, mediated through trophic interactions, are evaluated for the ecosystems as a whole and for selected similar species in different ecosystems. The analysis provides a simple qualitative way to explain which forcing factors have most influence on modeled dynamics. Both fishing and primary production forcing were required to obtain the best model fits to data. Fishing effects more strongly influenced 6 of 9 of the ecosystems, but primary production was more often found to be the main factor influencing the selected pelagic and demersal fish stock trends. Examination of sensitivity to ecological and model parameters suggests that the results are the product of complex food-web interactions rather than simple deterministic responses of the models.     

Structural changes in lake functioning induced from nutrient loading and climate variability

Climate variability is increasingly recognized as an important regulatory factor, capable of influencing the structural properties of aquatic ecosystems. Lakes appear to be particularly sensitive to the ecological impacts of climate variability, and several long time series have shown a close coupling between climate, lake thermal properties and individual organism physiology, population abundance, community structure, and food web dynamics. Thus, understanding the complex interplay among meteorological forcing, hydrological variability, and ecosystem functioning is essential for improving the credibility of model-based water resources/fisheries management. Our objective herein is to examine the relative importance of the ecological mechanisms underlying plankton seasonal variability in Lake Washington, Washington State (USA), over a 35-year period (1964–1998). Our analysis is founded upon an intermediate complexity plankton model that is used to reproduce the limiting nutrient (phosphate)–phytoplankton–zooplankton–detritus (particulate phosphorus) dynamics in the lake. Model parameterization is based on a Bayesian calibration scheme that offers insights into the degree of information the data contain about model inputs and allows obtaining predictions along with uncertainty bounds for modeled output variables. The model accurately reproduces the key seasonal planktonic patterns in Lake Washington and provides realistic estimates of predictive uncertainty for water quality variables of environmental management interest. A principal component analysis of the annual estimates of the underlying ecological processes highlighted the significant role of the phosphorus recycling stemming from the zooplankton excretion on the planktonic food web variability. We also identified a moderately significant signature of the local climatic conditions (air temperature) on phytoplankton growth (r = 0.41), herbivorous grazing (r = 0.38), and detritus mineralization (r = 0.39). Our study seeks linkages with the conceptual food web model proposed by Hampton et al. [Hampton, S.E., Scheuerell, M.D., Schindler, D.E., 2006b. Coalescence in the Lake Washington story: interaction strengths in a planktonic food web. Limnol. Oceanogr. 51, 2042–2051.] to emphasize the “bottom-up” control of the Lake Washington plankton phenology. The posterior predictive distributions of the plankton model are also used to assess the exceedance frequency and confidence of compliance with total phosphorus (15 μg L⁻¹) and chlorophyll a (4 μg L⁻¹) threshold levels during the summer-stratified period in Lake Washington. Finally, we conclude by underscoring the importance of explicitly acknowledging the uncertainty in ecological forecasts to the management of freshwater ecosystems under a changing global environment.     

Stable C and N isotope analysis elucidated the importance of zooplankton in a tropical seagrass bed of Santiago Island,Northwestern Philippines

The role of zooplankton in a tropical seagrass ecosystem was investigated in milkfish farms pollution-impacted and -unimpacted seagrass beds in Santiago Island coral reefs, Northwestern Philippines. The aim was to compare between the two sites: (1) abiotic factors and zooplankton community parameters, and (2) the trophic structure using C and N stable isotopes. Low water (98–119?mV) and sediment (–121 to ?138?mV) Oxidation Reduction Potential values indicated a reducing environment in the impacted site. Zooplankton in the impacted site showed the typical community response to eutrophication (low diversity, but high total abundance due to the dominance of the cyclopoid copepod Oithona oculata), generally few elevated δ¹⁵N values, but a significant shift towards depleted ¹³C due to the organic enrichment of fish-farm feeds. Apart from suggesting a highly complex food web with POM and zooplankton as main food sources in the unimpacted site, the Bayesian mixing model simulation generated reduced complexity in feeding interactions between basal sources, zooplankton, and fish including adults of a key fish species, Siganus fuscescens, in the impacted sites. In this study, C and N stable isotope analysis has clarified the importance of zooplankton as fish prey in a seagrass bed food web.     

Carbon-based balanced trophic structure and flows in the offshore Lake Ontario food web before (1987-1991) and after (2001-2005) invasion-induced ecosystem change

Replicate mass-balanced solutions to Ecopath models describing carbon-based trophic structures and flows were developed for the Lake Ontario offshore food web before and after invasion-induced disruption. The food webs link two pathways of energy and matter flow: the grazing chain (phytoplankton-zooplankton-fish) and the microbial loop (bacteria-protozoans) and include 19 species-groups and three detrital groups. Mass-balance was achieved by using constrained optimization techniques to randomly vary initial estimates of biomass and diet composition. After the invasion, production declined for all trophic levels and species-groups except Chinook salmon. The trophic level (TL) increased for smelt, adult sculpin, adult alewife and Chinook salmon. Changes to ecotrophic efficiencies indicate a reduction in phytoplankton grazing, increased predation pressure on Mysis, adult smelt and alewife and decreased predation pressure on protozoans. Specific resource to consumer TTE changed; increasing for protozoans (8.0-11.5%), Mysis (0.6-1.0%), and Chinook salmon (1.0-2.3%) and other salmonines (0.4-0.5%) and decreasing for zooplankton (20.2-15.1%), prey-fish (9.7-8.8%), and benthos (1.7-0.6%). Direct trophic influences of recent invasive species were low. The synchrony of the decline in PP and species-group production indicates strong bottom-up influence. Mass balance required an increase of two to threefold in lower trophic level biomass and production, confirming a previously observed paradoxical deficit in lower trophic level production. Analysis of food web changes suggest hypotheses that may apply to other similar large pelagic systems including, (1) as pelagic primary productivity declines, overgrazing of zooplankton results in an increase in protozoan production and a loss of trophic transfer efficiency, (2) habitat and food web changes increased Mysis predation on Diporeia and contributed to their recent decline, and (3) production of Chinook salmon, the primary piscivore, was uncoupled from pelagic production processes. This study demonstrates the value of food web models to better understand the impact of invasive species and to develop novel hypotheses concerning trophic influences.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

洱海浮游桡足类群落结构对季节性休渔的响应

为了解洱海浮游桡足类群落结构的季节动态特征,于2009年2~11月对洱海浮游桡足类进行逐月采样.结果显示:洱海浮游桡足类生物量的季节性分布呈"双峰型",峰值分别出现在休渔期中期和捕鱼期中期.在休渔期中期,桡足类以杂食性的剑水蚤Mesocyclops dissimilis为优势种;在捕鱼期中期,则以滤食性的哲水蚤舌状叶镖水蚤为优势种.季节性休渔不仅导致鱼类对桡足类的捕食压力出现季节性变化,而且通过影响枝角类的群落结构,改变枝角类对桡足类的竞争压力.在相对较低的捕食压力和较大的食物竞争压力影响下,桡足类以杂食性的剑水蚤为优势类群;在相对较大的捕食压力和较低的食物竞争压力下,以滤食性的哲水蚤为优势类群.     

Study of the nutrient and plankton dynamics in Lake Tanganyika using a reduced-gravity model

An eco-hydrodynamic (ECOH) model is proposed for Lake Tanganyika to study the plankton productivity. The hydrodynamic sub-model solves the non-linear, reduced-gravity equations in which wind is the dominant forcing. The ecological sub-model for the epilimnion comprises nutrients, primary production, phytoplankton biomass and zooplankton biomass. In the absence of significant terrestrial input of nutrients, the nutrient loss is compensated for by seasonal, wind-driven, turbulent entrainment of nutrient-rich hypolimnion water into the epilimnion, which gives rise to high plankton productivity twice in the year, during the transition between two seasons. Model simulations predict well the seasonal contrasts of the measured physical and ecological parameters. Numerical tests indicate that the half saturation constant for grazing by zooplankton and the fish predation rate on zooplankton affect the zooplankton biomass measurably more than that of phytoplankton biomass. This work has implications for the application of this model to predict the climatological biological productivity of Lake Tanganyika.     

Using social network analysis tools in ecology: Markov process transition models applied to the seasonal trophic network dynamics of the Chesapeake Bay

Ecosystem components interact in complex ways and change over time due to a variety of both internal and external influences (climate change, season cycles, human impacts). Such processes need to be modeled dynamically using appropriate statistical methods for assessing change in network structure. Here we use visualizations and statistical models of network dynamics to understand seasonal changes in the trophic network model described by Baird and Ulanowicz [Baird, D., Ulanowicz, R.E., 1989. Seasonal dynamics of the Chesapeake Bay ecosystem. Ecol. Monogr. 501 (59), 329–364] for the Chesapeake Bay (USA). Visualizations of carbon flow networks were created for each season by using a network graphic analysis tool (NETDRAW). The structural relations of the pelagic and benthic compartments (nodes) in each seasonal network were displayed in a two-dimensional space using spring-embedder analyses with nodes color-coded for habitat associations (benthic or pelagic). The most complex network was summer, when pelagic species such as sea nettles, larval fishes, and carnivorous fishes immigrate into Chesapeake Bay and consume prey largely from the plankton and to some extent the benthos. Winter was the simplest of the seasonal networks, and exhibited the highest ascendency, with fewest nodes present and with most of the flows shifting to the benthic bacteria and sediment POC compartments. This shift in system complexity corresponds with a shift from a pelagic- to benthic-dominated system over the seasonal cycle, suggesting that winter is a mostly closed system, relying on internal cycling rather than external input. Network visualization tools are useful in assessing temporal and spatial changes in food web networks, which can be explored for patterns that can be tested using statistical approaches. A simulation-based continuous-time Markov Chain model called SIENA was used to determine the dynamic structural changes in the trophic network across phases of the annual cycle in a statistical as opposed to a visual assessment. There was a significant decrease in outdegree (prey nodes with reduced link density) and an increase in the number of transitive triples (a triad in which i chooses j and h, and j also chooses h, mostly connected via the non-living detritus nodes in position i), suggesting the Chesapeake Bay is a simpler, but structurally more efficient, ecosystem in the winter than in the summer. As in the visual analysis, this shift in system complexity corresponds with a shift from a pelagic to a more benthic-dominated system from summer to winter. Both the SIENA model and the visualization in NETDRAW support the conclusions of Baird and Ulanowicz [Baird, D., Ulanowicz, R.E., 1989. Seasonal dynamics of the Chesapeake Bay ecosystem. Ecol. Monogr. 501 (59), 329–364] that there was an increase in the Chesapeake Bay ecosystem's ascendancy in the winter. We explain such reduced complexity in winter as a system response to lowered temperature and decreased solar energy input, which causes a decline in the production of new carbon, forcing nodes to go extinct; this causes a change in the structure of the system, making it simpler and more efficient than in summer. It appears that the seasonal dynamics of the trophic structure of Chesapeake Bay can be modeled effectively using the SIENA statistical model for network change.     

Feeding behavior of adult <Emphasis Type="Italic">Vinciguerria nimbaria</Emphasis> (Phosichthyidae), in the tropical Atlantic (0°–4°N, 15°W)

Adult Vinciguerria nimbaria are the main prey of tuna during the tuna fishing season (late autumn and winter) in the equatorial Atlantic (0–4°N, and ~15°W). V. nimbaria trophic behavior in the fishing grounds was studied in relation to hydrobiological factors to determine its role in the trophic food web. Sampling stations spaced by 20 nautical miles were set up along a 15°W north–south transect from 4°N to 0°40S. At each station, the temperature and vertical fluorescence profiles were recorded. Nitrate and chlorophyll a analyses were performed on water sampled at different levels in the euphotic zone. Vertical plankton hauls were carried out at depths of 0–100 and 0–200 m using a standard WP2 net fitted with a 200-μm mesh gauze. Vinciguerria nimbaria adults were collected using a young-fish mid-water trawl net (10 × 15 m opening mouth, 10 mm cod end mesh). The weight of the stomach contents, the stomach fullness index, the number of prey, the frequency of occurrence and the prey preponderance were recorded for 20 fish from each haul. An oligotrophic typical tropical structure (TTS) was found between 1° and 4°N where small zooplankton was relatively abundant above or near the thermocline. In the TTS, V. nimbaria behaved as an epipelagic fish, feeding on the dominant small prey during the daytime. In turn, it was a prey for tuna. In the equatorial zone, where zooplankton was more abundant than in the north equatorial zone, V. nimbaria behaved as a mesopelagic fish and as an opportunistic mesozooplankton feeder. It consumed a wide range of sizes of food, feeding on the most abundant species of zooplankton as well as the largest zooplankton species, possibly while migrating towards the surface in the late afternoon or in the deep layer.     

NEATS: A Network Economics Approach to Trophic Systems

The main principle of the economic approach to a trophic system we propose here lies in assuming that there is a transfer of food along a path between a prey and a predator if, for the predator, the benefits are greater than costs of predation on this path. Conversely, if the costs exceed the benefits, there are no flows. This trade-off, considered all along the food chains of an ecosystem, together with ecological processes (assimilation, somatic maintenance) results in a model coupling mass balance equations (biological constraints) and complementarity principles (Walras’ law). Here is the core of the Network Economics Approach to Trophic Systems (NEATS).     

EcoTroph: Modelling marine ecosystem functioning and impact of fishing

EcoTroph (ET) is a model articulated around the idea that the functioning of aquatic ecosystems may be viewed as a biomass flow moving from lower to higher trophic levels, due to predation and ontogenetic processes. Thus, we show that the ecosystem biomass present at a given trophic level may be estimated from two simple equations, one describing biomass flow, the other their kinetics (which quantifies the velocity of biomass transfers towards top predators). The flow kinetic of prey partly depends on the abundance of their predators, and a top-down equation expressing this is included in the model. Based on these relationships, we simulated the impact on a virtual ecosystem of various exploitation patterns. Specifically, we show that the EcoTroph approach is able to mimic the effects of increased fishing effort on ecosystem biomass expected from theory. Particularly, the model exhibits complex patterns observed in field data, notably cascading effects and ‘fishing down the food web’. EcoTroph also provides diagnostic tools for examining the relationships between catch and fishing effort at the ecosystem scale and the effects of strong top-down controls and fast-flow kinetics on ecosystems resilience. Finally, a dynamic version of the model is derived from the steady-state version, thus allowing simulations of time series of ecosystem biomass and catches. Using this dynamic model, we explore the propagation of environmental variability in the food web, and illustrated how exploitation can induce a decrease of ecosystem stability. The potential for applying EcoTroph to specific ecosystems, based on field data, and similarities between EcoTroph and Ecopath with Ecosim (EwE) are finally discussed.     

Long-term trends in the trophic structure of the North Sea fish community: evidence from stable-isotope analysis, size-spectra and community metrics

Fishing has wide-ranging impacts on marine ecosystems. One of the most pervasive signs of intensive fishing is "fishing down the food web", with landings increasingly dominated by smaller species from lower trophic levels. Decreases in the trophic level of landings are assumed to reflect those in fish communities, because size-selective mortality causes decreases in the relative abundance of larger species and in mean body size within species. However, existing analyses of fishing impacts on the trophic level of fish communities have focused on the role of changes in species composition rather than size composition. This will provide a biased assessment of the magnitude of fishing impacts, because fishes feed at different trophic levels as they grow. Here, we combine body size versus trophic level relationships for North Sea fishes (trophic level assessed using nitrogen stable-isotope analysis) with species-size-abundance data from two time-series of trawl-survey data (whole North Sea 1982-2000, central and northern North Sea 1925-1996) to predict long-term trends in the trophic structure of the North Sea fish community. Analyses of the 1982-2000 time-series showed that there was a slow but progressive decline in the trophic level of the demersal community, while there was no trend in the trophic level of the combined pelagic and demersal community. Analyses of the longer time-series suggested that there was no trend in the trophic level of the demersal community. We related temporal changes in trophic level to temporal changes in the slopes of normalised biomass size-spectra (which theoretically represent the trophic structure of the community), mean log₂ body mass and mean log₂ maximum body mass. While the size-based metrics of community structure showed long-term trends that were consistent with the effects of increased fishery exploitation, these trends were only correlated with trophic level for the demersal community. Our analysis suggests that the effects of fishing on the trophic structure of fish communities can be much more complex than previously assumed. This is a consequence of sampled communities not reflecting all the pathways of energy transfer in a marine ecosystem and of the absence of historical data on temporal and spatial changes in the trophic level of individuals. For the North Sea fish community, changes in size structure due to the differential effects of fishing on species and populations with different life histories are a stronger and more universal indicator of fishing effects than changes in mean trophic level.     

The impact of detritivorous fishes on a mangrove estuarine system

Sundarban mangrove ecosystem in India is one of the largest detritus based ecosystem of the world and it supplies the detritus and nutrients to the adjacent Hooghly-Matla estuarine complex. In this estuary a group of fish completely detritivorous in nature, belonging to the genus Mugil spp. is present. This group of fish is expected to have important effects on the trophic dynamics of ecosystems, but exact nature of these effects is not known. In order to study the impact that detritivory by fish may have on the estuarine food chain, we developed mathematical formulations. We run two models, one with phytoplankton, zooplankton, carnivorous fish, detritus and nutrient and without this group of fish and a second one after including this fish in the system. In our model this group of fish has no major impact on primary productions of the estuarine system but has extensive role in total fish production. Coexistence of detritivorous fish and carnivorous fish occurs within reasonable parameter range. We have tested different growth rates of phytoplankton, grazing rates and predation rates of zooplankton, detritivorous fish and carnivorous fish for total system equilibrium. Carnivorous fish predation rate on detritivorous fish and detritivorous fish grazing rate on detritus are very important. Different foraging ratios are also tested in this study. Foraging preference of carnivorous fish on detritivorous fish appears significant for the system equilibrium.     

The functional biology and trophic role of krill (Thysanoessa raschii) in a Greenlandic fjord

Despite being a key zooplankton group, knowledge on krill biology from the Arctic is inadequate. The present study examine the functional biology and evaluate the trophic role of krill in the Godth?bsfjord (64°N, 51°W) SW Greenland, through a combination of fieldwork and laboratory experiments. Krill biomass was highest in the middle fjord and inner fjord, whereas no krill was found offshore. The dominating species Thysanoessa raschii revealed a type III functional response when fed with the diatom Thalassiosira weissflogii. At food saturation, T. raschii exhibited a daily ration of 1% body C d^?1. Furthermore, T. raschii was capable of exploiting plankton cells from 5 to 400???m, covering several trophic levels of the pelagic food web. The calculated grazing impact by T. raschii on the fjord plankton community was negligible. However, the schooling and migratory behaviour of krill will concentrate and elevate the grazing in specific areas of the euphotic zone.     

Impact of Arenicola marina (Polychaeta) on the microbial assemblages and meiobenthos in a marine intertidal flat

The benthic microbial food web can be responsible for a large proportion of benthic carbon cycling yet there are few data on the trophic interactions between this food web and macrobenthos. A large-scale field experiment was conducted to investigate effects of eliminating the polychaete Arenicola marina on benthic microbes (prokaryotes, heterotrophic and autotrophic protists) and metazoan meiofauna in a marine intertidal flat of the North Sea, Germany. Over a period of 2 years, quantity and composition of micro- and meiobenthos from unmanipulated sites were compared to those from sites deplete of lugworms. These grazer treatments were cross-classified with different sediment characteristics (low- and mid-intertidal areas). Lugworm removal resulted in an initial increase in abundance of prokaryotes and nanoflagellates, which became less pronounced in the second year. Ciliates were not affected quantitatively, but in the absence of lugworms, diversity and the proportion of carnivorous forms increased. Meiobenthos (nematodes, ostracods and copepods) were affected only moderately. The observed changes are probably due to a combination of release from grazing/predation pressure, changes in the species composition of higher trophic levels (namely large polychaetes) and altered environmental conditions (such as depth of the oxygenated layer and sediment grain size). Spatial differences between sites of different tidal exposure/grain size appeared to be as large as temporal differences during the 2 years following the manipulation of the system. We conclude that in intertidal sediments, indirect effects due to habitat transformation are as important as direct biological interactions (grazing pressure and competition) for the dynamics of the benthic microbial food web.     

Assessment of impacts of invasive fishes on the food web structure and ecosystem properties of a tropical reservoir in India

A network model of trophic interactions in a tropical reservoir in India was developed with the objective to quantify matter and energy flows between system components and to study the impact of invasive fishes on the ecosystem. Structure of flows and their distribution within and between trophic levels were analysed by aggregating single flows into combined flows for discrete trophic levels. The trophic flows primarily occurred in the first four trophic level (TL) and the food web structure in this reservoir ecosystem was characterized by the dominance of low TL organisms, with the highest TL of only 3.57 for the top predator. Highest system omnivory index (SOI) was observed for indigenous catfishes (0.422), followed by the exotic fish Mozambique Tilapia (0.402). Nile Tilapia and Pearl spots show the highest niche overlap which suggests high competition for similar resources. The mixed trophic impact routine reveals that an increase in the abundance of the African catfish would negatively impact almost all fish groups such as Indian major carps, Pearl spots, indigenous catfishes and Tilapines. The other invasive fish Mozambique Tilapia adversely affects the indigenous catfishes. The most interesting observation in this study is that the most dominant invasive fish in this reservoir, the Nile Tilapia does not negatively impact any of the fish groups. In fact it positively impacts the Indian major carps. The direct and indirect effects of predation between system components (i.e. fish, invertebrates, phytoplankton and detritus) are quantitatively described and the possible influence and role in the ecosystem's functioning of the invasive fish species are discussed.