An analysis of temporal variability in abundance, diversity and growth rates within the coastal ichthyoplankton assemblage of South Georgia (sub-Antarctic)

The study of spatial and temporal distribution and diversity of ichthyoplankton (fish eggs and larvae) can provide fisheries-independent information on the population dynamics and recruitment processes of marine fish species. Ichthyoplankton studies in the Southern Ocean have to date been largely constrained to the summer months. We analysed ichthyoplankton data collected from a year round, long term (2002–2008), plankton trawl sampling programme in a large fjord system (Cumberland Bay) at South Georgia, sub-Antarctic (54.25°S, 36.5°W) to assess temporal changes in larval fish diversity and abundance. Larvae of 22 species, representing nine families, were identified although three, Krefftichthys anderssoni (Myctophidae), Lepidonotothen nudifrons/Trematomus hansoni (Nototheniidae) and Champsocephalus gunnari (Channichthyidae), dominated abundance in all years. Significant seasonal and interannual differences in the larval fish assemblage were revealed by multivariate analyses. Estimates of larval growth are provided for five abundant species. Considerable inter-specific differences in relative larval growth rate were recorded but interannual variability within species was small. However, in the commercially important C. gunnari, multiple larval cohorts, representing a protracted spawning season, were observed to grow at different rates, and this may be related to temperature and/or food availability. A comparison with historical growth data from South Georgia suggests there has been little change in growth rate for the main species over the last three decades.     

Veligers of the invasive bivalve Limnoperna fortunei in the diet of indigenous fish larvae in a eutrophic subtropical reservoir

Larval fish development depends largely on their ability to capture and ingest food items, and on food availability. In this context, invasive species, eutrophication and river impoundments have complex impacts on fish larvae. Using samples collected in 2005–2009 in the Salto Grande reservoir (Argentina–Uruguay), periodically affected by cyanobacterial blooms, we studied the impact of the larvae of the exotic bivalve Limnoperna fortunei (Dunker, 1857) (Bivalvia) on larval fish diets. Compared with other nearby waterbodies, the abundance of fish larvae was scarcer in the reservoir, especially during algal bloom periods. Only 20% of the larval fish with gut contents fed on L. fortunei veligers. Seven fish taxa (of a total of 12) consumed veligers of L. fortunei, but only two showed a preference for this prey. Taxonomic changes in the larval fish assemblages due to the river's impoundment, and temporal uncoupling between veliger densities (affected by the toxigenic effects of Microcystis spp.) and ichthyoplankton could account for the comparatively low trophic importance of the invasive bivalve's veligers. These results reflect the complexity of interactions brought about when the same invasive species invades different environments, underscoring that the impacts involved depend as much on the invader, as on the regional and ecological settings of the area invaded.     

Climate effects on fish body size–trophic position relationship depend on ecosystem type

The energetic demand of consumers increases with body size and temperature. This implies that energetic constraints may limit the trophic position of larger consumers, which is expected to be lower in tropical than in temperate regions to compensate for energy limitation. Using a global dataset of 3635 marine and freshwater ray‐finned fish species, we addressed if and how climate affects the fish body size–trophic position relationship in both freshwater and marine ecosystems, while controlling for the effects of taxonomic affiliation. We observed significant fish body size–trophic position relationships for different ecosystems. However, only in freshwater systems larger tropical fish presented a significantly lower trophic position than their temperate counterparts. Climate did not affect the fish body size–trophic position relationship in marine systems. Our results suggest that larger tropical freshwater fish may compensate for higher energetic constraints feeding at lower trophic positions, compared to their temperate counterparts of similar body size. The lower latitudinal temperature range in marine ecosystems and/or their larger ecosystem size may attenuate and/or compensate for the energy limitation of larger marine fish. Based on our results, temperature may determine macroecological patterns of aquatic food webs, but its effect is contingent on ecosystem type. We suggest that freshwater ecosystems may be more sensitive to warming‐induced alterations in food web topology and food chain length than marine ecosystems.     

重金属在海洋食物链中的传递

近年来 ,金属在不同海洋食物链中摄食富集的定量研究得到越来越多的关注。自然环境中生物体内金属的浓度并不一定和生物在食物链中所处的营养级有相关关系 ,金属在生物体内的富集还受到生物的同化、排出等过程以及其它生理生化因子的影响。在经典的海洋浮游生物食物链中 (浮游植物→桡足类→鱼类 ) ,桡足类往往可以很有效地排出体内的金属 ,同时鱼类的金属同化率又很低 ,所以该食物链中金属的浓度随食物链水平增加而减少。目前研究发现只有甲基汞和铯 Cs会被食物链所放大。在以腹足动物为顶级捕食者的底栖食物链中 ,因为生物结合金属的效率很高 ,高同化率和低排出率导致金属浓度在生物体内得到放大。重金属在生物体内的可利用性可以通过测定同化率、排出率等参数、并结合考虑生物对该金属的消化行为 ,运用一个简易的动态模型来估算。已有的研究中人们多考虑金属的化学性质对食物链传递的影响。着重介绍了近年来国外对金属在不同海洋食物链 (底栖和浮游 )中的传递的研究成果 ,强调在金属的生物可利用性评估中 ,要充分考虑到动物的生理、生化过程的影响 ,同时也必须认识到不同的海洋生物有着复杂且不同的金属代谢机制     

The importance of silicon for marine production*

A review of silicon, with emphasis on its forms, uptake, dissolution and role in marine primary production, is given. The importance of silicon in marine food webs is discussed, as well as the concentrations of silicon in various areas and the importance of changing N:Si:P ratios. The methodology for measuring silicate transformations has recently been improved by the introduction of the highly enriched ³²Si isotope. Results from uptake experiments using ³²Si in nutrient rich coastal water and in open ocean are presented. The uptake kinetic experiments showed that the silicate uptake usually is unsaturated. We propose that closer attention in the future should be paid to the importance of balanced nutrient composition as well as nutrient supply dynamics for the development of eutrophication versus efficient trophic transfer and fish production in nutrient enriched systems. Close attention should also be paid to the mechanisms that reduce the inputs of silicate to coastal waters.     

Fish as predators and prey: DNA-based assessment of their role in food webs

Fish are both consumers and prey, and as such part of a dynamic trophic network. Measuring how they are trophically linked, both directly and indirectly, to other species is vital to comprehend the mechanisms driving alterations in fish communities in space and time. Moreover, this knowledge also helps to understand how fish communities respond to environmental change and delivers important information for implementing management of fish stocks. DNA-based methods have significantly widened our ability to assess trophic interactions in both marine and freshwater systems and they possess a range of advantages over other approaches in diet analysis. In this review we provide an overview of different DNA-based methods that have been used to assess trophic interactions of fish as consumers and prey. We consider the practicalities and limitations, and emphasize critical aspects when analysing molecular derived trophic data. We exemplify how molecular techniques have been employed to unravel food web interactions involving fish as consumers and prey. In addition to the exciting opportunities DNA-based approaches offer, we identify current challenges and future prospects for assessing fish food webs where DNA-based approaches will play an important role.     

Trophic Structure of One Deep-sea Benthic Fish Community in the Eastern Canadian Arctic: Application of Food, Parasites and Multivariate Analysis

Synopsis As new arctic marine fisheries develop there is need for a comprehensive ecosystem approach to long-term management. This approach recognizes the importance of community interactions such as food web structure and trophic patterns. We determined whether hierarchical clustering (guild formation) is an effective method of trophic evaluation in deep-sea Artic fish communities using stomach content and parasite data with size class, and evaluated the application of endohelminth communities (parasite species transmitted in the food) as indicators of trophic status. Cluster analysis using food group abundance with size class of fish revealed the presence of 11 guilds within the community, however the same analysis using parasite data showed little correlation between food and parasites. Redundancy analysis (RDA) within the 11 guilds also revealed no significant correlations between food group and parasite abundance suggesting that this type of ordination is not suited for environments containing mainly generalist feeders. RDA of individual taxa without a priori guild designation found that taxa in benthic deep-sea communities are defined by their ability to exploit prey species in more than one habitat zone. Benthic fish species were significantly correlated with benthic food groups and parasites that utilize benthic intermediate hosts whereas benthopelagic–pelagic species fed on a higher diversity of prey species and were infected by a larger number of non-host specific parasites. Eigenanalysis and Monte Carlo results showed that parasites and food groups are highly correlated, indicating that parasite community analysis is an effective tool for predicting feeding strategies in Arctic marine environments. It also suggests that in most cases endoparasite infections alone could be used for trophic evaluation in the absence of stomach content data.     

Functional morphogenesis from embryos to adults: Late development shapes trophic niche in coral reef damselfishes

The damselfishes are one of the dominant coral reef fish lineages. Their ecological diversification has involved repeated transitions between pelagic feeding using fast bites and benthic feeding using forceful bites. A highly‐integrative approach that combined gene expression assays, shape analyses, and high‐speed video analyses was used to examine the development of trophic morphology in embryonic, larval, juvenile, and adult damselfishes. The anatomical characters that distinguish pelagic‐feeding and benthic‐feeding species do not appear until after larval development. Neither patterns of embryonic jaw morphogenesis, larval skull shapes nor larval bite mechanics significantly distinguished damselfishes from different adult trophic guilds. Analyses of skull shape and feeding performance identified two important transitions in the trophic development of a single species (the orange clownfish; Amphiprion percula): (a) a pronounced transformation in feeding mechanics during metamorphosis; and (b) more protracted cranial remodeling over the course of juvenile development. The results of this study indicate that changes in postlarval morphogenesis have played an important role in damselfish evolution. This is likely to be true for other fish lineages, particularly if they consist of marine species, the majority of which have planktonic larvae with different functional requirements for feeding in comparison to their adult forms.     

Ichthyoplankton dynamics and biodiversity in the Gulf of Alaska: Responses to environmental change

Climate variation can cause major changes in the marine food web. We analyzed over 24 years of ichthyoplankton data from the Gulf of Alaska to evaluate lower trophic level responses to environmental change and judge their usefulness as ecological indicators. We standardized abundance data for each of 77 ichthyoplankton taxa, and used the Bray-Curtis distance measure and Flexible Beta linkage method, which grouped them into 22 discrete clusters. Variance Partitioning Analysis stressed the importance of geographical and seasonal processes for ichthyoplankton dynamics, and helped us identify the specific region(s) and month(s) for each response variable (cluster abundance, diversity) in which annual variation was maximized. Response variables were linked to environmental explanatory variables (atmospheric pressure, temperature, salinity and circulation indices) by Canonical Correspondence Analysis. The North Pacific Index (atmospheric pressure) and meso-scale climate variables like the El Niño Index (temperature), wind, and freshwater input (circulation) had the strongest impacts on ichthyoplankton species clusters. Specifically, the El Niño Index was negatively correlated with several ichthyoplankton clusters that were dominated by cold water species. Circulation was predominantly positively related to diversity and ichthyoplankton clusters, with the exception of clusters that mainly consisted of offshore taxa. The immediate response of ichthyoplankton to environmental forcing might make them suitable ecological indicators of environmental change although additional work is needed to assess affects on survival and recruitment.     

Variability in the trophic position of larval fish in a coastal pelagic ecosystem based on stable isotope analysis

We used stable isotopes of carbon (C) and nitrogen (N) to investigatethe trophic position of six species of larval fish in the pelagicecosystem of coastal Newfoundland. Isotope profiles from phytoplankton,net plankton and macrozooplankton were consistent with previousstudies. All species of larval fish showed a length-dependentshift in ¹³C that indicates a move to a pelagic diet from thecombined pelagic and demersal eating habits of the adult spawners.The trophic position of four larval fish species (American plaice,yellowtail flounder, cunner, radiated shanny) was consistentwith them feeding primarily on copepods, as expected from stomachcontent analysis. The ¹⁵N-based trophic position of larval witchflounder and capelin indicates that they feed significantlyon phytoplankton and heterotrophic protists from the microbialloop, respectively; this evidence contrasts with stomach contentanalysis. Although links between larval fish and the microbialloop are not considered as common as is the link with crustaceanzooplankton, this and other studies challenge the long-heldbelief that marine fish larvae feed effectively exclusivelyon copepods.     

Seasonal plankton cycles in a temperate fjord and comments on the match-mismatch hypothesis

As a field test of the ‘match-mismatch hypothesis’(Cushing, 1972, 1975, 1990), we sampled the ichthyoplanktonand different size fractions of zooplankton at two stationsin a temperate fjord (Dabob Bay, Washington, USA) between May1985 and October 1987. We present the results of this studytogether with historical data on seasonal abundances of phytoplanktonand invertebrate predators. The peak abundance of fish larvaein late winter/early spring preceded the peak abundance of preyorganisms by an average of 3 months. Correlation analysis resultedin one negatively significant and three non-significant correlationsbetween ichthyoplankton and appropriately sized prey organisms.These field observations, showing a distinct temporal lag betweenabundances of larval fish and their zooplankton prey, do notsupport the match-mismatch hypothesis as originally formulated.We use these field data and a simulation model of the growthof young fish to discuss the role of two alternative processes—foodlimitation during late larval/juvenile stages and predation—thatmay constrain the timing of spawning of marine fish and whichmay have implications for recruitment variability as well. Thisleads us to propose that the match-mismatch hypothesis be extendedto include a ‘critical period’ that lasts longerand occurs later in the life history of marine fish than haspreviously been supposed.     

Vulnerability and diet breadth predict larval and adult parasite diversity in fish of the Bothnian Bay

Recent studies of aquatic food webs show that parasite diversity is concentrated in nodes that likely favour transmission. Various aspects of parasite diversity have been observed to be correlated with the trophic level, size, diet breadth, and vulnerability to predation of hosts. However, no study has attempted to distinguish among all four correlates, which may have differential importance for trophically transmitted parasites occurring as larvae or adults. We searched for factors that best predict the diversity of larval and adult endoparasites in 4105 fish in 25 species studied over a three-year period in the Bothnian Bay, Finland. Local predator–prey relationships were determined from stomach contents, parasites, and published data in 8,229 fish in 31 species and in seals and piscivorous birds. Fish that consumed more species of prey had more diverse trophically transmitted adult parasites. Larval parasite diversity increased with the diversity of both prey and predators, but increases in predator diversity had a greater effect. Prey diversity was more strongly associated with the diversity of adult parasites than with that of larvae. The proportion of parasite species present as larvae in a host species was correlated with the diversity of its predators. There was a notable lack of association with the diversity of any parasite guild and fish length, trophic level, or trophic category. Thus, diversity is associated with different nodal properties in larval and adult parasites, and association strengths also differ, strongly reflecting the life cycles of parasites and the food chains they follow to complete transmission.     

The effects of predation by planktivorous juvenile fish on the microbial food web

The feeding impact of planktivorous fish on microbial organisms is still poorly understood. We followed the seasonal dynamics of the food web in two natural fishponds for two years: one was stocked with planktivorous whitefish while the other had no planktivorous fish. The aim of the study was the simultaneous assessment of the feeding behaviours of planktivorous fish and of bacterivorous meta-/protozooplankters. We hypothesized that in the presence of planktivorous fish there would be fewer metazooplankton, more protozoans and decreased numbers of bacteria. Our results showed that the amount of metazooplankton eaten by the fish was indeed negatively correlated with metazooplankton biomass. The feeding impact of planktivorous fish in shaping the microbial loop was remarkable. The main grazers of bacteria in the fishpond were ciliates, whereas in the pond without fish these were heterotrophic nanoflagellates. In the fishless pond the role of the top predator shifted to the predaceous metazooplankter Leptodora kindtii which controlled the abundance of herbivorous metazooplankters. We found a negative relationship between the number of bacteria and flagellates in the fishless pond, while the number of bacterivorous ciliates was suppressed by predaceous ciliates. Therefore the bacteria-grazing activity was higher in the absence of planktivorous fish.     

The influence of autotrophy,heterotrophy and temperature on pelagic food web efficiency in a brackish water system

Climate change has been suggested to lead to higher temperature and increased heterotrophy in aquatic systems. The aim of this study was to test how these two factors affect metazooplankton and food web efficiency (FWE was defined as metazooplankton production divided by basal production). We tested the following hypotheses: (1) that lower metazooplankton production and lower FWE would be found in a food web based on heterotrophic production (bacteria) relative to one based on autotrophic production (phytoplankton), since the former induces a larger number of trophic levels; (2) the metazooplankton in the heterotrophic food web would contain less essential fatty acids than those from the autotrophic food web; and (3) that higher temperature would lead to increased FWE. To test these hypotheses, a mesocosm experiment was established at two different temperatures (5 and 10°C) with a dominance of either autotrophic (NP) or heterotrophic basal production (CNP). Metazooplankton production increased with temperature, but was not significantly affected by differences in basal production. However, increased heterotrophy did lead to decreased fatty acid content and lower individual weight in the zooplankton. FWE increased with autotrophy and temperature in the following order: 5CNP < 10CNP < 5NP < 10NP. Our results indicate that in the climate change scenario we considered, the temperature will have a positive effect on FWE, whereas the increase in heterotrophy will have a negative effect on FWE. Furthermore, the quality and individual weight of the metazooplankton will be reduced, with possible negative effects on higher trophic levels.     

A model of the predatory impact of larval marine fish on the population dynamics of their zooplankton prey

A simulation model of the population dynamics of two speciesof calanoid copepods (Calanus.r pacificus and Pseudocalanussp.) was forced with predation pressure from a genetic, hypotheticalpopulation of larval marine fish. Results of the model are sensitiveto changes in parameters describing the dynamics of both predatorand prey populations, including initial numbers, fecundity,growth, mortality, size of prey organisms and feeding selectivityof the predators; the relative importance of these parametersis tested by way of a brute-force sensitivity analysis. Usingresults from recent ichthyoplankton surveys in Dabob Bay, WA,USA, the model was also forced with predation from populationsof larval Pacific herring (Clupea harengus pallasi) and Pacificwhiting (Merluccius productus). Results of the various simulationruns lead to the conclusion that marine fish larvae can significantlyimpact the population dynamics of their calanoid copepod prey,but that the magnitude of this impact is highly dependent onspecies-specific values of various population parameters.     

Climate change alters the structure of arctic marine food webs due to poleward shifts of boreal generalists

Climate-driven poleward shifts, leading to changes in species composition and relative abundances, have been recently documented in the Arctic. Among the fastest moving species are boreal generalist fish which are expected to affect arctic marine food web structure and ecosystem functioning substantially. Here, we address structural changes at the food web level induced by poleward shifts via topological network analysis of highly resolved boreal and arctic food webs of the Barents Sea. We detected considerable differences in structural properties and link configuration between the boreal and the arctic food webs, the latter being more modular and less connected. We found that a main characteristic of the boreal fish moving poleward into the arctic region of the Barents Sea is high generalism, a property that increases connectance and reduces modularity in the arctic marine food web. Our results reveal that habitats form natural boundaries for food web modules, and that generalists play an important functional role in coupling pelagic and benthic modules. We posit that these habitat couplers have the potential to promote the transfer of energy and matter between habitats, but also the spread of pertubations, thereby changing arctic marine food web structure considerably with implications for ecosystem dynamics and functioning.     

The role of hyperiid parasites as a trophic link between jellyfish and fishes

The trophic interactions between the scyphozoan medusa Chrysaora plocamia and the palm ruff Seriolella violacea were investigated off northern Chile and showed that large numbers of hyperiid amphipods parasitizing the medusa may channel energy back to the fishes, which feed on the parasites. The biomass of hyperiids eaten by the fish was a function of the biomass of hyperiids parasitizing the medusa. This temporally available food supply may enhance fish recruitment. The large number of hyperiids parasitizing diverse jellyfish species represents a missing trophic link in current efforts to understand the effects of jellyfish blooms on marine food webs.     

大型年长鱼类对海洋生态系统生物资源养护的作用

随着人口增长和渔业资源需求的上升,人类沿着食物网营养级自上向下捕捞渔业资源的趋势仍在持续,导致海洋生物多样性丧失及其生态功能与服务价值迅速下降。由捕捞活动造成的渔业诱导进化进一步导致鱼类资源小型化已受到国内外学者的广泛关注。从大型年长鱼类这一独特视角出发,综述了大型年长鱼类在种群繁衍过程中的作用,其具备巨大的繁殖能量输出、丰富的亲本繁殖经验以及强大的年龄组繁殖力贡献,更有利于种群的延续;大型年长鱼类对于初次性成熟亲鱼生殖洄游具有一定的引领作用,并在季节性集群繁殖过程中占据主导地位,可通过抑制小个体同类繁殖维护种群结构的稳定;大型年长鱼类在生态系统中更是占据了较高营养级和广阔的生态位宽度,面对生态环境的改变,具有更强的适应调控能力;大型年长鱼类在种群基因交流过程中亦起到十分关键的作用。保护大型年长鱼类更有利于鱼类种群的快速恢复,这为生物资源养护和渔业管理政策制定方面提供了一定的参考。     

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).     

Disentangling effects of multiple stressors on matter flow in a lake food web

Understanding the relative importance of multiple stressors is valuable to prioritize conservation and restoration measures. Yet, the effects of multiple stressors on ecosystem functioning remain largely unknown in many fresh waters. Here, we provided a methodology combining ecosystem modeling with linear regression to disentangle the effects of multiple stressors on matter flow, an important ecosystem function. Treating a shallow lake as the model ecosystem, we simulated matter flow dynamics during 1950s–2010s with different combinations of stressors by Ecopath with Ecosim (EwE) modeling and determined the relative importance of each stressor by generalized linear mixed models. We found that matter flow of the lake food web was highly dynamic, attributing to effects of multiple anthropogenic stressors. Biological invasion played the strongest role in driving the matter flow dynamics, followed by eutrophication, while biomanipulation (i.e., phytoplankton control by planktivorous fish stocking) was of little importance. Eutrophication had a stronger role on primary producers, pelagic food chain, and top predators, while biological invasion on consumers in the middle of food chains. The former was more important in driving the quantity of matter flow, while the latter on trophic transfer efficiencies. Scenario forecasting showed that reducing nutrients contents could largely shape the matter flow pattern, while biomanipulation had little effect. Our findings provided new insights into understanding the mechanistic links between anthropogenic stressors and ecosystem functioning by combining ecosystem modeling with linear regression.