Rescaling the trophic structure of marine food webs

Measures of trophic position (TP) are critical for understanding food web interactions and human‐mediated ecosystem disturbance. Nitrogen stable isotopes (δ¹⁵N) provide a powerful tool to estimate TP but are limited by a pragmatic assumption that isotope discrimination is constant (change in δ¹⁵N between predator and prey, Δ¹⁵N = 3.4‰), resulting in an additive framework that omits known Δ¹⁵N variation. Through meta‐analysis, we determine narrowing discrimination from an empirical linear relationship between experimental Δ¹⁵N and δ¹⁵N values of prey consumed. The resulting scaled Δ¹⁵N framework estimated reliable TPs of zooplanktivores to tertiary piscivores congruent with known feeding relationships that radically alters the conventional structure of marine food webs. Apex predator TP estimates were markedly higher than currently assumed by whole‐ecosystem models, indicating perceived food webs have been truncated and species‐interactions over simplified. The scaled Δ¹⁵N framework will greatly improve the accuracy of trophic estimates widely used in ecosystem‐based management.     

Population dynamics and energy budget of Marionina southerni (Cernosvitov) (Enchytraeidae,Oligochaeta) in the shallow littoral of Lake Esrom,Denmark

Marionina southerni (Cernosvitov) was numerically the dominant oligochaete in a shallow, strongly exposed surf zone locality (0–1 m depth) in the mesotrophic Lake Esrom. It comprised 21,500 ind m^–2 or nearly 50% of the total oligochaete community, which otherwise was dominated by Nais spp. and the lumbriculid Stylodrilus heringianus Claparède. M. southerni appears to have a 1-y life cycle. Individual biomass ranged from 3 to 48 µg ash free dry weight. Annual net production at the site was 5.1 kcal m^–2 (size-frequency method) with a P/B ratio of 2.5. This is about 0.5% of the estimated mean zoobenthic production in the littoral zone from 0 to 2 m depth in Lake Esrom. At 5° C M. southerni showed a constant oxygen uptake down to 25% oxygen saturation, but practically no regulatory respiration was found at 20 °C. In the field M. southerni was never found in sediment with less than 60% oxygen saturation. The annual community respiration of M. southerni was 14.2 kcal m^–2, and annual assimilation thus made up 19.3 kcal m^–2 with a net production efficiency of 26.5%.     

昆承湖优势种鱼类时空-营养生态位

为了解昆承湖优势种鱼类资源利用情况,首先利用生态位方法计算了时间、空间及营养三个资源维度的生态位宽度及重叠值,然后根据时空-营养生态位宽度值将优势种鱼类划分为广位种、中位种和窄位种,最后讨论了生态位宽度及重叠的可能原因。结果显示：刀鲚Coilia nasus、蒙古鲌Chanodichthys mongolicus、似鱎Toxabramis swinhonis、似鳊Pseudobrama simoni、鳙Hypophthalmichthys nobilis、鲢Hypophthalmichthys molitrix、花鱼骨Hemibarbus maculatus、似刺鳊鮈Paracanthobrama guichenoti、大鳍鱊Acheilognathus macropterus和鲫Carassius auratus为优势种。在时间维度：鲢的生态位宽度最大,似鳊的最小;生态位重叠具有显著意义的有24对,占总的53.33%。在空间维度,似刺鳊鮈最大,鲢最小;生态位重叠具有显著意义的有36对,占总的80%。在营养维度,最大的为鲫,最小的为花鱼骨;生态位重叠具有显著意义的有8对,占17.78%...     

Life-history strategies and production of Tinodes waeneri (L.) (Trichoptera) in Lake Esrom,Denmark

The life cycle and production of Tinodes waeneri (L.) was examined at five shallow littoral localities in Lake Esrom, Denmark, during 1979/80. Five larval instars were demonstrated. The differences in temporal composition of instars indicated three different life cycles. A bivoltine strategy was shown for the most exposed locality which also had the highest average abundance equal to 11 500 ind. m^–2. A univoltine population was found in the more sheltered NW part of the lake with an average density of 1 500 ind. m^–2. At the three remaining stations the life cycles were partly bivoltine with wintering populations of mixed cohorts and densities between 10 500 and 3 000 ind. m^–3. Two distinct flight periods with maxima in June and August were demonstrated for bi- and partly bivoltine populations. Differences in rates of growth and elimination added to the variability in life cycle. Growth rates peaked in August (5.0–7.4% d^–1) at an average temperature of 20°C. Estimates of production ranged from 1.9 to 17.5 g AFDW m^–2 a^–1 with an average of 8.0 g AFDW m^–2. The P/B ratios, which were within the expected range for uni- and bivoltine populations of trichopterans, were 5.73 for the bivoltine population and 3.31 for the univoltine population.Estimates of consumption revealed that the populations could be sustained by autochthonous epilithic production in the habitats. Variability in the quality and seasonal availability of food were judged to influence the type of life cycle.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.     

Primary productivity and trophic status of a Kashmir Himalayan Lake

The paper deals with general limnology, phytoplankton production and trophic status of a Himalayan lake of Kashmir. The primary production as measured by in situ ¹⁴C istope technique ranged from 36 to 906 mgC.m^–2 d^–1 with characteristic seasonal variation. The photosynthetic efficiency was from 0.02 to 0.48% during the same period. The yearly (1975–1976) production was estimated at 134 gC.m^–2 m^–2, which shows that the lake has not yet reached a high level of eutrophication. The waterbody has been classified as mesotrophic.Formed part of thesis for which Ph. D. was awarded to MAK by Kashmir University, India     

The Life History and Occurrence of Sialis lutaria L. (Megaloptera) in the Littoral Zone of Lake Esrom,Denmark

The life history and distribution of Sialis lutaria L. in the exposed littoral zone of Lake Esrom, Denmark were examined during 1979/80. Sialis showed an annual life cycle with mean annual abundance of 20 and 40 ind m⁻² at 0.4 and 2.0 m depth, respectively. Growth peaked during July with rates of about 10 % d⁻¹. Net annual production averaged 0.2 g DW at 0.4 m depth and 0.8 g DW at 2.0 m depth. The annual P/B-ratios were estimated in the range of 4.6-7.1.     

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.     

Bioaccumulation and trophic dilution of human pharmaceuticals across trophic positions of an effluent-dependent wadeable stream

Though pharmaceuticals are increasingly observed in a variety of organisms from coastal and inland aquatic systems, trophic transfer of pharmaceuticals in aquatic food webs have not been reported. In this study, bioaccumulation of select pharmaceuticals was investigated in a lower order effluent-dependent stream in central Texas, USA, using isotope dilution liquid chromatography–tandem mass spectrometry (MS). A fish plasma model, initially developed from laboratory studies, was tested to examine observed versus predicted internal dose of select pharmaceuticals. Pharmaceuticals accumulated to higher concentrations in invertebrates relative to fish; elevated concentrations of the antidepressant sertraline and its primary metabolite desmethylsertraline were observed in the Asian clam, Corbicula fluminea, and two unionid mussel species. Trophic positions were determined from stable isotopes (δ¹⁵N and δ¹³C) collected by isotope ratio-MS; a Bayesian mixing model was then used to estimate diet contributions towards top fish predators. Because diphenhydramine and carbamazepine were the only target compounds detected in all species examined, trophic magnification factors (TMFs) were derived to evaluate potential trophic transfer of both compounds. TMFs for diphenhydramine (0.38) and carbamazepine (1.17) indicated neither compound experienced trophic magnification, which suggests that inhalational and not dietary exposure represented the primary route of uptake by fish in this effluent-dependent stream.     

Fish abundance and trophic structure from the commercial catch in Lake Patzcuaro, Mexico

The composition, abundance and trophic structure of the fish community in Lake Patzcuaro is analyzed, using data from eight samples taken from the commercial fishery acquired between August 1990 and April 1991. A total of 65 767 individuals, caught with gillnets selective to all fish and size, were analyzed. Eleven species (8 native and 3 exotic) were recorded during the study. Zooplanktivorous were the most representative species, amounting to 91% of the total number. Also, they were the most characteristic species (36.4%) when feeding habits were considered, followed by carnivore and omnivore (27.3% each), and herbivore (9.1%). Three species groups and three time-associated groups were detected by means of classification methods. Lake Patzcuaro's ichthyofauna is to be considered as a transitional community that has not reached equilibrium, particularly after the introduction of exotic species and local extinction of native species. However, Lake Patzcuaro has the greatest fishery production of all tropical lakes in the Central Plain of Mexico and it is mainly sustained by native species. The differential use of feeding resources by the fish species and the diversity in their habits must be taken into account to design strategies to maintain the fish structure (avoiding new introduction of exotic species) and the fish production in Lake Patzcuaro.     

Colonisation rate and adaptive foraging control the emergence of trophic cascades

Ecological communities are assembled and sustained by colonisation. At the same time, predators make foraging decisions based on the local availabilities of potential resources, which reflects colonisation. We combined field and laboratory experiments with mathematical models to demonstrate that a feedback between these two processes determines emergent patterns in community structure. Namely, our results show that prey colonisation rate determines the strength of trophic cascades – a feature of virtually all ecosystems – by prompting behavioural shifts in adaptively foraging omnivorous fish predators. Communities experiencing higher colonisation rates were characterised by higher invertebrate prey and lower producer biomasses. Consequently, fish functioned as predators when colonisation rate was high, but as herbivores when colonisation rate was low. Human land use is changing habitat connectivity worldwide. A deeper quantitative understanding of how spatial processes modify individual behaviour, and how this scales to the community level, will be required to predict ecosystem responses to these changes.     

基于稳定同位素技术的长湖鱼类营养结构研究

本研究应用碳、氮稳定同位素技术分析了2017年秋季（10月）和2018年春季（3月）湖北省荆州长湖鱼类营养结构特征,构建了δ¹³C和δ¹⁵N稳定同位素双位图,并计算了7个相关的量化指标。结果显示：所采集的16种鱼类,δ¹³C均值范围为（-27.7±0.8）‰~（-24.8±0.1）‰,δ¹⁵N的均值范围为（11.4±0.3）‰~（16.6±0.5）‰。营养生态位分析显示,长湖鱼类以偏肉食性鱼类黄颡鱼的营养级最高（3.28±0.2）,草鱼的营养级最低（1.74±0.1）。稳定同位素的量化指标表明,长湖春季鱼类群落核心生态位空间（standard ellipse area, SEA）、生态位总空间（total area, TA）和基础食物来源（δ¹³C range, CR）均高于秋季,两个季度的营养多样性（centrifugal distance, CD）和鱼类群落的整体密度（mean nearest neighbor distance, MNND）相似,但春季的营养长度（δ¹⁵N range, NR）和鱼类群落营养生态位分布范围（standard deviation of nearest neighbor distance, SDNND）值低于秋季。说明春季长湖的生态位总空间（TA上升）和基础食物来源（CR上升）较为丰富,但因鱼类种类的食性相近,其群落的总食物链降低（NR下降）。该研究可为分析拆围后长湖鱼类群落结构特征积累基础数据,也为长湖的渔业管理策略提供依据。     

Fast response of freshwater consumers to a new trophic resource: Predation on the recently introduced Asian bivalve Limnoperna fortunei in the lower Paraná river,South America

Abstract Comparison of Limnoperna fortunei numbers and biomass in screened (5, 15 and 40 mm) and unscreened cages deployed for 18 months in the lower Paraná delta indicates that predators harvest 26–79% (numbers), or 20–85% (biomass) of the mussel population. Predation impact decreases with mussel size. On average, 6 kg of whole live mussel × m⁻² × year⁻¹ (0.36 g of dry mussel tissue × m⁻² × day⁻¹) were eliminated from the unscreened cages. Cages with 15 and 40 mm screens lost between 1 and 2 kg × m⁻² × year⁻¹. Aquatic mammals, birds, and especially fish, are probably the main consumers of large mussels. Small L. fortunei are most probably eaten by fish and also by several invertebrates, including crustaceans, leeches and gastropods. It is suggested that L. fortunei intercepts a significant fraction of the organic carbon that the Paraná‐Uruguay rivers flush into the ocean, locally boosting numbers of benthophagous animals, deposit feeders and, indirectly, higher level predators. Our results indicate that only 15 years after its first introduction in South America this invasive species is very actively consumed by local predators, but predatory suppression of the mussel seems very unlikely. Comparisons with the effects reported for the zebra mussel (Dreissena polymorpha) in Europe and North America suggest that L. fortunei is consumed more actively and that its negative impact on the local fauna is more restricted. These differences are attributed to the fact that while D. polymorpha feeds chiefly on plankton, a limited resource, L. fortunei feeds on detrital particulate organic matter, whose supply in these large South American rivers largely exceeds consumption.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

Energy metabolism of Chironomus anthracinus (Diptera: Chironomidae) from the profundal zone of Lake Esrom,Denmark, as a function of body size,temperature and oxygen concentration

The profundal zone of Lake Esrom, Denmark has a dense population of Chironomus anthracinus, which survives 2–4 months of oxygen depletion each summer during stratification. The metabolism of 3rd and 4th instar larvae was examined in regard to variation in biomass and temperature. Respiration at air saturation was described by a curvilinear multiple regression relating oxygen consumption to individual AFDW and temperature. At 10 °C and varying oxygen regimes the O₂ consumption and CO₂ production of 4th instar larvae were almost unaltered from saturation to about 3 mg O₂ l^–1, but decreased steeply below this level. The respiratory quotient increased from 0.82 at saturation to about 3.4 at oxygen concentrations near 0.5 mg O₂ l^–1. This implied a shift from aerobic to partially anaerobic metabolism. At 0.5 mg O₂ l^–1 the total energy production equalled 20% of the rate at saturation of which more than one third was accounted for by anaerobic degradation of glycogen. This corresponded to a daily loss of 12 µg mg AFDW^–1 or approximately 5% of the body reserves. At unchanged metabolic rate the glycogen store would last three weeks, but long term oxygen deficiency causes a further suppression of the energy metabolism in C. anthracinus.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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The role of zoobenthos in energy flow in deep,oligotrophic Lake Thingvallavatn,Iceland

Macrozoobenthos in Thingvallavatn is dominated by 42 taxa. The vertical distribution delimits 5 communities: (1) the surf zone community from 0–2 m, (2) the upper stony littoral community from 2–6 m, (3) the lower stony littoral community from 6–10 m, (4) the Nitella zone community from 10–20 m, and (5) the profundal zone community from 20–114 m. Total mean lakewide production was 78 kJ m^–2 yr^–1. Herbivores, detritivores, and carnivores contributed 59%, 38% and 3%, respectively. Respiration and ingestion were estimated according to the literature. Net production efficiency averaged 0.50. Ingestion was dominated by herbivores in the littoral zones (46–81%), while detritivores made up 93% in the profundal zone. Total zoobenthic production averaged 6% of estimated available food with a range from 10–11% in the three upper littoral zones to only 2% in the Nitella zone. The profundal fauna converted 6% of the estimated sedimentation of organic matter to secondary production. On a lakewide basis the zoobenthis utilized one third of the estimated potential food resources. Zoobenthic production made up 32% of total secondary production.     

Ecological effects of sex differ with trophic positions in a simple food web

Sexual differences in parental investment, predation pressure, and foraging efforts are common in nature and affect the trophic flow in food webs. Specifically, the sexual differences in predator and prey behavior change in trophic inflow and outflow, respectively, while those in parental investment alter the reproductive allocation of acquired resources in the population. Consequently, these factors may play an important role in determining the system structure and persistence. However, few studies have examined how sexual differences in trophic flow affect food web dynamics. In this study, I show the ecological role of sex by explicitly incorporating sexual differences in trophic flow into a three‐species food web model. The results demonstrated that the ecological waste of males, that is, the amount of trophic inflow into males with less parental investment, plays an important role in system persistence and structure. In particular, the synergy between sexual differences in parental investment and trophic inflows and outflows is important in determining web persistence: Significant impacts of male‐biased trophic flows require the condition of anisogamy. In addition, the dynamic effects of the ecological waste of males differ with trophic level: The coexistence of a food web occurs more frequently with biased inflows into predator males, but occurs less frequently with biased inflows into consumer males. The model analysis indicates that investigating the pattern of sexual differences among trophic positions can enrich our understanding of food web persistence and structure in the real world.     

Tracing energy flow in stream food webs using stable isotopes of hydrogen

1. Use of the natural ratios of carbon and nitrogen stable isotopes as tracers of trophic interactions has some clear advantages over alternative methods for food web analyses, yet is limited to situations where organic materials of interest have adequate isotopic separation between potential sources. This constrains the use of natural abundance stable isotope approaches to a subset of ecosystems with biogeochemical conditions favourable to source separation. 2. Recent studies suggest that stable hydrogen isotopes (δD) could provide a robust tracer to distinguish contributions of aquatic and terrestrial production in food webs, but variation in δD of consumers and their organic food sources are poorly known. To explore the utility of the stable hydrogen isotope approach, we examined variation in δD in stream food webs in a forested catchment where variation in δ¹³C has been described previously. 3. Although algal δD varied by taxa and, to a small degree, between sites, we found consistent and clear separation (by an average of 67‰) from terrestrial carbon sources. Environmental conditions known to affect algal δ¹³C, such as water velocity and stream productivity did not greatly influence algal δD, and there was no evidence of seasonal variation. In contrast, algal δ¹³C was strongly affected by environmental factors both within and across sites, was seasonally variable at all sites, and partially overlapped with terrestrial δ¹³C in all streams with catchment areas larger than 10 km². 4. While knowledge of isotopic exchange with water and trophic fractionation of δD for aquatic consumers is limited, consistent source separation in streams suggests that δD may provide a complementary food web tracer to δ¹³C in aquatic food webs. Lack of significant seasonal or spatial variation in δD is a distinct advantage over δ¹³C for applications in many aquatic ecosystems.     

Size‐based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn,Iceland

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