Generalist feeding strategies in Arctic freshwater fish: A mechanism for dealing with extreme environments

Generalist feeding strategies are favoured in stressful or variable environments where flexibility in ecological traits is beneficial. Species that feed across multiple habitat types and trophic levels may impart stability on food webs through the use of readily available, alternative energy pools. In lakes, generalist fish species may take advantage of spatially and temporally variable prey by consuming both benthic and pelagic prey to meet their energy demands. Using stomach content and stable isotope analyses, we examined the feeding habits of fish species in Alaska's Arctic Coastal Plain (ACP) lakes to determine the prevalence of generalist feeding strategies as a mechanism for persistence in extreme environments (e.g. low productivity, extreme cold and short growing season). Generalist and flexible feeding strategies were evident in five common fish species. Fish fed on benthic and pelagic (or nektonic) prey and across trophic levels. Three species were clearly omnivorous, feeding on fish and their shared invertebrate prey. Dietary differences based on stomach content analysis often exceeded 70%, and overlap in dietary niches based on shared isotopic space varied from zero to 40%. Metrics of community‐wide trophic structure varied with the number and identity of species involved and on the dietary overlap and niche size of individual fishes. Accumulation of energy from shared carbon sources by Arctic fishes creates redundancy in food webs, increasing likely resistance to perturbations or stochastic events. Therefore, the generalist and omnivorous feeding strategies employed by ACP fish may maintain energy flow and food web stability in extreme environments.     

Differences in stable isotope ratios of Dall’s porpoises (Phocoenoides dalli) between coastal and oceanic areas of the North Pacific

Carbon and nitrogen stable isotope ratios in the muscle of Dall’s porpoises were measured. Samples were collected from the catches of the hand harpoon fishery, incidental catches of drift net, and scientific research on the use of drift nets. Samples were from the North Pacific, Sea of Japan, Sea of Okhotsk, and Bering Sea. Although no variation in δ¹⁵N was observed, δ¹³C was significantly different between population groups near Japan and the oceanic North Pacific and Bering Sea. The difference may be due not only to local variation in prey species, but also to an overall difference in carbon stable isotope ratios that originate from coastal benthic or oceanic pelagic‐based food webs. We differentiated Dall’s porpoise population groups from both areas using carbon stable isotope ratios with an error rate of <5%. Although further study is needed, our results suggest that carbon stable isotope ratios could possibly be an indicator of whether a Dall’s porpoise belongs to a coastal benthic or oceanic pelagic food web.     

A comparison of changes in stable isotope ratios in the epidermal mucus and muscle tissue of slow‐growing adult catfish

Although stable isotope analysis is a powerful tool for determining diet, migration patterns and the structure of food webs in aquatic systems, the slow response of isotopic ratios in the widely used muscle tissue often hampers this approach, particularly in slow‐growing or adult fishes. We conducted a diet‐switch experiment to compare the changes in the stable carbon isotope ratio (δ¹³C values) in the epidermal mucus and muscle tissue of five‐year‐old catfish (Silurus asotus). The isotope ratios in noninvasively sampled mucus changed more rapidly than those in the muscle tissue. As isotopic change in mucus was relatively rapid, this technique can be used over a finer timescale than traditional isotopic analyses using the muscle tissue. The isotopic change half‐life in our experimental condition was 200 days, which would not be short enough for some research purposes. Examining mucus along with the muscle would enable food habits of slow‐growing fishes to be determined over different timescales. However, the rate of isotopic change in the mucus was negatively affected by the size of fish and was slower than the previously reported rates in juvenile steelhead. These findings suggest that mucus turnover rates need to be determined prior to the field data interpretation.     

Community and trophic structures of abalone <Emphasis Type="Italic">Haliotis diversicolor</Emphasis> habitat in Sagami Bay,Japan

Abstract:   The community and trophic structures of the subtidal habitat of the abalone Haliotis diversicolor were examined in Nagai, Sagami Bay, Japan. Conventional community indices showed no significant differences among three sampling stations. The overall continuum of stable isotope ratios was structured into three different trophic linkages: (i) brown algae-dependent benthic food chain; (ii) red algae-dependent benthic food chain; and (iii) planktonic food chain. Brown algae and red algae likely play different roles with respect to carbon sources in the habitat. Conventional fractionation values indicated that the abalone H. diversicolor (δ¹³C = −12.4 ± 1.0‰, δ¹⁵N = 9.3 ± 0.5‰) feeds on the lamina of Undaria pinnatifida during juvenile and adult stages (8.0–65.6 mm shell length). Stable isotope signatures suggested that the juveniles of other abalone species as well as some amphipods and a sea cucumber Holothuria decorata are competitors of H. diversicolor , whereas some Muricidae gastropods such as Thais bronni and Ergalatax contractus are predators. The isotopic differences among macroalgal species and the subsequent transfer to consumers indicate that stable isotopic analysis is an effective means of studying food webs in an open rocky shore community with little influence from external primary production such as terrestrial vegetation.     

水生食物网研究方法的发展和应用

水生生物在食物网中的相互关系及其变化是生态学研究的重要领域之一,其实质是研究生物之间的捕食与被捕食关系。传统的水生食物网研究方法是基于形态学观察的胃含物分析,具有较大的局限性。随着科学技术的发展,稳定同位素技术、特征脂肪酸组成分析和DNA条形码等分子技术的引入有效的弥补了胃含物分析的不足,并可以更深入的了解水生生物的摄食信息,因此得到了广泛的应用。本文采用文献计量法分析了水生食物网研究方法的发展动态,分别介绍了稳定同位素技术、特征脂肪酸组成分析和DNA条形码技术与胃含物结合应用的研究进展,着重归纳总结了其在水生生物食性研究中的应用现状及发展前景,详细地指出了4种水生食物网研究方法的优势和局限性、适用范畴和对实验样品的需求。胃含物分析作为传统食物网研究方法未来仍是不可缺少的一部分,其向我们传递了最为直接的摄食信息。在此基础上,稳定同位素技术、特征脂肪酸组成分析和DNA条形码等分子技术作为胃含物分析方法的补充手段,将更有利于重建复杂的水生食物网结构,进而为渔业管理提供必要的帮助。     

Plankton composition and biomass development: a seasonal study of a semi‐intensive outdoor system for rearing of turbot

Plankton food web dynamics were studied during a complete production season in a semi‐intensive land‐based facility for rearing of turbot (Scophthalmus maximus) larvae. The production season was divided into three production cycles of 3–5 weeks. Phytoplankton biomass (using chlorophyll a as biomass proxy) peaked in each production cycle. However, the maximum biomass decreased from spring (18 μg chlorophyll a L^?1) to fall (ca. 7 μg chlorophyll a L^?1), simultaneous with a decline in the concentration of dissolved nitrogen in the inoculating water. During the three production cycles, we observed decreasing copepod nauplii concentration from spring to fall in the rearing tanks. The decreasing nitrogen gave increasing carbon to chlorophyll a ratio in the seston (from 23 in spring to 73 in fall). The pool of free amino acids in seston was constant. We suggest that the decreasing nitrogen input in the inoculating water reduces the quality of the phytoplankton and thus the growth potential of the prey for fish larvae, copepods.     

Seasonal and spatial variations of the food web structure in a shallow eutrophic lake assessed by stable isotope analysis

Stable isotope analysis was used to investigate seasonal and spatial variations of the food web structure in a large eutrophic lake ecosystem (Lake Taihu, China). Basal food sources, invertebrates and fish were sampled in two lake regions with different environmental conditions and spatial variations in the isotopic composition of lake food webs were found. Overall, more depleted δ¹³C and enriched δ¹⁵N isotope values of organic matter sources and consumers were found in the phytoplankton-dominated lake region than in the macrophyte-dominated region. Wide seasonal variations in the isotopic ratios were also observed in the lake biota, with a general trend towards enriched δ¹³C and δ¹⁵N values in summer and depleted values in winter. This pattern could be explained by a combination of environmental (e.g., irradiance and nutrient inputs) and biotic (e.g., availability of food sources and plasticity in prey item choice) features. Results of isotope mass balance suggest that macrophytes provide some trophic support in the macrophyte-dominated area, but in both lake regions it is more likely that aquatic food webs are phytoplankton based rather than macrophyte based under eutrophic conditions.     

Trophic Ecology of Arapaima sp. in a ria lake—river–floodplain transition zone of the Amazon

Neotropical floodplains are usually productive systems that are maintained by the nutrient, detritus and sediment inputs provided by the main river channel flood pulse. Ria lakes represent a special feature and habitat in the Amazonian floodplains, being characterised by a dendritic morphology and dependence on terrestrial inputs provided by an intricate stream network. Our objective was to evaluate the trophic ecology of the arapaima (Arapaima sp.). We combined stomach content analysis with measurements of carbon and nitrogen stable isotope values from dorsal muscle to infer the ontogenetic changes in trophic level and isotopic niche width in floodplain and ria lakes. Arapaima diet was dominated by fish from low trophic positions. While most of the stomachs sampled in the study ria lake were full, empty stomachs predominated in samples taken in the floodplain lakes. These differences indicate that ria lakes may provide better feeding grounds for arapaima during the dry season, presumably because ria lakes are interconnected with a large stream network and the main river channel year round. Nitrogen stable isotope results further indicated an ontogenetic dietary shift in arapaima, with piscivory increasing as a function of length in both environments. Carbon stable isotope analysis indicated that energy sources used by arapaima varied by environment, with arapaima using a greater diversity of food sources in ria lakes than in floodplain lakes. Information about the main carbon sources is useful for fishery management because stakeholders may choose to conserve key vegetal groups to ensure the productivity and diversity of aquatic ecosystems.     

Epilithic biofilm as a key factor for small‐scale river fisheries on Caribbean islands

Abstract Numerous freshwater species are consumed or exploited through artisanal fisheries in the rivers of the islands of Guadeloupe, French West Indies. Autochthonous production of organic matter is limited in these flowing rivers and is mainly represented by scarce filamentous green algae and an abundant epilithic biofilm growing on wet stones in the river bed. Stable isotope analysis was used to quantify the relative importance of biofilm and other riverine allochthonous and autochthonous food sources in the diet of tropical shrimps (Palaemonidae, Atyidae and Xiphocarididae) and fishes (Gobiidae and Eleotridae) consumed by the local people. The epilithic biofilm was exploited by most species, constituted an important source of autochthonous carbon and was an important source of organic matter production at the base of freshwater food webs in Caribbean rivers. Biofilm percentages in the diet reached 32% for molluscs, 85% for atyid shrimps, 29% for xiphocaridid shrimps, 14% for palaemonid shrimps and 13% for fish. Assessment of biofilm in nine rivers showed that blue and red cyanobacteria were quantitatively dominant with a moderately rich diatom flora. These results address the interactions between river biofilm and Caribbean freshwater fauna where trophic links between consumers and their potential resources have poorly been documented.     

基于碳、氮稳定同位素技术的大亚湾紫海胆食性分析

为掌握大亚湾紫海胆(Heliocidaris crassispina)的食性特征,应用碳、氮稳定同位素技术对2015年8月所采集紫海胆样本的稳定同位素特征、营养级和食性特征进行了初步研究。结果表明,大亚湾紫海胆平均δ~(13)C值为-(13.35±1.21)‰,平均δ~(15)N值为(9.14±0.38)‰,平均营养级为2.11±0.14。不同壳径紫海胆之间的碳、氮稳定同位素比值无显著性差异(P0.05)。大亚湾海域紫海胆生活环境周围生物δ~(13)C值分布范围为-20.76‰~-9.93‰,δ~(15)N值分布范围为-0.16‰~14.99‰,营养级范围为1.34~3.77。大亚湾主要生物种类可划分为悬浮物、初级生产者和初级消费者、次级消费者、顶级消费者4个营养组群,其中紫海胆属于次级消费者。8月份调查海域珊瑚稀少,大型海藻密度低且死亡降解形成颗粒有机物(Particulate Organic Matter,POM),陆源POM随降雨大量流入大亚湾,导致紫海胆在8月份摄食偏向碎屑食物链,主要食物来源为POM,平均贡献率为67.3%;其余摄食种类为沉积物(Sediment Organic Matter,SOM)、裂叶马尾藻(Scagassum siliquastrum)、底栖硅藻、浮游动物及浮游植物,平均贡献率分别为9.7%、9.3%、6.7%、3.7%及3.3%。大亚湾紫海胆摄食种类与其栖息地底栖生物存在重叠,具有一定的食物竞争关系。研究表明,分析紫海胆食性特征对了解其所在生态系统中营养级水平具有重要意义。     

Responsiveness of fish mass–abundance relationships and trophic metrics to flood disturbance,stream size,land cover and predator taxa presence in headwater streams

Characterisation of food webs, by summarising energy transfer and trophic relationships, allows more functional measurement of ecosystems and may reveal threats (e.g., land‐cover change) in sensitive environments that are not obvious from conventional biomonitoring. However, typical methods used to achieve this are time‐consuming and expensive. Therefore, we tested the usefulness of fish‐focused food‐web proxies as functional measures, specifically mass–abundance relationships of fish assemblages and stable isotope (SI)‐derived metrics in headwater stream reaches. These metrics have been trialled before for similar use in other settings, but have yielded varying results, and have not been employed in tandem in temperate freshwaters. Sampling reaches (N = 46) were spread across a variety of streams, and the effects of habitat predictors at multiple scales on metrics were assessed using model selection. We found that habitat size positively correlated with food‐chain lengths in streams, possibly because of increased abundance of fish at multiple trophic levels in habitats with more space. Additionally, flood disturbance was negatively associated with fish mass–abundance and carbon range, likely due to the harshness of flood‐prone streams. Riparian land‐cover variables were correlated with multiple metrics, indicating the importance of terrestrial–aquatic linkages. Additionally, variations in all metrics were influenced by the presence of native, predatory longfin eels. Overall, we conclude that mass–abundance relationships and SI‐derived metrics are sensitive to drivers of trophic organisation and likely reflect processes occurring at multiple spatial scales in freshwaters. Thus, these metrics could be an insightful monitoring tool for managers because they reflect functional measures of aquatic ecosystems.     

Trophic structure of a pond community dominated by an invasive alien species: Insights from stomach content and stable isotope analyses

1. Invaders affect native species across multiple trophic levels, influencing the structure and stability of freshwater communities. Based on the ‘trophic position hypothesis’, invaders at the top of the food web are more harmful to native species via direct and indirect effects than trophically analogous native predators are.
2. However, introduced and native predators can coexist, especially when non-native species have no ecological and behavioural similarities with natives, occupy an empty niche, or natives show generalist anti-predator strategies that are effective at the community level.
3. At present, conservation efforts are focused on eradicating invaders; however, their removal may lead to unwanted and unexpected outcomes, especially when invaders are well established and strongly interspersed with natives. This highlights the need to consider invaders in a whole-ecosystem context and to consider the evolutionary history and behavioural ecology of natives and invaders before active management is applied.
4. Here, stomach content and stable isotope analyses were combined to investigate a pond system dominated by invaders in order to understand the effects of the interactions among upper level predators and lower level members of the food web on the whole community structure.
5. Both diet and isotope analyses showed that several invaders contributed to the diet of natives and invaders. A significant isotope overlap was found among upper level predators. However, stomach content analysis suggested that predators reduced the potential competition differentiating the food spectrum by including additional prey in their diet. Both native and non-native upper level predators, by preying on invaders, seem not to exert a strong suppressive effect through predation and competition on native species.
6. This research confirms the importance of studying food webs to identify ecological conditions that forecast the potential for deleterious impacts before management is applied. In cases where invaders cannot be eradicated, management efforts should follow a conciliatory approach promoting the coexistence of native species with invaders.

     

鱼类食性和食物网研究评述

鱼类食性和食物网的研究对于了解鱼类群落的结构与功能具有重要意义。本文主要结合国内外有关鱼类食性和食物网研究的报道，概括性叙述了在进行鱼类的食物组成、食性比较、摄食的时空变化、摄食的选择性、营养级、摄食量、食物网及其营养动力学研究时所使用的方法，并对我国开展此类研究的前景和存在的问题进行了讨论。     

Organic matter derived from kelp supports a large proportion of biomass in temperate rocky reef fish communities: Implications for ecosystem‐based management

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应用稳定同位素技术分析枸杞岛近岸海域褐菖鲉的食性

为获知褐菖鲉的食性,本研究应用碳、氮稳定同位素技术分析了不同季节和发育阶段褐菖鲉的摄食习性,并评估了浮游动物、端足类、虾蟹类和小型鱼类等饵料生物对褐菖鲉的营养贡献。结果显示,枸杞岛近岸海域4个季节褐菖鲉的δ~(13)C值为–18.2‰~–14.7‰,平均值为–15.6‰;δ~(15)N值为3.3‰~9.9‰,平均值为7.9‰;单因素方差分析显示,褐菖鲉的δ~(13)C值和δ~(15)N值在不同季节之间差异都不显著,说明不同季节之间褐菖鲉的食性差异不大。对褐菖鲉全长和碳、氮稳定同位素比值进行线性回归分析,显示其δ~(13)C值和δ~(15)N值随全长变化都极显著,表明不同发育阶段褐菖鲉的食性差异极显著。聚类分析将不同全长的褐菖鲉分为3组,其中0~19 mm为一组,20~79和80~109 mm为一组,110~199 mm为一组,ANOSIM分析3组之间的差异极显著。褐菖鲉的δ~(13)C值与虾蟹类和鱼类等饵料类群δ~(13)C值较接近,应用Iso Source软件分析饵料类群的营养贡献时发现,麦秆虫对褐菖鲉的的营养贡献均值最大,为22.3%,鱼类、虾蟹类等饵料类群的营养贡献均值都超过了12%。枸杞岛近岸海域褐菖鲉的平均营养级为3.18,属于低级肉食性鱼类,主要摄食麦秆虫、鱼类和虾蟹类,是底栖动物食性,而非浮游动物食性。     

阿根廷滑柔鱼雌性个体肌肉和性腺组织能量积累及其生殖投入

利用组织能量密度测定技术和残差指标分析方法,分析了阿根廷滑柔鱼雌性个体肌肉组织和性腺组织的能量积累及其分配规律。结果显示,足腕、胴体、卵巢、缠卵腺和输卵管复合体等组织的能量密度平均值分别为(22.75±0.60)、(23.42±0.68)、(25.15±2.08)、(21.49±1.46)和(25.12±3.02)k J/g。不同性腺成熟度等级,足腕、胴体和缠卵腺等组织的能量密度均不存在显著性差异,卵巢和输卵管复合体等组织的能量密度则差异显著。足腕、胴体等肌肉组织能量在性腺成熟度III期时达到最大值,卵巢、缠卵腺和输卵管复合体等性腺组织在I~III期时的能量积累缓慢,随后能量积累增长迅速,在生理性成熟后期(V期)或功能性成熟期(VI期)时达到最大值。随着性腺生长发育,肌肉组织能量占比逐渐下降,性腺组织能量占比逐渐增加。在性腺成熟度III~V期时,雌性个体肌肉组织能量部分转化用于性腺组织的生长发育,但这种生殖投入方式不影响肌肉组织的完整性。研究表明,阿根廷滑柔鱼生殖投入以外源性为主,但发育过程中肌肉组织存储能量部分转化以满足性腺发育对能量的需求。     

Planktivorous vendace (Coregonus albula) utilise algae‐derived fatty acids for biomass increase and lipid deposition

Spatial and seasonal variation in the abundance and composition of phytoplankton and other basal resources (bacteria, terrestrial detritus) influence the availability of essential polyunsaturated fatty acids (PUFA) for upper trophic levels in lake food webs. We studied accumulation, composition and content of fatty acids in planktivorous vendace (Coregonus albula) muscle tissue during the summer feeding period (May to late September) in six boreal lakes. Vendace muscle fatty acid content increased from ~17.7 mg/g DW to ~28.1 mg/g DW from spring to summer, corresponding to a ~59% increase in total fatty acids. PUFA accounted for 45%–65% of all fatty acids, and the most abundant fatty acid was docosahexaenoic acid (DHA, 22:6n‐3). The DHA content remained relatively constant throughout the sampling period (~7–8 mg/g DW), but the proportion of DHA decreased from ~40% in spring to ~30% in summer and autumn. The decrease in DHA proportion was due to accumulation of other fatty acids, primarily C₁₈ PUFA (predominantly synthetised by algae) and monounsaturated fatty acids. A similar fatty acid accumulation pattern was observed in lipid storing copepods, an important prey for planktivorous vendace. The abundance of bacterial and terrestrial biomarker fatty acids in vendace muscle and prey items was low (<5% and <0.1% respectively). The increase in weight and the deposition of internal energy stores in vendace were achieved predominantly by algae‐based food chain. This has ecological implications, as vendace requires sufficient lipid stores for reproduction in autumn and survival over winter.     

Stable isotope analysis suggests the existence of multiple populations of streaked spinefoot (<Emphasis Type="Italic">Siganus javus</Emphasis> L.) in Bandon Bay,Southern Thailand

The stock unit used in fisheries resource assessment and management is generally based on the morphological and genetic characteristics of a particular population or species to avert problems caused by the treatment of multiple populations as one stock, which can lead to the overestimation of population sizes and genetic pollution. Furthermore, since the linkage of microhabitats is an important factor affecting the reproduction of marine organisms in coastal areas, an understanding of the food web in each microhabitat is essential to establish sustainable fisheries management practices. We investigated spatial variations in the food sources and feeding habits of immature stage of Siganus javus using genetic population analyses and stable isotope analyses (δ¹³C and δ¹⁵N). These species are commonly harvested by small-scale fisheries, and it inhabits Bandon Bay in the Surat Thani Province of Southern Thailand. Genetic variation within sampling sites was greater than that between sites. The δ¹³C values of S. javus differed between sites, which suggest that the different ecological habitats exhibit different rates and patterns of carbon flow even among sites located in the same bay. Our results suggest that studies combining genetic population analyses and stable isotope analyses are required to confirm the delineation of fine-scale management units intended for the development of coastal fishery resources.     

Brown shrimp (Farfantepenaeus aztecus) density distribution in the Northern Gulf of Mexico: an approach using boosted regression trees

The estuarine‐dependent brown shrimp, Farfantepenaeus aztecus, is a significant commercial fishery and important species in the Gulf of Mexico (GOM) ecosystem as well as being a key component in energy transfer between benthic and pelagic food web systems. Because of the economical and ecological importance of brown shrimp, we developed a spatial population model to identify places of high shrimp density under a set of spatial, environmental and temporal variables in the Northern Gulf of Mexico (NGOM). We used fisheries‐independent data collected by the Southeast Area Monitoring and Assessment Program (SEAMAP) from 1992 to 2007 (summer and fall seasons). The relationship between the predictor variables and shrimp density was modeled using Boosted Regression Trees (BRT). Within the environmental variables included in the model, bottom type and depth of the water column were the most important predictors of shrimp density in the NGOM. Spatial predictions performed using the trained BRT model for summer and fall seasons showed a spatial segregation of shrimp density. During the summer, higher densities were predicted near the Texas and Louisiana coast and during the fall, higher densities were predicted further offshore. The model performed well and allowed successful prediction of brown shrimp hot spots in the NGOM. Model results allow fisheries managers to evaluate the potential impact from fisheries on the resource and to develop future fisheries management strategies, understand the biology of brown shrimp as well as assess the potential impacts of oil spills or climate change.     

Food habits of introduced brown trout and native masu salmon are influenced by seasonal and locational prey availability

A knowledge of food habits is important for evaluating interspecific competition and predation between sympatric species. Data on food availability should be combined with data on food habits in this type of survey. Although food availability differs between habitats or seasons, these differences had never been considered in previous studies. We conducted year-round field surveys throughout a stream to compare the food habits of an introduced salmonid, brown trout Salmo trutta, and a native salmonid, masu salmon Oncorhynchus masou. Masu salmon did not constitute a large proportion of the diet of brown trout and vice versa. Thus, predation will likely not affect the population level of either species. The dietary overlap between brown trout and masu salmon varied depending on the presence of Gammaridae and terrestrial invertebrates; i.e., the intensity of interspecific competition for food resources may differ according to food conditions.