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。     

应用稳定同位素技术研究崂山湾夏季鱼类群落的摄食生态

崂山湾是黄海沿岸众多海湾之一,也是当前增殖放流的重要水域,研究该区域鱼类群落结构及其食物网结构,有着非常重要的生态意义和实用价值。本研究根据2014年6月和8月在崂山湾海域进行的底拖网调查,分析该海域夏季鱼类群落的优势种组成,并通过测定稳定同位素对其摄食生态进行研究。结果表明,崂山湾夏季鱼类群落的优势鱼种共有10种,包括六丝矛尾鳅虎鱼、中华栉孔鳅虎鱼、红狼牙鳅虎鱼、皮氏叫姑鱼、白姑鱼、小黄鱼、绿鳍鱼、短吻红舌鳎、斑鲨和赤鼻棱鳀;夏季崂山湾不同月份鱼类群落的优势种组成差异较大。各优势鱼种间δ13C值存在显著差异,结合胃含物分析结果表明6月鱼类群落食物的主要来源均为底层饵料,而8月鱼类群落的食物来源则更广泛。崂山湾夏季2个月鱼类群落的优势鱼种间的δ15N值均存在显著差异,最大分别相差4.61‰和3.70‰。崂山湾6月鱼类群落的优势种包括了中营养级鱼类和高营养级鱼类,鱼类群落的平均营养级为3.90,主要是底栖动物食性鱼类;而8月鱼类群落的优势种包括了低营养级鱼类和中营养级鱼类,鱼类群落的平均营养级为3.37,包括浮游动物食性和底栖动物食性2种食性类型,以底栖动物食性鱼类为主。线性回归分析表明崂山湾夏季10种优势鱼种除皮氏叫姑鱼、斑鲨、短吻红舌鳎和绿鳍鱼的营养级与体长无线性相关外,其余优势鱼种的营养级均与体长呈显著的正相关。这主要由于它们随着体长的增加,摄食高营养级的饵料增加,低营养级的饵料减少,自身的营养级也随之升高。     

基于鱼类食性的白洋淀食物网研究

2009年6月至2010年5月,对白洋淀8个典型采样点进行了调查,分析了白洋淀鱼类的食性、营养级和食物网特征,绘制了鱼类食物网及能量流动途径。结果显示,白洋淀现有鱼类25种,其食性类型分为草食性、浮游生物食性、底栖生物食性、游泳动物食性、浮游生物+底栖生物、底栖生物和游泳动物食性共6种;白洋淀的鱼类营养级可分为草食性、杂食性、低级肉食性、中级肉食性和高级肉食性共5种。与历史资料相比,白洋淀鱼类食物网趋于简单,中级肉食性鱼类匮乏,能量流动出现越级现象,食物网结构不完整,说明其渔业资源遭受严重破坏。     

渤海鱼类的食物关系

本研究通过对2010~2011年渤海大面调查所获得的渔获物进行胃含物分析,了解当前渤海鱼类的食物关系及其变化。结果显示,渤海生态系统的27种鱼类有12种低营养级鱼类、12种中营养级鱼类和3种高营养级鱼类,包括了杂食性鱼类、浮游动物食性鱼类、底栖动物食性鱼类、混合动物食性鱼类和鱼食性鱼类,各鱼种营养级较20世纪90年代变化不大。3种高营养级鱼类均为鱼食性鱼类,饵料生境宽度值均很低,属于狭食性鱼类;矛尾虾虎鱼(Chaeturichthys stigmatias)和小黄鱼(Pseudosciaena polyactis)是渤海饵料生境宽度最大的2种鱼,同时也是当前渤海生态系统食物网中最重要的饵料种类,其广食性有利于食物网各营养层次的物质、能量流动。当前渤海食物网中浮游食物链削弱,主要食物链转变为“植物、有机碎屑→鼓虾→鱼类”和“底栖动物→虾虎鱼、小黄鱼→大型经济鱼类”。     

莱州湾鱼类群落的营养结构及其变化

2011年5月~2012年4月期间对莱州湾进行了9次底拖网调查,对采集的20种鱼类4 854个胃含物样品进行了分析。结果表明,莱州湾鱼类群落包括浮游动物食性、底栖动物食性、鱼食性、杂食性和广食性5种食性类型,其中底栖动物食性鱼类是莱州湾各月份鱼类群落的重要食性类型,钩虾和双壳类等底栖动物饵料在莱州湾生态系统的食物关系中起着关键的作用。秋末和初春的莱州湾鱼类群落以底栖动物食性鱼类为主,食性类型单一;随着水温升高,鱼类群落的食性类型逐渐变得丰富。分析表明,莱州湾海域鱼类群落的营养结构有利于增殖放流品种后期的生长存活,建议跟踪调查增殖放流点周边海域的食物关系和饵料基础,同时结合多学科调查,选择最佳的增殖放流地点和时间,才能切实有效地保证增殖放流的效果。     

渤海鱼类的食物关系

本研究通过对2010~2011年渤海大面调查所获得的渔获物进行胃含物分析,了解当前渤海鱼类的食物关系及其变化。结果显示,渤海生态系统的27种鱼类有12种低营养级鱼类、12种中营养级鱼类和3种高营养级鱼类,包括了杂食性鱼类、浮游动物食性鱼类、底栖动物食性鱼类、混合动物食性鱼类和鱼食性鱼类,各鱼种营养级较20世纪90年代变化不大。3种高营养级鱼类均为鱼食性鱼类,饵料生境宽度值均很低,属于狭食性鱼类;矛尾虾虎鱼(Chaeturichthys stigmatias)和小黄鱼(Pseudosciaena polyactis)是渤海饵料生境宽度最大的2种鱼,同时也是当前渤海生态系统食物网中最重要的饵料种类,其广食性有利于食物网各营养层次的物质、能量流动。当前渤海食物网中浮游食物链削弱,主要食物链转变为"植物、有机碎屑→鼓虾→鱼类"和"底栖动物→虾虎鱼、小黄鱼→大型经济鱼类"。     

傀儡湖鱼类群落结构及其食物组成研究

为了分析傀儡湖鱼类对水环境的影响,于2010年1-12月研究了傀儡湖的渔获量、组成比例和主要鱼类食物组成。结果表明,傀儡湖渔获物中,发现有9科、26种鱼类,其中鲤科鱼类最多,有18种,占鱼类种数的69.23%;鱼类组成中,鲢和鳙为优势种,分别占总渔获量的（20.82±8.06）%和（41.58±19.24）%;主要鱼类...     

基于生态网络结构的浙江南部近海鱼类群落关键种识别

对2016-2019年浙江南部近海底拖网采集的主要鱼类进行食性分析,并结合前期摄食习性资料,以种间营养关系为基础构建浙江南部近海鱼类群落的食物网拓扑,以拓扑网络指标结合KPP运算确定该海域的鱼类群落关键种.结果表明:(1)对浙江南部近海的34种主要鱼类开展胃含物分析,共形成营养连接数276对.(2)拓扑食物网中带鱼(T...     

基于鱼类食性的白洋淀食物网研究

2009年6月至2010年5月,对白洋淀8个典型采样点进行了调查,分析了白洋淀鱼类的食性、营养级和食物网特征,绘制了鱼类食物网及能量流动途径。结果显示,白洋淀现有鱼类25种,其食性类型分为草食性、浮游生物食性、底栖生物食性、游泳动物食性、浮游生物＋底栖生物、底栖生物和游泳动物食性共6种;白洋淀的鱼类营养级可分为草食性、...     

基于稳定同位素技术的阳澄湖鱼类群落营养结构研究

为探究阳澄湖水域鱼类群落的营养结构特征, 于 2017 年夏季(7 月)、秋季(10 月)、2018 年春季(4 月)对该水域进行了渔业资源调查, 并利用稳定同位素技术对鱼类群落的碳(δ¹³C)、氮(δ¹⁵N)值特征及其营养结构进行了分析。 研究结果显示, 阳澄湖水域共采集到鱼类 42 种, 隶属于 5 目 10 科 33 属。鱼类的 δ¹³C、δ¹⁵N 和营养级均值分别为 (–24.85±1.77)‰、(15.34±2.19)‰和 3.06±0.64。其中, δ¹⁵N 值与营养级均存在极显著的鱼类食性和季节性差异(P< 0.01), 而 δ¹³C 值在鱼类食性和季节性上未出现显著差异(P>0.05)。阳澄湖鱼类群落的营养结构存在季节差异, 鱼类群落的营养层次(NR)、基础食物来源(CR)、生态位总空间(TA)、群落的整体密度(NND)、聚集均匀程度(SDNND) 和核心生态位空间(SEAc)从 2017 年夏季至秋季再到 2018 年春季, 呈逐渐上升的趋势; 表明 2018 年春季食物源多样性高, 营养冗余程度低。本研究初步构建了阳澄湖鱼类连续营养谱, 丰富了对阳澄湖鱼类群落营养结构方面的研究, 以期为湖泊生态管理和渔业可持续发展提供参考。     

Using a topological approach to identify keystone species of fish in eutrophic lake ecosystems: A case of Zhushan Bay,Taihu Lake

Eutrophication is a global aquatic environmental problem. Traditional biomanipulation targeting keystone species has been effective for restoring eutrophic lakes. In this study, we conducted monthly fishery assessments and identified keystone species in Zhushan Bay of Taihu Lake, the most representative lake in the middle and lower reaches of the Yangtze River. We built the topological structure of Zhushan Bay's food web based on feeding relationships and calculated network indices of fish groups in the food web using network analysis, combined with Key Player Problem operation. Culter mongolicus, Culter alburnus, Protosalanx hyalocranius and Salangichthys tangkahkeii were the keystone species in Zhushan Bay's food web. Culter mongolicus and Culter alburnus had the greatest impact on the food web after removal. Protosalanx hyalocranius had the fastest speed of transmitting information. Salangichthys tangkahkeii was the key baitfish with the strongest ability to determine structure of species in higher trophic levels. Our study provided a reference for scientific management of shallow lake ecosystems and biological solutions for restoration of eutrophic lakes.     

Contrasting changes in freshwater fish assemblages and food webs follow modification of tropical waterways

Fresh waters are increasingly threatened by flow modification. Knowledge about the impacts of flow modification is incomplete, especially in the tropics where ecological studies are only starting to emerge in recent years. Using presence/absence data dated approximately four decades apart (~1966 to ~2010) from 10 tropical rivers, we assessed the changes in freshwater fish assemblage and food web after flow modification. The sites were surveyed with methods best suited to habitat conditions (e.g., tray/push netting for low‐order forest streams, visual surveys for canalised rivers and net casting for impounded rivers). With the presence/absence data, we derived and compared six measures of fish assemblage and food web structure: species richness, proportion of native species, overall functional diversity, native functional diversity, food web complexity and maximum trophic level. We found that changes in community assemblage and food web structure were not generalisable across modification regimes. In canalised sites, species richness and maximum trophic levels were lower in the second time period while the opposite was true for impounded sites. However, proportion of native species was consistently lower in the second time period across modification regimes. Changes in fish assemblages and food webs appear to be driven by species turnover. We recorded 79 cases of site‐specific extirpation and 117 cases of site‐specific establishment. Our data further suggest that turnover in assemblage is again contingent on flow‐modification regime. While the process was stochastic in canalised rivers, benthopelagic species were more likely to be extirpated from impounded rivers where species lost were replaced by predominantly alien fish taxa.     

Big floods,big knowledge gap: Food web dynamics in a variable river system

While flow regime is widely recognised as an over‐arching feature structuring aquatic ecosystems, the influence of flood events on feeding habits in fish assemblages inhabiting variable environments remains largely unknown. In this study, we investigated how diets of a fish assemblage changed in response to fluctuations in hydrology in a highly variable, wet‐dry tropical Australian river. We compared dietary habits, trophic guilds and intra‐ and interspecific dietary overlap and diet breadth across different seasons. Wet‐season floods acted as major geomorphic and ecological perturbations that exerted substantial effects on the trophic dynamics of the terapontid assemblage, particularly through the removal of in‐stream plant production and detritus. Forced by major shifts in food source availability due to seasonal flooding, fishes responded to these perturbations by marked changes in diet, with significant guild switching following wet‐season floods. Rather than the predictable, gradual, changes in consumer–resource interactions associated with the gentler seasonal flood pulse evident in other tropical river systems, study results instead emphasised rapid and profound changes to species' diets and trophic interactions following abrupt flood pulses. These sudden shifts in food sources may also explain some of the lack of clarity in discrimination of specific carbon sources or species' trophic levels evident in isotopic food web studies from the region. Results indicate that the effects of flood pulses on assemblage trophic interactions may differ among ecosystem types and across the broad potential scale of hydrological disturbance mediated by flood events.     

Resource use of crucian carp along a lake productivity gradient is related to body size,predation risk,and resource competition

Generalist fish species can feed on a wide resource spectrum and across trophic levels depending on resource availability and trophic interactions. Crucian carp (Carassius carassius) represents a good candidate species to investigate variation in the trophic ecology of generalist fish as it can be found in highly variable fish communities and its resource use is well documented. In this study, we explored the trophic ecology of crucian carp at the individual and population levels using stable isotope and gut content analysis. We tested if trophic resource use varied according to lake productivity, predation risk, intra- and interspecific competition, or individual fish size. We found that crucian carp resource preference was highly variable among and within lakes. In predator-free lakes, small crucian carp occurred in high densities, showed increased interindividual specialisation, and relied mainly on pelagic zooplankton. In presence of predators, large crucian carp occurred in low densities and included greater proportions of benthic macroinvertebrates in their diet. This shift in resource use was further favoured in productive, shallow lakes where littoral prey were probably abundant. Resource partitioning was an important factor determining crucian carp niche use, as fish had higher trophic position in absence of other cyprinids. Crucian carp showed highly dynamic resource use and food preferences in response to variable environmental conditions. Overlooking complex diet preferences of generalist fish may lead to an oversimplification of freshwater community dynamics.     

Impaired littoral energy pathways cause a shift to pelagic resources by fish in recovering lake food webs

Fish communities may increase in biomass and productivity due to energy subsidies from the littoral invertebrate community. In lakes recovering from acidification and metal contamination, such as those in Sudbury, Ontario, Canada, impaired benthic invertebrate communities (i.e., low diversity with higher abundance of small‐bodied taxa) allowed a critical test of the role of these littoral pathways on fish diet. We compared fish abundance, diversity, diet and biomass in eight recovering and eight reference lakes and related availability of the main littoral and pelagic invertebrate groups to fish diet regime using stable isotope analysis. A Bayesian mixing model (MixSIR) was used to estimate diet likelihood, and convex hull analysis was used to estimate trophic niche space of fish communities. Fish biomass did not differ between impaired and reference lakes despite substantial differences in potential diet. Fish depended strongly on littoral benthos in the reference lakes but consumed more pelagic food in the impaired lakes. The trophic niche of the focal, most common fish species (i.e., yellow perch, smallmouth bass, pumpkinseed and brown bullhead) was larger in the impaired lakes. We attributed these differences to low diversity at the highest trophic levels of fish communities in the impaired lakes as well as to depauperate benthic invertebrate communities. In contrast to the food webs of most temperate lakes, fish in impaired lakes preyed less on littoral invertebrates yet still managed to maintain a reference lake level of biomass standing crop by relying more on pelagic resources – macro zooplankton such as Chaoborus.     

Effects of fish species richness and assemblage composition on stream ecosystem function

Abstract –  Biodiversity is declining across aquatic ecosystems because of biological invasions and species extinctions. Because fishes have pervasive effects on ecosystems through species-specific food web interactions, alterations to species and functional richness, and composition of natural assemblages could have negative effects on aquatic ecosystem function. In this study, I tested the effects of fish species and functional richness, and assemblage composition on primary production (PPR), benthic invertebrate density, and benthic particulate organic matter (BPOM) in a 42-day experiment in artificial stream mesocosms. I found that fish species richness and assemblage composition were important predictors of PPR in stream mesocosms. However, the effect of species richness on PPR increased with time, suggesting that richness-related effects might strengthen as the magnitude of community-level interactions increases in ecosystems. There was no effect of fish species or functional richness or assemblage composition on benthic invertebrate densities or BPOM. These data provide additional support that fishes can be important regulators of ecosystem function in aquatic systems, and suggest that positive effects of fishes on ecosystems can be strengthened by increased species richness and composition of the assemblage. This study broadens the applicability of the biodiversity ecosystem-function literature to a new suite of taxa, supporting the overall hypothesis that ecosystem functions and services are likely to decline in response to species extinctions.     

Trophic ecology of brown trout (Salmo trutta L.) in subarctic lakes

In subarctic lake systems, fish species like brown trout are often important predators, and their niche performance is a key characteristic for understanding trophic interactions and food web functioning at upper trophic levels. Here, we studied summer habitat use and stomach contents of brown trout under both allopatric and sympatric conditions in six subarctic lakes to reveal its trophic role, and population‐ and individual‐level niche plasticity. In allopatry, brown trout mainly used the littoral habitat, but also less commonly used the pelagic zone. In sympatry with stickleback, there was always a considerable habitat overlap between the two species. In contrast, sympatric populations of brown trout and Arctic charr generally revealed a distinct habitat segregation. In the sympatric systems, in general, there was a distinct resource partitioning between the trout and charr, whereas the observed diet overlap between trout and stickleback was much larger. Trout modified their individual dietary specialisation between the littoral and pelagic zone, always being lower in the pelagic. Piscivorous behaviour of trout was only found in sympatric systems, possibly contributing to a competitive advantage of trout over charr and stickleback. Hence, the trophic level of trout was strongly related to the fish community composition, with a higher trophic level in sympatric systems where piscivorous behaviour was frequent. These changes in the trophic level of trout linked with the observed food resource partitioning might be an important mechanism in the ecosystem functioning of subarctic lakes to allow coexistence among sympatric‐living fish species.     

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.     

Effects of littoral habitat complexity and sunfish composition on fish production

Carey MP, Maloney KO, Chipps SR, Wahl DH. Effects of littoral habitat complexity and sunfish composition on fish production. Ecology of Freshwater Fish 2010: 19: 466–476. © 2010 John Wiley & Sons A/S Abstract – Habitat complexity is a key driver of food web dynamics because physical structure dictates resource availability to a community. Changes in fish diversity can also alter trophic interactions and energy pathways in food webs. Few studies have examined the direct, indirect, and interactive effects of biodiversity and habitat complexity on fish production. We explored the effects of habitat complexity (simulated vegetation), sunfish diversity (intra‐ vs. inter‐specific sunfish), and their interaction using a mesocosm experiment. Total fish production was examined across two levels of habitat complexity (low: 161 strands m^?2 and high: 714 strands m^?2) and two sunfish diversity treatments: bluegill only (Lepomis macrochirus) and bluegill, redear sunfish (Lepomis microlophus), and green sunfish (Lepomis cyanellus) combination. We also measured changes in total phosphorus, phytoplankton, periphyton, and invertebrates to explain patterns in fish production. Bluegill and total fish production were unaffected by the sunfish treatments. Habitat complexity had a large influence on food web structure by shifting primary productivity from pelagic to a more littoral pathway in the high habitat treatments. Periphyton was higher with dense vegetation, leading to reductions in total phosphorus, phytoplankton, cladoceran abundance and fish biomass. In tanks with low vegetation, bluegill exhibited increased growth. Habitat complexity can alter energy flow through food webs ultimately influencing higher trophic levels. The lack of an effect of sunfish diversity on fish production does not imply that conserving biodiversity is unimportant; rather, we suggest that understanding the context in which biodiversity is important to food web dynamics is critical to conservation planning.