基于SURF指数识别海州湾食物网的关键饵料生物

关键种(keystone species)在生态系统中发挥着不可替代的重要作用,对于群落结构的稳定与演替起着决定性的作用。基于2011年3—12月在海州湾及其邻近海域进行的渔业资源底拖网调查资料以及胃含物分析数据和参考历史文献数据,以物种间的摄食关系作为基础,采用SURF(Supportive Role to Fishery ecosystems)指数识别海州湾食物网中的关键饵料生物。在矩阵中分析每个物种作为饵料生物为捕食者提供的摄食比例及捕食者数量,计算每个物种的关键指标(SURFi)的值,结果表明前五位数值较高的物种是细鳌虾(Leptochela gracilis)、毛虾(Acetessp.)、疣背宽额虾(Latreutes planirostris)、日本鼓虾(Alpheus japonicus)和鳀鱼(Engraulis japonicus),它们是海州湾的关键饵料生物,在整个食物网中起到关键的控制作用,它们的数量波动会对海州湾其他物种产生直接或间接的影响。加强关键饵料生物的保护,对于维持海州湾生态系统的健康和稳定至关重要。     

海州湾5种关键饵料生物的主要捕食者及其捕食压力

为了解海州湾关键饵料生物在食物网中的被捕食压力及自然死亡率波动情况,研究基于2011和2013—2020年在海州湾及其邻近海域进行的渔业资源底拖网调查资料和胃含物分析数据,以物种间的营养联系为基础,选择细螯虾(Leptochela gracilis)、日本鼓虾(Alpheus japonicus)、枪乌贼(Loligo sp.)、小黄鱼(Larimichthys polyactis)和口虾蛄(Oratosquilla oratoria)5种关键饵料生物为研究对象,通过计算捕食压力指数(Predation pressure index, PPI),分析它们在海州湾食物网中的主要捕食者及其捕食压力,并计算了纳入捕食压力指数的自然死亡系数。结果表明,细螯虾的捕食者对其年平均捕食压力指数最高,小黄鱼的捕食者对其年平均捕食压力指数最低。小眼绿鳍鱼(Chelidonichthys kumu)对细螯虾和枪乌贼的年平均捕食压力指数最高,分别为168.89和75.77;(Miichthys miiuy)对日本鼓虾和口虾蛄的年平均捕食压力指数最高,分别为39.41和9.85;海鳗(Muraenesox c...     

基于Delta-GAMMs模型研究海州湾5种关键饵料生物被摄食的影响因素

海洋生态系统的营养动力学是决定渔业生产力的主要因素,也是实施基于生态系统的渔业管理的重要基础。为了评估生物因子和非生物因子对海州湾5种关键饵料生物被摄食的影响,本研究基于2011和2018年秋季在海州湾及其邻近海域进行的渔业资源底拖网调查和胃含物分析数据,构建了Delta-GAMMs模型,选择细螯虾、日本鼓虾、枪乌贼、小黄鱼和口虾蛄5种关键饵料生物作为研究对象,基于出现频率百分比和捕食压力指数筛选其主要捕食者,基于方差膨胀因子和全子集回归筛选影响5种关键饵料生物被摄食的关键因子。结果表明：5种关键饵料生物在捕食者食物中的出现频率范围为8.5%～42.2%,重量百分比范围为4.2%～40.9%。Delta-GAMMs二项模型和正态模型的平均偏差解释率分别为16.1%和23.8%。捕食者体长、捕食者种群密度和底层水温是影响饵料生物-捕食者相互作用的重要因素。其中,捕食者体长是最重要的影响因素,关键饵料生物的被摄食率和被摄食量均随捕食者体长的增加呈上升趋势,关键饵料生物的被摄食率和被摄食量随着捕食者种群密度的增加而下降。此外,底层水温、水深、纬度和底层盐度等环境因素对种间营养关系的影响,因饵...     

群落水平食物网能流季节演替特征

稳定碳、氮同位素比值分析技术是研究生态系统中物质循环与能量流动的有效技术.δ13C可以用来判断食物网中不同生物的能量来源;δ15N主要用于确定生物在食物网中所处的营养位置.通常用δ13C—δ15N图来表征某一特定时间或空间的食物网结构,但是这种方法在比较不同时间和空间食物网结构中功效较差.同时这种定性描述食物网结构也无法满足食物网复杂变化下的假说验证.应用环形统计方法,以太湖梅梁湾鱼类群落为例,定量评价了群落水平食物网能流季节演替特征.结果表明太湖梅梁湾鱼类营养生态位移动的季节特征明显.进一步物种水平分析结果表明,各种鱼类角度和幅度随季节均有变化.Rayleigh检验结果表明,群落中不同种类在秋冬、冬春和夏秋均有显著的一致的方向变化;而春夏期间不显著.Watson-William检验结果表明,群落水平的鱼类营养生态位移动在秋冬和冬春季节转换中有显著差异.引起鱼类群落水平营养生态位在食物网空间中季节性移动的主要因素为可利用资源稳定同位素的季节变化和不同鱼类种群自身的食性季节性转变.同时,由于梅梁湾食物网鱼类群落杂食性水平高,季节性浮游初级生产力成为食物网能量流动的重要驱动作用.因此,在富营养化生态系统中,食物网群落水平营养生态位季节波动也暗示了系统稳定性的下降.定量评价食物网变化有助于认识和理解食物网结构与功能在生态学和生态系统管理等方面的重要.     

浅水湖泊四种渔具捕捞选择性及对鱼类群落影响比较研究

了解不同种类和大小的鱼类对各种捕捞方式的脆弱性, 对于制定最佳的捕捞策略和合理开发鱼类资源至关重要。研究从渔获物组成、捕捞效率和个体大小等方面, 对长江中游典型湖泊牛山湖的四种渔具(网簖、刺网、电捕和鸬鹚)的捕捞选择性进行了比较分析。使用相对重要性指数(IRI)评价渔具捕捞对鱼类物种的选择性, 同时通过计算渔获物的平均营养水平(MTL)和饵料鱼与凶猛性鱼类的重量比, 分析捕捞对鱼类群落营养结构的潜在影响。结果表明, 网簖和电捕捕获的鱼类种数多、体长范围广, 对鱼类资源的破坏性最大, 应当加以限制或废除; 鸬鹚是捕获鳜(尤其是高龄鳜)的一种有效方式, 但对幼鳜和其他幼鱼也有较大损害, 应控制其捕捞强度; 刺网主要捕获几种经济鱼类, 并可平衡捕食者和饵料鱼之间数量关系, 是利用鱼类资源的一种较好方式, 关键在于对刺网网目大小和捕捞强度的限制。建议湖泊捕捞应从单一物种转向多物种综合管理, 并考虑物种之间的营养生态关系, 以更好地保护和可持续利用渔业资源。     

崂山湾人工鱼礁区星康吉鳗摄食生态及食物网结构

根据2015年4月至2017年1月于崂山湾人工鱼礁区地笼网和延绳钓捕获的279尾星康吉鳗样本,从胃含物组成、食性类型、摄食等级、营养生态位和营养级等方面对其摄食生态进行研究,同时结合海区许氏平鲉、大泷六线鱼、斑头鱼、褐菖鲉、花鲈等9种鱼类的胃含物分析结果,构建人工鱼礁区鱼类关键种的简化食物网模型.食性研究结果表明:星康吉鳗共摄食7类30余种饵料,虾类是其最主要的饵料类群,其次为鱼类和头足类,大泷六线鱼、方氏云鳚、鹰爪虾、玉筋鱼和日本鼓虾等是其优势饵料.星康吉鳗的饵料生物组成随肛长和季节发生显著变化.四季均以鱼类和虾类为主,春季胃含物中包括头足类,秋季包括头足类和蟹类,冬季亦有蟹类出现.肛长≤120 mm的星康吉鳗主要摄食鱼卵和鹰爪虾,120~130 mm肛长组主要摄食玉筋鱼和日本鼓虾,肛长>130 mm的星康吉鳗主要摄食大泷六线鱼和方氏云鳚.其摄食强度也随季节和肛长而变化,空胃率的季节性差异显著,平均胃饱满系数的季节性差异不显著,不同肛长组的空胃率和平均胃饱满系数均不存在显著差异.人工鱼礁区简化食物网结构显示:鱼类关键种的营养级均在3级以上,星康吉鳗的营养级为4.636,处于海区食物网的最顶端.虾类、蟹类、端足类和软体动物等是鱼类关键种的主要饵料,甲壳类、方氏云鳚、大泷六线鱼和玉筋鱼是高营养级鱼类花鲈和星康吉鳗的主要饵料.     

浅水湖泊四种渔具捕捞选择性及对鱼类群落影响比较研究（英文）

了解不同种类和大小的鱼类对各种捕捞方式的脆弱性,对于制定最佳的捕捞策略和合理开发鱼类资源至关重要。研究从渔获物组成、捕捞效率和个体大小等方面,对长江中游典型湖泊牛山湖的四种渔具(网簖、刺网、电捕和鸬鹚)的捕捞选择性进行了比较分析。使用相对重要性指数(IRI)评价渔具捕捞对鱼类物种的选择性,同时通过计算渔获物的平均营养水平(MTL)和饵料鱼与凶猛性鱼类的重量比,分析捕捞对鱼类群落营养结构的潜在影响。结果表明,网簖和电捕捕获的鱼类种数多、体长范围广,对鱼类资源的破坏性最大,应当加以限制或废除;鸬鹚是捕获鳜(尤其是高龄鳜)的一种有效方式,但对幼鳜和其他幼鱼也有较大损害,应控制其捕捞强度;刺网主要捕获几种经济鱼类,并可平衡捕食者和饵料鱼之间数量关系,是利用鱼类资源的一种较好方式,关键在于对刺网网目大小和捕捞强度的限制。建议湖泊捕捞应从单一物种转向多物种综合管理,并考虑物种之间的营养生态关系,以更好地保护和可持续利用渔业资源。     

海州湾鱼类群落功能多样性的时空变化

功能多样性是联系物种、生态环境和生态系统功能的基础.从功能多样性的角度研究群落结构,有助于更好地理解生物多样性与生态系统功能之间的关系.本研究根据2011—2017年(除2012年外)春、秋季海州湾渔业资源底拖网调查数据,选取反映鱼类摄食、运动、生态适应性、繁殖行为和种群动态特征的13个功能性状,利用功能丰富度指数、功能均匀度指数、功能离散度指数和群落特征加权平均数指数(CWM),研究了海州湾鱼类群落功能多样性的季节、年际和空间的变化.结果表明: 海州湾鱼类群落功能多样性指数具有显著的季节差异,其中秋季功能丰富度指数显著高于春季,春季功能离散度指数显著高于秋季,鱼类洄游是造成功能多样性指数季节变化的主要原因;CWM分析表明,春季鱼类群落优势种为营养级较高、运动能力较强、生长系数、恢复力和脆弱性较高的冷/暖温性鱼类,秋季则相反,春、秋季鱼类群落均以产浮性卵鱼类为主;鱼类群落功能多样性指数呈现一定的年际变化,其中春、秋季各功能多样性指数在不同年际均呈波动趋势,表明海州湾鱼类群落稳定性较低;鱼类群落功能多样性指数呈现显著的空间差异,其中20 m以深远岸水域功能离散度指数较高.海州湾鱼类群落功能多样性具有明显的时空变化特征,鱼类生态位与其对资源的利用均随季节、年际和空间而变化.     

海州湾鱼类群落平均营养级和大型鱼类指数的变化特征

为探究近年来海州湾鱼类群落的变化特征,本研究根据2011年及2013-2017年春季(5月)和秋季(9-10月)在海州湾及其邻近海域进行的渔业资源底拖网调查,分析了海州湾鱼类群落平均营养级(MTL)和大型鱼类指数(LFI),对海州湾鱼类群落结构特征进行研究.结果表明:海州湾的优势鱼种主要有大泷六线鱼、方氏云鳚、尖海龙、小黄鱼、长蛇鲻等,且优势鱼种季节性变化明显.海州湾鱼类群落的平均营养级存在明显的年际和季节变化,总体上秋季的MTL高于春季,而且秋季MTL变化具有滞后性.LFI计算结果表明,近年来海州湾及其邻近海域大个体鱼类资源量有所减少,鱼类群落结构呈现出明显的小型化趋势.     

三峡水库生态渔业发展策略与关键技术研究分析

针对三峡水库蓄水后水体资源丰富、水生态系统发育尚不完善、支流库湾藻类水华问题较严重、鱼类群落结构有待调控、水生生物资源未有效利用和转化的现状, 阐述了三峡水库生态渔业作为生态系统保护途径和绿色产业的必要性与重要性, 提出了三峡水库生态渔业发展的总体目标与基本原则, 认为三峡水库发展生态渔业应以生态安全保障和水质养护为首要任务, 严格控制外来物种的引种移植, 以土著鱼类自然繁殖保护和捕捞管理为主, 动态调控放流增殖的鱼类种类和数量为辅, 建立以鱼类群落合理配置和食物网结构优化为手段的水库生态系统调控技术体系, 促进高效的物质循环和能量流动, 实现环境保护和渔业增效的双赢。作者围绕渔业放流增殖、野生鱼类资源保护、捕捞管理、局部库区渔业调控、渔业生物控藻、社区渔业协调管理、生态渔业总体规划等方面, 分析了现阶段三峡水库生态渔业的重点研究任务与关键技术,同时建议加强相关生态学理论与方法研究、技术示范和成果应用, 为三峡水库以渔养水、渔-水和谐的综合管理提供决策依据。       

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Fish predation can induce mesohabitat-specific differences in food web structures in small stream ecosystems

Diverse benthic communities in streams include a wide variety of predators with different habitat preferences, e.g. for pools or riffles. We hypothesised that these preferences result in mesohabitat-specific predator community structures with quantitative differences concerning predation intensity by vertebrate and invertebrate predators, importance of intraguild predation, or top–down pressure. This hypothesis was evaluated for a small submontane stream by means of mesohabitat-specific quantification of prey consumption by two benthivorous fish species (Gobio gobio and Barbatula barbatula) and several invertebrate predators. The estimation was based on daily food rations and diet composition of predators and mesohabitat-specific predator biomass. We found clear differences between the two mesohabitat types. Predator food webs were less complex in pools than in riffles. Fish predation was more important than invertebrate predation in pools, and intraguild predation had a higher relative importance in these mesohabitats. These differences were probably caused by the mesohabitat use of G. gobio, the largest top predator, which preferred pools. Consequently, the predator food webs were more similar between the mesohabitats when fish were absent. Top–down pressure on primary consumers by all predators together was lowest in pools without fish, but the effect was not significant. Omnivory (including cannibalism) was intense, but its potentially destabilising effects were probably counterbalanced by mesohabitat connectivity. From the results of our experimental study, we conclude that even in small stream ecosystems, food web structures and predation pathways can differ between mesohabitats and that a mesohabitat-specific consideration will help to explain the variety of top–down effects on benthic communities.     

Spatial dynamics of keystone predation

1. I investigated the effects of dispersal on communities of keystone predators and prey. I obtained two key results. 2. First, a strong trade-off between competitive ability and predator susceptibility allows consumer coexistence over a large resource productivity range, but it also lowers the predator-susceptible superior competitor's abundance and increases its risk of extinction. Thus, unexpectedly, dispersal plays a more important role in coexistence when predator-mediated coexistence is strong rather than weak. The interplay between the trade-off, small population sizes resulting from transient oscillations, and dispersal leads to qualitatively different species distributions depending on the relative mobilities of the consumers and predator. These differences yield comparative predictions that can be tested with data on trade-off strength, dispersal rates, and species distributions across productivity gradients. 3. Second, there is an asymmetry between species in their dispersal effects: the predator-resistant inferior competitor's dispersal has a large effect, but the predator-susceptible superior competitor's dispersal has no effect, on coexistence and species' distributions. The inferior competitor's dispersal also mediates the predator's dispersal effects: the predator's dispersal has no effect when the inferior competitor is immobile, and a large effect when it is mobile. The net outcome of the direct and indirect effects of the inferior competitor's dispersal is a qualitative change in the species' distributions from interspecific segregation to interspecific aggregation. 4. The important point is that differences between species in how they balance resource acquisition and predator avoidance can lead to unexpected differences in their dispersal effects. While consumer coexistence in the absence of dispersal is driven largely by the top predator, consumer coexistence in the presence of dispersal is driven largely by the predator-resistant inferior competitor.     

ORGANIZATION OF PREDATOR-PREY COMMUNITIES AS AN EVOLUTIONARY GAME

We consider a simple predator-prey model of coevolution. By allowing coevolution both within and between trophic levels the model breaks the traditional dichotomy between coevolution among competitors and coevolution between a prey and its predator. By allowing the diversity of prey and predator species to emerge as a property of the evolutionarily stable strategies (ESS), the model breaks another constraint of most approaches to coevolution that consider as fixed the number of coevolving species. The number of species comprising the ESS is influenced by a parameter that determines the predator's niche breadth. Depending upon the parameter's value the ESS may contain: 1) one prey and one predator species, 2) two prey and one predator, 3) two prey and two predators, 4) three prey and two predators, 5) three prey and three predators, etc. Evolutionarily, these different ESSs all emerge from the same model. Ecologically, however, these ESSs result in very different patterns of community organization. In some communities the predator species are ecologically keystone in that their removal results in extinctions among the prey species. In others, the removal of a predator species has no significant impact on the prey community. These varied ecological roles for the predator species contrasts sharply with the essential evolutionary role of the predators in promoting prey species diversity. The ghost of predation past in which a predator's insignificant ecological role obscures its essential evolutionary role may be a frequent property of communities of predator and prey.     

Macrogeographic variation of a pond predator's top-down effects in a common garden environment

Geographic variation in species behavior and life history has been well documented in biology. Species with wide geographic distributions (i.e., across a continent) but small home ranges (i.e., <1 km²) likely experience wide variability in abiotic environments across the entirety of their range, possibly exhibiting strong local adaptation. Understanding variation across a large geographic scale is especially important when considering species that have strong ecological importance, such as keystone species. Yet, few studies have compared the potential cascading ecological effects of a predator with a keystone role in at least part of its range. To understand how keystone ability in pond food webs can vary across a large geographic range, we conducted an artificial pond experiment with a known keystone predator in at least part of its range, the marbled salamander (Ambystoma opacum). To do so, we collected size-matched salamander larvae from three geographically distant populations (>650 km apart) in Ohio, Mississippi, and North Carolina and placed them in mesocosms with a suite of spring breeding amphibian prey species. We observed differential survival of some prey species leading to differences in spring-breeding amphibian diversity among the three predator populations, indicating that keystone predation may vary at a geographic scale. Prey diversity was lowest with predators from northern (Ohio) populations of salamanders. Further understanding of large-scale variability in ecologically important predators and the potential effects of translocating wide-ranging ambystomatid species is needed to direct future conservation efforts and preserve biodiversity.     

From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973-2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.     

Predation pressure shapes brain anatomy in the wild

There is remarkable diversity in brain anatomy among vertebrates and evidence is accumulating that predatory interactions are crucially important for this diversity. To test this hypothesis, we collected female guppies (Poecilia reticulata) from 16 wild populations and related their brain anatomy to several aspects of predation pressure in this ecosystem, such as the biomass of the four major predators of guppies (one prawn and three fish species), and predator diversity (number of predatory fish species in each site). We found that populations from localities with higher prawn biomass had relatively larger telencephalon size as well as larger brains. Optic tectum size was positively associated with one of the fish predator’s biomass and with overall predator diversity. However, both olfactory bulb and hypothalamus size were negatively associated with the biomass of another of the fish predators. Hence, while fish predator occurrence is associated with variation in brain anatomy, prawn occurrence is associated with variation in brain size. Our results suggest that cognitive challenges posed by local differences in predator communities may lead to changes in prey brain anatomy in the wild.     

Predatory fish loss affects the structure and functioning of a model marine food web

The rate of species loss is increasing on a global scale and predators are most at risk from human-induced extinction. The effects of losing predators are difficult to predict, even with experimental single species removals, because different combinations of species interact in unpredictable ways. We tested the effects of the loss of groups of common predators on herbivore and algal assemblages in a model benthic marine system. The predator groups were fish, shrimp and crabs. Each group was represented by at least two characteristic species based on data collected at local field sites. We examined the effects of the loss of predators while controlling for the loss of predator biomass. The identity, not the number of predator groups, affected herbivore abundance and assemblage structure. Removing fish led to a large increase in the abundance of dominant herbivores, such as Ampithoids and Caprellids. Predator identity also affected algal assemblage structure. It did not, however, affect total algal mass. Removing fish led to an increase in the final biomass of the least common taxa (red algae) and reduced the mass of the dominant taxa (brown algae). This compensatory shift in the algal assemblage appeared to facilitate the maintenance of a constant total algal biomass. In the absence of fish, shrimp at higher than ambient densities had a similar effect on herbivore abundance, showing that other groups could partially compensate for the loss of dominant predators. Crabs had no effect on herbivore or algal populations, possibly because they were not at carrying capacity in our experimental system. These findings show that contrary to the assumptions of many food web models, predators cannot be classified into a single functional group and their role in food webs depends on their identity and density in 'real' systems and carrying capacities.     

Interactions of multiple predators with different foraging modes in an aquatic food web

Top predators can have different foraging modes that may alter their interactions and effects on food webs. Interactions between predators may be non-additive resulting from facilitation or interference, whereas their combined effects on a shared prey may result in emergent effects that are risk enhanced or risk reduced. To test the importance of multiple predators with different foraging modes, we examined the interaction between a cruising predator (largemouth bass, Micropterus salmoides) and an ambush predator (muskellunge, Esox masquinongy) foraging on a shared prey (bluegill sunfish, Lepomis macrochirus) with strong anti-predator defense behaviors. Additive and substitution designs were used to compare individual to combined predator treatments in experimental ponds. The multiple predator interaction facilitated growth of the cruising predator in the combined predator treatments, whereas predator species had substitutable effects on the growth of the ambush predator. The combined predator treatments created an emergent effect on the prey; however, the direction was dependent on the experimental design. The additive design found a risk-reducing effect, whereas the substitution design found a risk-enhancing effect for prey fish. Indirect effects from the predators weakly extended to lower trophic levels (i.e., zooplankton community). Our results highlight the need to consider differences in foraging mode of top predators, interactions between predators, and emergent effects on prey to understand food webs.