东海及黄海南部带鱼的海底水温分布特征分析及分析方法改进

为了解东海及黄海南部带鱼随海底水温变化的分布特征并优化其分析方法,本实验基于2014—2015年4个季度大面积调查获得的带鱼资源分布数据,在前期研究的基础上对水温范围分析方法进行了优化,避免了分析当中的一些主观因素,创建了一种新的生物主体温度范围分析方法,并与传统频率方法和先前生物主体方法进行了比较。结果显示,频率法覆盖站点比例相对较好,但生物量覆盖率相对较差;先前生物主体法,生物量累积覆盖率最高,站点覆盖率也相对不错,缺点是操作过程加入的主观因素较多,且分布范围不连续、包含个别狭小范围;新方法,因其温度范围相对集中、短小,导致站点和生物量两方面的覆盖率均不是很高,但其所确定温度范围内的生物量密度最高,找到了能容纳最多生物量的最小范围,且分布范围相对连续、集中。生物空间分布的特征会直接影响温度范围分析方法的结果,如带鱼空间分布相对均匀,导致其频率分布中的一个分支与生物主体分布保持基本一致,而小黄鱼的空间分布相对集中,导致其频率分布与生物主体分布存在较大差别。分析还发现,生物量较高的群体一般均分布在水温较高的水域,这可能与群体良好的营养状况和快速生长的需求相关;生物空间上分布的不均匀特征,可能与物种的食物需求和捕食特点相联系;根据带鱼高生物量群体的时空分布特征,分析推测该群体可能对应带鱼的春夏生殖鱼群和秋季生殖鱼群。     

黄海南部近岸海域鱼类群落结构与区系划分

黄海南部海域具有复杂的水团结构和海流环境,使该海域鱼类群落产生较为复杂的空间结构。为研究黄海南部近岸海域鱼类群落结构特征,实验根据2014—2015年对该海域进行4个季度的底拖网调查数据,利用多元统计方法分析了该海域鱼类群落的分布格局并比较了各群落的相对生物量、生物多样性、优势种等结构特征。应用聚类分析(cluster)和多维标度排序(MDS)分析将黄海南部近岸海域划分为海州湾群落与江苏近岸群落;海州湾群落的平均相对资源量和生物多样性均高于江苏近岸群落,而暖温性与暖水性鱼类所占比重相对较低;海州湾群落的优势种主要为方氏云鳚、大泷六线鱼和日本鳀等,季节变化显著;江苏近岸群落的优势种为短吻红舌鳎、途、棘头梅童鱼和黑鳃梅童鱼等,季节变化不明显。研究表明,黄海南部近岸海域可划分为海州湾区系与江苏近岸区系,其鱼类群落结构特征具有显著差异。海流、水团、底质类型及水深等环境因子是形成鱼类群落结构差异的主要因素。     

Evaluation of a spatially sex‐specific assessment method incorporating a habitat preference model for blue marlin (Makaira nigricans) in the Pacific Ocean

Blue marlin, widely distributed throughout the Pacific Ocean, are sexually dimorphic, have certain preferred habitats, and migrate seasonally. These characteristics have been ignored in previous stock assessment models. A population dynamics model that includes spatial structure, and sex and age structure was therefore constructed and fitted to fisheries data for blue marlin, along with information on the relative density of the population over space derived from a habitat preference model that uses the oceanographic and biological variables sea‐surface temperature, mixed layer depth, sea‐surface height anomaly, and chlorophyll‐a concentration. Monte Carlo simulation was then used to examine the estimation performance of the stock assessment method. Estimates of management‐related quantities including current spawning stock biomass are substantially biased when the assessment method ignores seasonal movement and sexual dimorphism. We also found that (i) uncertainty about the relationship between catch rate and abundance influences estimation performance to a larger extent than uncertainty in catches, (ii) the outcomes of the assessment are sensitive to the values assumed for natural mortality and stock‐recruitment steepness, and (iii) the ratio of current spawning stock biomass to that at pre‐exploitation equilibrium appears to be the most robust among the quantities considered. We conclude that assessment methods for blue marlin in the Pacific Ocean need to take account of seasonal migration and sex structure to improve stock assessments.     

The relationship between the spatial distribution of fish,zooplankton and other environmental parameters in the Solina reservoir,Poland

The hydroacoustic estimation of fish abundance and distribution in the Solina reservoir was related to other biological and hydrological parameters. The increase of fish abundance corresponded to increased levels of eutrophication expressed by Secchi-disc visibility and chlorophyll a concentration. A negative correlation between fish and zooplankton abundances was observed, with changes in the zooplankton population structure, caused by fish grazing. Vertical distribution of zooplankton and fish were affected by the thermocline. Maximal concentrations were observed in a well-illuminated surface layer at the highest temperature and oxygen concentration.     

山东半岛东南部海域星康吉鳗资源密度时空分布及其与环境因子之间关系

为了解山东半岛东南部海域星康吉鳗资源密度时空变化及其与环境因子之间的关系,实验根据2016年10月和2017年1月、5月、8月山东半岛东南部海域4个航次底拖网调查数据,利用广义可加模型(GAM)分析了星康吉鳗资源密度时空分布特征及其与环境因子之间的关系。结果发现,星康吉鳗资源密度及分布具有明显的季节变化。春季,山东近海星康吉鳗资源密度为66.38 kg/h,夏季资源密度达到一年中最大值,为87.31 kg/h,秋季资源密度为79.01 kg/h,冬季资源密度大幅度降低,仅为10.44 kg/h。GAM模型结果显示,水深和海水底层温度对星康吉鳗资源分布影响最大。春季,星康吉鳗资源密度与水深、底温呈正相关关系,其分布范围较广,主要分布在海州湾中部海域(35°N沿线分布最多);夏季,其空间分布受水深影响,主要集中分布在水深20～30 m的山东半岛南部近岸海域;秋季,水深、底温、饵料生物量与星康吉鳗资源密度呈正相关,此时星康吉鳗分布较分散。冬季,星康吉鳗资源密度与水深呈正相关,此时主要分布在受黄海暖流影响的海州湾北部海域以及123.5°E～124°E海域。研究表明,星康吉鳗资源分布与其洄游习性和海域水温等水文特征的季节性变动有关,其分布特征在春季、冬季分别受青岛冷水团与黄海暖流影响显著。本研究有助于了解山东近海星康吉鳗群体的生活习性,为其资源的养护和管理提供依据。     

胶州湾鱼类生物量粒径谱特征

为了解胶州湾海域鱼类群落粒径结构特征,本研究根据2016-2017年4次底拖网调查数据构建了胶州湾鱼类生物量粒径谱,比较分析了各季节鱼类生物量粒径谱特征参数的异质性。结果显示,胶州湾Sheldon型鱼类生物量粒径谱呈现单峰模式,鱼类粒径范围为-3~10粒径级,粒径分布相对均匀,且峰值位于5~6粒径级,主要由大泷六线鱼和褐菖鲉等小型鱼类构成。各季节鱼类标准化生物量粒径谱曲率和曲度差异显著,其中春季曲率最大且曲线较平缓,夏季曲率最小且曲线相对陡峭。谱线曲度和曲率的季节变化主要与粒径小的细纹狮子鱼、大泷六线鱼、玉筋鱼及长丝虾虎鱼等地方常栖性种类和赤鼻棱鳀、白姑鱼等季节洄游性种类的出现有关。Abundance-biomass comparison(ABC)曲线表明,春季处于未干扰状态,冬季处于中度干扰状态,夏秋季处于严重干扰状态。研究表明,胶州湾鱼类群落总体处于严重干扰状态,鱼类生物量谱呈现单峰型,粒径结构以小型粒径鱼类为主,且地方常栖性和季节洄游性鱼类等补充群体的数量和粒径大小影响鱼类生物量粒径谱的峰型和曲率。     

Niche partitioning of fish assemblages in a mountain stream with frequent natural disturbances – an examination of microhabitat in riffle areas

Abstract –  Microhabitat usage by different species of fish was studied to examine the potential for niche partitioning in riffle areas of a mountain river characterised by frequent natural disturbances. We randomly sampled 96 riffle‐area plots (mostly 1 m × 1 m) in a stream in south‐western Taiwan for fish abundance and microhabitat characteristics during the dry season from November 2008 to March 2009. Density and electivity values associated with microhabitat usage indicate the presence of niche partitioning among four major species of fish based largely on flow velocity and water depth. Hemimyzon formosanum, a herbivore, prefers higher velocity and shallower water but avoids sandy substrate. Onychostoma alticorpus, a herbivore, prefers lower velocity and deeper water. Rhinogobius nantaiensis, an omnivore, prefers higher velocity, shallower water and gravel‐sized substrate but avoids boulder substrate. Acrossocheilus paradoxus, an omnivore, prefers lower velocity and deeper water but avoids pebble‐sized substrate. The existence of differentiation of microhabitat preference and overlap of microhabitat usage suggests that interspecific competition is a factor affecting the structure of this fish assemblage, which may remain at early successional stages because of frequent disturbances in the stream. Study results also show that density and electivity detect preference differently. Density detects microhabitat preference for two herbivores while electivity detects microhabitat preference for two omnivores.     

Two species of seabirds foraged in contrasting marine habitats across the cold-water belt along the coast of northern Hokkaido in the southwestern Okhotsk Sea

To understand the environmental factors affecting the density of foraging seabirds across the cold-water belt in the southwestern Okhotsk Sea, we conducted a 1-day (180-km transect length) shipboard seabird survey off the northeastern coast of Hokkaido during summer in 2019, along with acoustic observations of potential prey (zooplankton and fish) biomass, thermosalinograph measurements, and CTD observations. Planktivorous short-tailed shearwaters Ardenna tenuirostris (66% of total seabirds) and piscivorous rhinoceros auklets Cerorhinca monocerata (28%) were predominant, but foraged in contrasting habitats. A large foraging flock of shearwaters was observed in the cold-water belt zone, including its front with coastal Soya Warm Current Water and the offshore Fresh Surface Okhotsk Sea Water, where surface chlorophyll a concentrations were the highest but not related to their prey (zooplankton) biomass at any spatial scale between 4.6 and 9.2 km. In contrast, the density of auklets was high in the coastal Soya Warm Current Water, where the acoustically determined fish biomass was large, and showed a positive relationship with the fish biomass especially in the lower layer (29–104 m depth) at any spatial scale. This species-specific difference in response to prey biomass might be related to prey-searching behaviors; i.e., rhinoceros auklets search prey underwater visually, but short-tailed shearwater can use both visual and olfactory cues to locate zooplankton patches from the air.

     

Diet and fish choice of Eurasian otters (<Emphasis Type="Italic">Lutra lutra</Emphasis> L.) in fish wintering ponds in Hungary

The diet composition and fish preference of piscivorous Eurasian otters (Lutra lutra) were studied in two fish farm systems in Hungary using spraint (otter faeces) analysis during two wintering periods. The primary food source of otters in both fish farms was fish (97–99% of biomass). The main fish prey was small-sized, below 100 g in weight (96% in both areas), while fish prey above 500 g comprised only 0.1–0.4% of the diet. The bulk of the otters’ diet consisted of less-valued species, especially non-native Prussian carp (Carassius auratus gibelio). Consumption of commercial fish species ranged between 15 and 31% of the total diet. Otters preferred fish below 100 g in weight (Ivlev’s electivity index, E _i = 0.65–0.70), and showed a lesser preference for (or avoided) fish above 100 g in weight (E _i = −0.37–1.00). With regard to species distribution, otters preferred small (below 100 g) grass carp (Ctenopharyngodon idella), zander (Sander lucioperca), pike (Esox lucius), Prussian carp, topmouth gudgeon (Pseudorasbora parva), while they consumed common carp (Cyprinus carpio), the most important commercial species, proportionally to its abundance in the environment (E _i = −0.18–0.29).     

Hydrological connectivity in coastal inland systems: lessons from a Neotropical fish metacommunity

de Macedo-Soares PHM, Petry AC, Farjalla VF, Caramaschi EP. Hydrological connectivity in coastal inland systems: lessons from a Neotropical fish metacommunity.
Ecology of Freshwater Fish 2010: 19: 7–18. © 2009 John Wiley & Sons A/S
Abstract –  We assessed the influence of hydrological connectivity in structuring fish communities through seasonal samplings of environmental variables and fishes in a coastal lagoon and associated pools in the Restinga de Jurubatiba National Park, Brazil. Community structure attributes such as species richness, numerical density and biomass, Shannon–Wiener diversity index and evenness were compared between periods of the lowest and highest hydrological connectivity, while the environmental gradient and fish zonation were explored through ordination techniques. The greater hydrological connectivity established in the rainy season promoted the homogenisation of most environmental variables and fish species, which differed markedly from the arrangement observed in the dry season. Despite variation in fish species composition, community attributes showed non-significant differences between the dry and rainy seasons. The patterns of composition and numerical density in pools were strongly influenced by local factors, especially salinity, dissolved oxygen, total phosphorous concentration and water colour in the dry season, in addition to total nitrogen concentration and depth in the rainy season. Comparable to the role played by flood pulses in river-floodplain systems, the hydrological connectivity in these tropical coastal waterbodies seems to strongly influence fish community structure, and, therefore to determine regional biodiversity.     

浙江中部披山海域蟹类群落结构特征及其与环境因子的关系

根据2015年11月、2016年2、5、8月在浙江中部披山海域进行渔业资源调查所获得的资料,分析了浙江中部披山海域蟹类的种类组成、优势种、生物多样性等群落结构特征,同时,研究了群落结构的相似性,以及群落结构与环境因子之间的关系。结果显示,所得渔获物共鉴定出蟹类19种,隶属于6科9属,其中优势种为三疣梭子蟹、日本蟳、双斑蟳和隆线强蟹等4种。不同季节的蟹类种类组成差异较大,而优势种类组成变化较大,其中,蟹类种类数以夏季最多,冬季最少。从不同水深区域的渔获种类分布趋势分析,蟹类种类数以5~15 m水深带较多,小于5 m和15~20 m水深带较少。生物多样性指数夏季最高,冬季最低。从不同水深区域分析,多样性指数在10~15 m水深带较高,5~10 m水深带较低。冗余分析结果认为,表层水温、表层盐度和底层盐度是影响调查海域蟹类种类组成和群落结构特征的主要环境因子。     

舟山群岛外海域春秋季鱼类群落结构及生物多样性

为了解舟山群岛外海域鱼类资源状况,于2018年4月(春季)和10月(秋季)在舟山群岛外海域进行的渔业资源调查,用得到的数据分析了鱼类种类组成、优势种、群落结构及生物多样性。结果显示:(1)舟山群岛外海域鱼类有106种,隶属于12目47科80属;(2)本次调查鱼类种类数春季为63种,秋季为85种,总体与历史调查结果相比春季明显减少,而秋季却略有增加;(3)2个季节的鱼类优势种更替明显,春季优势种为黄鮟、日本红娘鱼和细条天竺鲷,秋季优势种为细条天竺鲷和日本发光鲷;(4)鱼类生物多样性指数值为秋季大于春季,春、秋季均匀度指数(J')对比多样性指数(H')和丰富度指数(D)都较小,且春、秋季生物多样性指数值基本都是随着水深的增加而增加,均以80 m以上水深区间较高;(5)通过聚类分析与非度量多维标度排序分析可得,春季在35%的相似水平上分为3个群落,秋季在45%相似水平上分为2个群落,秋季物种相似度比春季高,春季在生物量和丰度水平上均比秋季低。研究表明,舟山群岛外海域的春季鱼类群落结构可能处于未受干扰状态,秋季鱼类群落结构可能处于严重干扰状态。     

Modelling changes in the length–frequency distributions of fish larvae using field estimates of predator abundance and size distributions

The goal of this study is to determine if an individual-based size-dependent model can realistically simulate changes in the length–frequency distributions of several species of fish larvae collected in Conception Bay in 1993 and 1994, using field estimations of growth and predator abundance. We first model the length–frequency distribution of field samples with the best possible estimates of mean growth rate. Then, we add predation mortality given the characteristics of the predator community observed during our surveys, which was composed of macrozooplankton and adult capelin. The larval fish community is generally not affected by predation by macrozooplankton, as the average instantaneous mortality rate predicted by the model was 0.004 day^–1. Fish larvae appear to be more vulnerable to predation by the population of adult capelin. We estimate that an abundance of adult capelin ranging between 0.2 and 1.0 individuals per 1000 m^–3 may have a substantial impact on the larval fish community. The predictions of an individual-based model are directly related to the accuracy of estimates of the mean growth rates of the larval fish cohorts. We find that it is difficult to differentiate size-selective removal of individuals from random selection by analysing changes of the length–frequency distributions of the larval fish community.     

Association between biogeographical factors and boreal lake fish assemblages

Abstract  Five regions in insular Newfoundland Canada, comprising 152 lakes, were studied to identify associations between species composition (presence and absence), geographical location and environmental variables (pH, area, depth, alkalinity, secchi disc depth and shoreline development factor). Correspondence analysis and canonical variate analysis were used to distinguish regional patterns. Five biologically and environmentally distinct areas were identified. The degree of association between biological, environmental and geographical distances were contrasted using Mantel's test. Regional fish community structure was significantly correlated with large-scale geographical distance but not with environmental parameters or small scale distance. It was proposed that large scale processes such as post-glacial dispersion, climate and recent species introductions are important determinates in structuring regional fish assemblages. Differences in individual lake character were important determinates in intraregional variability in fish assemblage type. Sampling strategies for regional modelling and management are discussed.     

Assessment of changes in the ichthyofauna in a tropical reservoir in south-eastern Brazil: Consequences of global warming?

Increases in temperatures over the last century, more intensively after the eighties, were recorded in several databases for the south-eastern Brazil. These increases are likely to change fish communities in aquatic systems by decreasing abundances, biomass and sizes of some species more sensitive to climate change. Reservoirs may be particularly susceptible to the effects of climate change, as they isolate previously connected areas limiting fish dispersal and migration, as well as increasing water temperature and thermal stratification. We assessed temporal changes in the abundance and biomass of the ichthyofauna in an isolate reservoir (inflowing waters from small tributaries in a highland region) aiming to associate changes with climate effects over three decades (1990, 2000, 2010). Two abundant native species, a loricariid Loricariichthys castaneus (Castelnau, 1855) and a cichlid Geophagus brasiliensis (Quoy & Gaimard, 1824), were selected to assess eventual response to climate changes in their geographical distribution in the Southeast Atlantic Hydrographic Region, considering different carbon emission scenarios using ecological niche models. A decrease in the densities and biomass of the ichthyofauna and of the two selected species were observed in both summer and winter over the three decades, coinciding with increases in temperatures. These decreases were correlated with increases in positive anomalies of temperature, which may be an indicator of climatic changes, although with low explanation of the data variation. The mean size of L. castaneus decreased between the nineties and 2010, whereas no significant trend was found for the size of G. brasiliensis. Our predictions of new area for future adequacy indicated a loss in the distribution area for both species (mainly for G. brasiliensis), considering the most pessimistic scenario for 2050 and 2070, possibly due to climatic changes. Both hydroelectric dams and global warming pose threats to freshwater fish diversity, and both will interact. Changes observed in the fish assemblage over the last decades were essential for an assessment of the consequences of global warming in this type of reservoir combining larger scale studies with future projections.     

Biogeographical region and environmental conditions drive functional traits of estuarine fish assemblages worldwide

Assessing trait–environment relationships is crucial for predicting effects of natural and human‐induced environmental change on biota. We compiled a global database of fish assemblages in estuaries, functional traits of fishes and ecosystem features of estuaries. And we quantified the relative importance of ecosystem features as drivers of patterns of fish functional traits among estuaries worldwide (i.e. drivers of the proportions of fish traits). In addition to biogeographical context, two main environmental gradients regulate traits patterns: firstly temperature, and secondly estuary size and hydrological connectivity of the estuary with the marine ecosystem. Overall, estuaries in colder regions, with larger areas and with higher hydrological connectivity with the marine ecosystem, have higher proportions of marine fish (versus freshwater), macrocarnivores and planktivores (versus omnivores, herbivores and detritivores) and larger fish, with greater maximum depth of distribution and longer lifespan. The observed trait patterns and trait–environment relationships are likely generated by multiple causal processes linked to physiological constraints due to temperature and salinity, size‐dependent biotic interactions, as well as habitat availability and connectivity. Biogeographical context and environmental conditions drive species richness and composition, and present results show that they also drive assemblage traits. The observed trait patterns and trait–environment relationships suggest that assemblage composition is determined by the functional role of species within ecosystems. Conservation strategies should be coordinated globally and ensure protection of an array of estuaries that differ in ecosystem features, even if some of those estuaries do not support high species richness.     

Spatial distribution pattern and physical – biological interactions in the larval notothenioid fish assemblages from the Bransfield Strait and adjacent waters

The Bransfield Strait and adjacent waters represent one of the most important areas of larval retention off the Antarctic Peninsula. The species composition of larval fish assemblages has been described in detail in previous surveys carried out in the area, but the role of environmental parameters influencing the spatial distribution of early life stages was poorly known. By applying generalized additive models and multivariate analyses, we evaluated the role of environmental variables in shaping the small‐scale distribution of larval fish and investigated the spatial structure of the larval assemblage. It consisted of a few dominant notothenioid species, such as Champsocephalus gunnari, Lepidonotothen squamifrons, Lepidonotothen larseni, Pleuragramma antarctica and Trematomus scotti, and several other rarely caught species. Sea water temperature, salinity and sampling depth were the most important factors determining the spatial distribution of fish with different relative contributions, together explaining more than 80% of total deviance observed. Species richness was mostly affected by salinity, probably due to the narrow range of salinity preference by the species. Cluster analysis of abundance and presence data identified six and five distinct groups, respectively, each of them with substantial contributions of single or rarely two species. Differences in reproductive strategies of adult populations and spatial distribution of early life stages driven by different larval behaviour in response to environmental factors contribute to maintaining a well‐structured larval fish assemblage, ensuring spatial and food niche partitioning.     

夏季台湾海峡北部及邻近海域鱼类群落结构及环境解释

根据2014年7月台湾海峡中部、北部及邻近水域渔业资源底拖网调查资料,分析了该水域的鱼类群落结构,包括种类组成、区系特点、优势种、多样性及空间分布等,探讨了鱼类群落结构与环境因子的关系,并阐述了鱼类群落的群聚结构。结果表明,调查海域共捕获鱼类84种,隶属于16目、48科、69属。从种类数上看,适温类型以暖水种为主,区系类型以大陆架浅水底层鱼类以及大陆架岩礁性鱼类为主;从生物量组成上看,区系类型则以大陆架浅水中低层鱼类为主。优势种为带鱼(Trichiurus japonicus)和七星底灯鱼(Benthosema pterotum),合计渔获质量占总渔获质量的46.92%,对总渔获质量的回归贡献值分别为0.30和0.62;高生物量区集中分布在台湾海峡北口海域,在空间上表现为明显的生态位重叠。层次聚类分析将群落优势物种划分为2个主要的生态类型:沿海类型和广布类型,沿海类型的代表性物种包括龙头鱼、黄鲫、鳓;广布类型物种有带鱼、窄颅带鱼、七星底灯鱼、竹荚鱼、刺鲳、麦氏犀鳕。Shannon-Wiener多样性、Pielou均匀度和Margalef丰富度的变化范围分别为0.22~2.31、0.11~0.72、0.36~4.04,平均值分别为1.44、0.51、1.59,丰富度与总渔获质量表现为显著负相关(r=?0.65),与深度显著正相关(r=0.48)。物种?环境典范对应分析表明,与物种空间结构关系密切的影响因子为底盐、底层无机氮、底温、表底温差和水深;而影响群落优势群体分布的因子为底温、底盐和底层无机氮。     

Niche processes and conservation implications of fish community assembly in a rice irrigation system

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The effect of depth, distance from dam and habitat on spatial distribution of fish in an artificial reservoir

Abstract –  Spatial distribution of the fish community in the deep canyon-shaped Římov Reservoir, Czech Republic, was studied using overnight gillnetting fishing in 1999–2007. Effects of depth, distance from the dam to the tributary and habitat type (benthic and pelagic) on fish community structure, catch per unit of effort (CPUE), biomass per unit of effort (BPUE) and average weight were tested. Fish were recorded in all sampled depths and parts of the reservoir. Redundancy analysis revealed that effects of three environmental variables were significant and most variability was explained by depth, then by distance from the dam and habitat type. CPUE and BPUE of all species decreased with depth and responses of individual species to depth were similar for juvenile and adult fish. Number of species, CPUE and BPUE of all species except perch increased heading towards the tributary and peaked close to or at the tributary part of the reservoir. Responses of juvenile fish to distance from the dam differed from that of adult fish. Structure of fish community differed in benthic and pelagic habitats with species preferably occupying epipelagic (bleak, asp, rudd and juvenile bleak, roach and bream) or littoral waters (perch, pikeperch, ruffe, roach, bream and juvenile percids). This study showed that fish distribution in the reservoir followed distinct patterns, which were probably shaped by a combination of physiological constraints plus trade-off between food resources and competition.