Development of an Index of Trophic Completeness for benthic macroinvertebrate communities in flowing waters

The analysis of the trophic structure of benthic macroinvertebrate communities can be used in biological assessments of the condition of river ecosystems. Using the trophic, or functional approach, the Index of Trophic Completeness (ITC) was developed. The goal was to overcome the problems and drawbacks of using conventional diversity or biotic indices in biological assessments of rivers, such as limitation to distinct geographical regions or focus on species richness without regard for ecosystem functioning. Following an extensive review of the literature on the trophic characteristics of benthic macroinvertebrates, a large number of species (±300) were characterized according to a number of trophic criteria: plant:animal ratio in the diet, feeding mechanism, food size, food acquisition behaviour, and energy and substance transfers. On the basis of their trophic characteristics, the species could be divided into 12 trophic groups. After examination of data from geographically diverse rivers, it was concluded that any undisturbed riverine benthic macroinvertebrate community should be represented by members of each of these 12 trophic groups, with each group fulfilling a function in the benthic community. Being a community which plays a central role in the functioning of the aquatic ecosystem, the benthic invertebrates are expected to respond to disturbances to the hydrobiocoenose. The outcome of an ITC assessment is clearly presentable in the form of a pie graph with 12 wedges, each representing one of the 12 defined trophic groups. Functionally complete communities are represented by 12 wedges; a blank wedge indicates that a trophic group is not represented. This paper describes the preliminary developments in the ITC method, its potential as a biological assessment method in rivers in different geographical zones, and presents examples of trial mappings of Russian and European rivers. The application of the ITC to these rivers demonstrated the absence of ITC trophic groups at sites under the influence of anthropogenic activity.     

Patterns of fish species richness in the Seine River basin,France

Variation in fish assemblage structure was examined in different rivers of the Seine Basin. Factor analysis was used to identify similarities among samples and species, and to show ichtyological changes along an upstream-downstream gradient. Fish species richness was correlated with catchment area, distance from the source and number of individuals. Relationships between species richness and number of individuals in a sample were largely an artifact of sampling.Species richness increased with river size, reached a maximum in midsize rivers, then decreased in large rivers. This species richness pattern was consistent with the model of the River Continuum Concept. However, possible causes could include greater pollution effects in large rivers. Distribution of feeding guilds was related to river size. Species richness and proportion of omnivores and piscivores increased with river size, whereas species richness and proportion of invertivores declined downstream. These patterns suggest that trophic diversity of fish assemblages may be related to food availability.     

Using diatom life-forms and ecological guilds to assess organic pollution and trophic level in rivers: a case study of rivers in south-eastern France

The European Union’s Water Framework Directive has set a target of achieving good ecological status for all aquatic environments in Europe by 2015. In order to determine the quality of aquatic environments, biological indicators such as diatoms are often used. However, biotic diatom indices can be difficult and time consuming to use because of complexity of species determination. We investigated whether the biological traits of diatoms in rivers (life-forms, size classes and ecological guilds) could be used to assess organic pollution and trophic level. We worked on a data set comprising 315 diatom species, determined at 328 river stations of south-east France and a variety of parameters. The abundances of some biological traits differed significantly between the different organic pollution and trophic levels, particularly stalked diatoms, and the motile and low-profile guilds.     

Delineating reef fish trophic guilds with global gut content data synthesis and phylogeny

Understanding species’ roles in food webs requires an accurate assessment of their trophic niche. However, it is challenging to delineate potential trophic interactions across an ecosystem, and a paucity of empirical information often leads to inconsistent definitions of trophic guilds based on expert opinion, especially when applied to hyperdiverse ecosystems. Using coral reef fishes as a model group, we show that experts disagree on the assignment of broad trophic guilds for more than 20% of species, which hampers comparability across studies. Here, we propose a quantitative, unbiased, and reproducible approach to define trophic guilds and apply recent advances in machine learning to predict probabilities of pairwise trophic interactions with high accuracy. We synthesize data from community-wide gut content analyses of tropical coral reef fishes worldwide, resulting in diet information from 13,961 individuals belonging to 615 reef fish. We then use network analysis to identify 8 trophic guilds and Bayesian phylogenetic modeling to show that trophic guilds can be predicted based on phylogeny and maximum body size. Finally, we use machine learning to test whether pairwise trophic interactions can be predicted with accuracy. Our models achieved a misclassification error of less than 5%, indicating that our approach results in a quantitative and reproducible trophic categorization scheme, as well as high-resolution probabilities of trophic interactions. By applying our framework to the most diverse vertebrate consumer group, we show that it can be applied to other organismal groups to advance reproducibility in trait-based ecology. Our work thus provides a viable approach to account for the complexity of predator–prey interactions in highly diverse ecosystems.

The diversity of life on our planet has produced a remarkable variety of biological traits that characterize different species, and such traits are widely considered an alternative to taxonomy to increase our understanding of biodiversity and ecosystem functioning. This study presents an unbiased and fully reproducible framework to delineate trophic guilds in reef fishes.     

Structure of tropical river food webs revealed by stable isotope ratios

Fish assemblages in tropical river food webs are characterized by high taxonomic diversity, diverse foraging modes, omnivory, and an abundance of detritivores. Feeding links are complex and modified by hydrologic seasonality and system productivity. These properties make it difficult to generalize about feeding relationships and to identify dominant linkages of energy flow. We analyzed the stable carbon and nitrogen isotope ratios of 276 fishes and other food web components living in four Venezuelan rivers that differed in basal food resources to determine 1) whether fish trophic guilds integrated food resources in a predictable fashion, thereby providing similar trophic resolution as individual species, 2) whether food chain length differed with system productivity, and 3) how omnivory and detritivory influenced trophic structure within these food webs. Fishes were grouped into four trophic guilds (herbivores, detritivores/algivores, omnivores, piscivores) based on literature reports and external morphological characteristics. Results of discriminant function analyses showed that isotope data were effective at reclassifying individual fish into their pre-identified trophic category. Nutrient-poor, black-water rivers showed greater compartmentalization in isotope values than more productive rivers, leading to greater reclassification success. In three out of four food webs, omnivores were more often misclassified than other trophic groups, reflecting the diverse food sources they assimilated. When fish δ¹⁵N values were used to estimate species position in the trophic hierarchy, top piscivores in nutrient-poor rivers had higher trophic positions than those in more productive rivers. This was in contrast to our expectation that productive systems would promote longer food chains. Although isotope ratios could not resolve species-level feeding pathways, they did reveal how top consumers integrate isotopic variability occurring lower in the food web. Top piscivores, regardless of species, had carbon and nitrogen profiles less variable than other trophic groups.     

Fruit of the forest - larval sea lamprey Petromyzon marinus are fuelled by allochthonous resources

We used stable isotopes of carbon, hydrogen and nitrogen to quantify the trophic position and resource use of larval sea lamprey Petromyzon marinus, four benthic macroinvertebrate functional feeding guilds (scraper, shredder, collector and predator) and other fishes in three rivers in eastern Canada. Larval lamprey and most invertebrate guilds foraged as primary consumers in all rivers whereas all other fishes predominantly foraged as secondary consumers. Larval lamprey obtained 75–85% of their resources from allochthonous derived material. This level exceeded all invertebrate guilds, which assimilated approximately 50% allochthonous and 50% autochthonous materials and fishes, which predominantly assimilated between 25% and 60% allochthonous material. Larval lamprey occupied a unique position within the river food webs analysed and show remarkable fidelity to a trophic niche specialising on terrestrially derived detritus.     

Using fish guilds to assess community responses to temperature and flow regimes in unregulated and regulated Canadian rivers

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Functional Diversity of Bacterioplankton in Three North Florida Freshwater Lakes over an Annual Cycle

The phylogenetic diversity of freshwater bacterioplankton is widely known; however, there is minimal information on the functional diversity of the bacterial communities in these systems. Understanding the functional diversity of freshwater bacterial communities is important because heterotrophic bacteria can be impacted by anthropogenic perturbation, which in turn can alter biogeochemical cycling. The objective of this study was to use Biolog EcoPlates to acquire spatial and temporal community-level physiological profiles (CLPPs) for three freshwater lakes of different trophic levels and to assess the phylogenetic affiliation of the bacteria responsible for utilizing the various carbon guilds within them by denaturing gradient gel electrophoresis (DGGE). CLPP results showed that bacterial communities utilized the carbon guilds similarly between sites within the three lakes. However, when the metabolic profile of each lake was compared, Lake Bradford and Moore Lake were more similar to one another than to Lake Munson, the eutrophic lake. Additionally, although the bacteria that utilized the five carbon guilds included representatives from the classes α-, β-, γ-Proteobacteria, Flavobacteria and Sphingobacteria, Lake Munson had the largest number of Flavobacteria and γ-Proteobacteria in comparison to Moore Lake and Lake Bradford. Overall, Biolog analysis was useful in identifying differences in the functional diversity of bacterial communities between lakes of different trophic statuses and can be used as a tool to assess ecosystem health.     

Seasonal and spatial variations of fish trophic guilds in a shallow,semi-enclosed tropical estuarine bay

A study on seasonal and spatial variations of feeding habits and trophic guilds of dominant fish species in Pattani Bay during March 2003 to February 2004 was aimed at classifying diet composition, identifying dominant food components of each species, categorizing trophic guilds of the community and evaluating effects of habitat characteristics and seasonality on guild organization. Most fishes showed high food intake, fed on a diverse range of food items but relied heavily on calanoid copepods and shrimps. All species, with the exception of Epinephelus coioides, were classified as specialist feeders. Four main dietary guilds were classified. Three of these were classified as the guilds dominated by at least two major food items. Significant variations in trophic guilds of 28 fish species based on habitat types and seasons were also identified. They could be divided into three seasonal groups and three site-groups and a single site. Trophic organization for each season and habitat ranged from two to four groups. This information identifies groups of fishes that seasonally and spatially utilize different food resources within a semi-enclosed estuarine bay ecosystem.     

Fish zonations and guilds as the basis for assessment of ecological integrity of large rivers

Longitudinal zonation concepts describe the downstream changes in chemico-physical and biological properties of rivers. Including information on ecological fish guilds can enhance the usefulness of fish zonation concepts, in a way that they can be used as tools for assessment and management of the ecological integrity of large rivers. We present an ecological characterization of fish zones and fish communities in near-natural and in regulated large rivers in Europe (the River Doubs in France and the Rivers Rhine and Meuse in the Netherlands), using guild classifications of several life-history traits of fish and national Red Lists of threatened species. The Doubs data set was also analyzed using indices of the sensitivity of fish species to environmental degradation and indices for eurytopy. In these rivers, the number of ecological guilds per zone increases downstream, and there are clear shifts in the structure of the guilds. Flow preference and reproduction ecology of river fish are closely linked. The proportion of rheophilic species in the fish community decreases downstream, and the proportions of limnophilic and eurytopic species increase. Lithophilic and psammophilic spawners are dominant in the upper zones, whereas the lower zones are dominated by phytophilic and phytolithophilic spawners. The proportion of zoobenthivorous and periphytivorous species decreases downstream, and the proportion of zooplanktivorous and phytivorous species increases. However, because the European fish fauna mainly consists of feeding generalists, the discriminative abilities of simplistic feeding guild classifications are not very high. Guilds of sensitive, stenoecious species that share life history strategies that are highly adapted to specific riverine conditions (rheophils and limnophils) have declined far more than generalist species that can survive in a wide range of habitats that are not characteristic of natural river ecosystems. Because of the subsequent over-abundance of the eurytopic species the original longitudinal fish zonations are hardly recognizable anymore in heavily impacted large rivers such as the River Rhine. Hence these rivers do not meet the criteria for ecological integrity. Within a specific fish region, a suitable way of analyzing and monitoring the impact of human disturbance on the structure of the fish community is by comparing the guild structure of the present state of a fish zone with that of the reference situation.     

Young-of-the-year fish assemblages as an alternative to adult fish monitoring for ecological quality evaluation of running waters

Studies on the Upper Mississippi River, particularly over the last 15 years, have contributed to our understanding of trophic processes in large rivers. The framework established by earlier population-specific studies, however, cannot be overlooked. Examination of the feeding habits of fish ranging from planktivores to piscivores gave the first indication that trophic processes were influenced by the spatial complexity and annual hydrological patterns of river-floodplain ecosystems. Experimental studies, which have often been considered impossible or impractical in large rivers, demonstrated the potential for biotic controls of system dynamics through predator–prey and competitive interactions. Such studies have been particularly helpful in understanding the potential impact of non-native species, including zebra mussels and Asian carp, to biodiversity and secondary production. Our understanding of riverine ecosystem function expanded greatly as food web studies began the application of a new tool—natural stable isotopes. Studies employing stable isotopes illustrated how food webs in a number of large rivers throughout the world are supported by the autochthonous production of microalgae. This study, coupled with other studies testing the prevailing models of riverine ecosystem function, has brought us to a point of better understanding the nature of river ecosystem functions. It is through looking back at the earlier studies of fish diet that we should realize that the temporal and spatial complexities of river ecosystem function must still be addressed more fully. This and a better grasp of the significance of the arrangement of patches within the riverine landscape will prove beneficial, as we assess the appropriate scale of river rehabilitation with an eye on how rehabilitation promotes productivity within complex ecosystems, including the Upper Mississippi River.     

筑坝河流磷素的迁移转化及其富营养化特征

人类活动过量营养物质输入是导致河流富营养化的主要原因,而河道过度的人为调控则进一步复杂化了河流的营养状态变化。闸坝是河流人为调控的重要工程措施之一,提高水资源利用效率的同时严重干扰了河流自然的生物地球化学循环,产生诸多负面生态环境效应。磷素的迁移转化对河流的营养限制作用受到越来越多的关注,国内外已有研究在筑坝河流磷的富营养化特征方面,已经取得了较为深刻的认识:水库闸坝建设滞留大量磷素,导致河流水体磷含量升高、营养物质比例变化,沉积物储存过量磷素形成的内源释放威胁,以及进一步浮游植物和有害藻类的生长响应等,使得筑坝河流的富营养化生态风险升高;在此基础上,也提出了根据降雨分配和闸控库区储水,合理设置闸坝泄流方式,以改善筑坝河流富营养化生态风险的重要管理思路。对于闸坝调控作用与水体富营养化的定量关系还有待进一步的探讨,而且随着河流资源开发和人为调控力度的增强,河流闸坝建设所产生的系列生态环境问题日益严峻,对此提出还需要系统研究的方向:闸坝调控作用下河流磷素的富营养化机制及其与氮、碳等元素的耦合作用,筑坝河流沉积物内源污染的综合管理,以及闸控景观河流的生态建设和修复等。     

Hydrogeomorphology and river impoundment affect food-chain length of diverse Neotropical food webs

Food-chain length is a central characteristic of ecological communities that affects community structure and ecosystem function. What determines the length of food chains is not well resolved for most ecosystems. Herein, we examine environmental correlates of food-chain length based on the productivity hypothesis, compare food-chain lengths among aquatic ecosystem types and identify bi-directional effects of river impoundment on food-chain length in the Paraná River Basin of South America. Both temperature regime, a surrogate of productivity, and ecosystem type significantly affected food-chain length in independent analyses. However, when analyzed together, only ecosystem type explained significant variation in food-chain length. Food chains were longest in reservoirs, and shortest in high-gradient rivers. The proximate mechanism driving this pattern appears to be body-size ratios of primary consumers to apex predators, which differ among trophic pathways. Food chains based on phytoplankton production may have an additional size-structured link not present in food chains based on other basal sources such as detritus and algae. Hydrogeomorphology is the ultimate mechanism influencing food-chain length because it affects the relative importance of basal carbon sources supporting higher trophic levels, which through differences in the number of trophic links along the different size-structured pathways, appears to drive the observed patterns in food-chain length. We discuss a hypothesis of food-chain length that integrates energy flow and size-structure, facilitates inclusion of temporal dynamics and which is readily testable in both 'closed' and 'open' ecosystems.     

Influence of lateral gradients of hydrologic connectivity on trophic positions of fishes in the Upper Mississippi River

1. Riverscapes consist of the main channel and lateral slackwater habitats along a gradient of hydrological connectivity from maximum connection in main channel habitats to minimum connection in backwaters. Spatiotemporal differences in water currents along this gradient produce dynamic habitat conditions that influence species diversity, population densities and trophic interactions of fishes. 2. We examined the importance of lateral connectivity gradients for food web dynamics in the Upper Mississippi River during spring (high flow, moderately low temperatures) and summer (low flow, higher temperatures). We used literature information and gut contents analyses to determine feeding guilds and stable isotope analysis to estimate mean trophic position of local fish assemblages. During June and August 2006, we collected over 1000 tissue samples from four habitats (main channel, secondary channels, tertiary channels and backwaters) distributed within four hydrologic connectivity gradients. 3. Mean trophic position differed among feeding guilds and seasons, with highest values in spring. Mean trophic position of fish assemblages, variability in trophic position and food chain length (maximum trophic position) of the two dominant piscivore species (Micropterus salmoides and M. dolomieu) in both seasons were significantly associated with habitat along the lateral connectivity gradient. Food chain length peaked in tertiary channels in both seasons, probably due to higher species diversity of prey at these habitats. We infer that food chain length and trophic position of fish assemblages were lower in backwater habitats in the summer mainly because of the use of alternative food sources in these habitats. 4. A greater number of conspecifics exhibited significant among‐habitat variation in trophic position during the summer, indicating that low river stages can constrain fish movements in the Upper Mississippi River. 5. Results of this study should provide a better understanding of the fundamental structure of large river ecosystems and an improved basis for river rehabilitation and management through knowledge of the importance of lateral complexity in rivers.     

Linking Flow Regime,Floodplain Lake Connectivity and Fish Catch in a Large River-Floodplain System,the Volga–Akhtuba Floodplain (Russian Federation)

River-floodplain systems are amongst the most productive—but often severely impacted—aquatic systems worldwide. We explored the ecological response of fish to flow regime in a large river-floodplain system by studying the relationships between (1) discharge and inundated floodplain area, with a focus on spatial and temporal patterns in floodplain lake connectivity, and (2) flood volume and fisheries catch. Our results demonstrate a non-linear relationship between discharge and floodplain inundation with considerable hysteresis due to differences in inundation and drainage rate. Inundation extent was mostly determined by flood volume, not peak discharge. We found that the more isolated lakes (that is, lakes with a shorter connection duration to the river) are located at higher local elevation and at larger hydrological distance from the main rivers: geographical distance to the river appears a poor predictor of lake isolation. Although year-to-year fish catches in the floodplain were significantly larger with larger flood volumes in the floodplain, they were not in the main river, suggesting that mechanisms that increase catch, such as increased floodplain access or increased somatic growth, are stimulated by flooding in the floodplain, but not in the river. Fish species that profit from flooding belong to different feeding guilds, suggesting that all trophic levels may benefit from flooding. We found indications that the ecological functioning of floodplains is not limited to its temporary availability as habitat. Refugia can be present within the floodplain itself, which should be considered in the management of large rivers and their floodplain.     

Contribution of microspatial factors to benthic diatom communities

Large rivers are worldwide under severe pressure and there is a lack of information on large river restoration. The present paper represents a meta-analysis of available data on river rehabilitation projects performed at the Austrian Danube River consisting of six rehabilitation projects addressing 19 sites. The overall goal was to analyse the response of fish assemblages to different rehabilitation types based on (1) morphological type (“Instream Habitat Enhancement”, “backwater Enhancement”, “extended Enhancement”), (2) length of rehabilitation measure (3) time after construction (4) applied monitoring design. Biological metrics evaluated included number of fish species and relative density, habitat guilds and Leitbild species. In total, number of species increases by 55% comparing rehabilitated with unrestored sites. The number of species of all habitat guilds is higher after rehabilitation. The proportion of rheophilic species increased and the community evolved toward a more type-specific community, according to the Leitbild. Significant differences between measure types were not detected. The rehabilitation success depends mainly on its spatial extent. Highest positive response of number of rheophilic species is achieved by a length >3.9 km. The results show that habitat rehabilitation of large rivers is effective if the spatial extent of the measure is in accordance with river size.     

Communities at the extreme: Aquatic food webs in desert landscapes

Studying food webs across contrasting abiotic conditions is an important tool in understanding how environmental variability impacts community structure and ecosystem dynamics. The study of extreme environments provides insight into community‐wide level responses to environmental pressures with relevance to the future management of aquatic ecosystems. In the western Lake Eyre Basin of arid Australia, there are two characteristic and contrasting aquatic habitats: springs and rivers. Permanent isolated Great Artesian Basin springs represent hydrologically persistent environments in an arid desert landscape. In contrast, hydrologically variable river waterholes are ephemeral in space and time. We comprehensively sampled aquatic assemblages in contrasting ecosystem types to assess patterns in community composition and to quantify food web attributes with stable isotopes. Springs and rivers were found to have markedly different invertebrate communities, with rivers dominated by more dispersive species and springs associated with species that show high local endemism. Qualitative assessment of basal resources shows autochthonous carbon appears to be a key basal resource in both types of habitat, although the particular sources differed between habitats. Food‐web variables such as trophic length, trophic breadth, and community isotopic niche size were relatively similar in the two habitat types. The basis for the similarity in food‐web structure despite differences in community composition appears to be broader isotopic niches for predatory invertebrates and fish in springs as compared with rivers. In contrast to published theory, our findings suggest that the food webs of the hydrologically variable river sites may show less dietary generalization and more compact food‐web modules than in springs.     

Abundance trends for river macroinvertebrates vary across taxa,trophic group and river typology

There is mounting evidence that terrestrial arthropods are declining rapidly in many areas of the world. It is unclear whether freshwater invertebrates, which are key providers of ecosystem services, are also declining. We addressed this question by analysing a long-term dataset of macroinvertebrate abundance collected from 2002 to 2019 across 5009 sampling sites in English rivers. Patterns varied markedly across taxonomic groups. Within trophic groups we detected increases in the abundance of carnivores by 19% and herbivores by 14.8%, while we estimated decomposers have declined by 21.7% in abundance since 2002. We also found heterogeneity in trends across rivers belonging to different typologies based on geological dominance and catchment altitude, with organic lowland rivers having generally higher rates of increase in abundance across taxa and trophic groups, with siliceous lowland rivers having the most declines. Our results reveal a complex picture of change in freshwater macroinvertebrate abundance between taxonomic groups, trophic levels and river typologies. Our analysis helps with identifying priority regions for action on potential environmental stressors where we discover macroinvertebrate abundance declines.     

Indicators quantifying small pelagic fish interactions: application using a trophic model of the southern Benguela ecosystem

Three indicators quantifying interactions between species are developed for an upwelling system to provide useful measures for the comparison of marine ecosystem structure and function. Small pelagic fish are dominant in upwelling systems, and by definition, they are pivotal in a wasp-waist upwelling system. The indicator of interaction strength (IS) quantifies the effect that a change in biomass of one group has on abundance of other groups. The functional impact (FI) indicator quantifies the trophic impacts of species on their own and other functional groups or feeding guilds. The trophic replacement (TR) indicator quantifies the trophic similarity between a species that is removed from an ecosystem and other species in that ecosystem, i.e. it quantifies the ability of one group to trophically replace another. A trophic model of the southern Benguela ecosystem is used as an example for the application of the indicators. The strong similarities in trophic functioning of the southern Benguela ecosystem in the anchovy-dominated system of the 1980s, and the 1990s when there was a shift towards greater sardine abundance, are explained by the mutual trophic replacement abilities of anchovy and sardine. Differences between the proposed indicators and mixed trophic impact assessment are highlighted, mainly resulting from the static versus dynamic nature of the models upon which they are based. Trophic indicators such as those presented here, together with other kinds of ecosystem indicators, may assist in defining operational frameworks for ecosystem-based fisheries management.