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.     

Fish mediate high food web connectivity in the lower reaches of a tropical floodplain river

High levels of hydrological connectivity during seasonal flooding provide significant opportunities for movements of fish between rivers and their floodplains, estuaries and the sea, possibly mediating food web subsidies among habitats. To determine the degree of utilisation of food sources from different habitats in a tropical river with a short floodplain inundation duration (~2 months), stable isotope ratios in fishes and their available food were measured from three habitats (inundated floodplain, dry season freshwater, coastal marine) in the lower reaches of the Mitchell River, Queensland (Australia). Floodplain food sources constituted the majority of the diet of large-bodied fishes (barramundi Lates calcarifer, catfish Neoarius graeffei) captured on the floodplain in the wet season and for gonadal tissues of a common herbivorous fish (gizzard shad Nematalosa come), the latter suggesting that critical reproductive phases are fuelled by floodplain production. Floodplain food sources also subsidised barramundi from the recreational fishery in adjacent coastal and estuarine areas, and the broader fish community from a freshwater lagoon. These findings highlight the importance of the floodplain in supporting the production of large fishes in spite of the episodic nature and relatively short duration of inundation compared to large river floodplains of humid tropical regions. They also illustrate the high degree of food web connectivity mediated by mobile fish in this system in the absence of human modification, and point to the potential consequences of water resource development that may reduce or eliminate hydrological connectivity between the river and its floodplain.     

Hydrologic Connectivity Affects Fish Assemblage Structure,Diversity, and Ecological Traits in the Unregulated Gambia River,West Africa

The Gambia River of West Africa is a large unobstructed river, characterized by a natural flow regime and lateral connectivity across its floodplain. Construction of a major dam, however, is planned. We compared patterns of fish diversity, habitat use, assemblage structure, and the distribution of trophic position and body morphology in riverine and floodplain habitats in Niokolo Koba National Park, located downstream of the planned dam site. A total of 49 fish species were captured, revealing a lognormal distribution as expected for species‐rich assemblages. Fish species exhibited a range of habitat use patterns, from generalist to highly habitat‐specific, and appeared to migrate laterally among habitats between seasons. Species richness was homogenous among habitats in the wet season yet appeared to increase with isolation from the main river in the dry season. Fish assemblage structure was best explained by the interaction between habitat type and season, underlining the importance of the natural flow regime and lateral connectivity among floodplain habitats. The abundance of fishes having elongate bodies increased with isolation from the main channel in the wet season only. The distribution of fishes having compressed cross‐sectional morphology decreased with isolation from the main channel in the dry season only. These patterns of trait distribution support the conclusion that variation in hydrologic connectivity structures the fish assemblage. Our results suggest that altered flow regimes and loss of floodplain habitats after damming could lead to both decreased taxonomic and functional diversity of the fish assemblage.     

Comparing the fish assemblages and food‐web structures of large floodplain rivers

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Diel variation in food intake and diet composition of three native fish species in floodplain lagoons of the Macintyre River,Australia

The feeding periodicity of three species of fishes in floodplain lagoons of the Macintyre River varied during 24 h cycles, however, peak feeding activity (around midday and late afternoon) was similar among the species studied, despite differences in their trophic position. In contrast, sampling season and site seemed to have a stronger effect on the composition of the diet than time of day. These findings have implications for studies of trophic dynamics and food web structure in floodplain lagoons.     

Fish larvae, growth and biomass relationships in an Australian arid zone river: links between floodplains and waterholes

1. Floodplain inundation provides many benefits to fish assemblages of floodplain river systems, particularly those with a predictable annual flood pulse that drives yearly peaks in fish production. In arid‐zone rivers, hydrological patterns are highly variable and the influence of irregular floods on fish production and floodplain energy subsidies may be less clear‐cut. To investigate the importance of floodplain inundation to a dryland river fish assemblage, we sampled fish life stages on the floodplain of Cooper Creek, an Australian arid‐zone river. Sampling was focused around Windorah during a major flood in January 2004 and in isolated waterholes in March 2004 following flood drawdown. 2. Of the 12 native species known to occur in this region, 11 were present on the floodplain, and all were represented by at least two of three life‐stages – larvae, juveniles or adult fish. Late stage larvae of six fish species were found on the floodplain. There were site‐specific differences in larval species assemblages, individual species abundances and larval distribution patterns among floodplain sites. 3. Significant growth was evident on the floodplain, particularly by larval and juvenile fish, reflecting the combination of high water temperatures and shallow, food rich habitats provided by the relatively flat floodplain. 4. Low variation in biomass, species richness and presence/absence of juvenile and adult fish across four floodplain sites indicates consistently high fish productivity across an extensive area. 5. Similarities and differences in fish biomass between the floodplain and isolated post‐flood waterholes suggest high rates of biomass transfer (involving the most abundant species) into local waterholes and, potentially, biomass transfer by some species to other waterholes in the catchment during floodplain inundation and after floods recede. 6. The high concentration of fish on this shallow floodplain suggests it could be a key area of high fish production that drives a significant proportion of waterhole productivity in the vicinity. The Windorah floodplain provides favourable conditions necessary for the spawning of some species and juvenile recruitment of the majority of species. It is also appears to be a significant conduit for the movements of fish that underpin high genetic similarity, hence population mixing, of many species throughout the Cooper Creek catchment. The high floodplain fish production in turn provides a significant energy subsidy to waterholes after floodwaters recede. 7. The identification of key sites of high fish production, such as the Windorah floodplain, may be important from a conservation perspective. Key management principles should be: maintenance of the natural flooding regime; identification of the most productive floodplain areas; and maintenance of their connectivity to anastomosing river channels and the remnant aquatic habitats that ultimately sustain this fish assemblage through long‐term dry/drought and flood cycles.     

Effects of seasonality and fish movement on tropical river food webs

Tropical rivers and their associated floodplain habitats are dynamic habitat mosaics to which fishes are challenged to respond in an adaptive manner. Migratory fishes create linkages among food webs that are partitioned along a nested hierarchy of spatial scales. Such linkages are examined across a hierarchy of spatio-temporal scales, ranging from small streams to entire drainage basins, for rivers in South America and Africa. Migratory herbivorous fishes originating from eutrophic, productive ecosystems may subsidize resident predators of oligotrophic river ecosystems, which may result in cascading direct and indirect Effects on other species in local food webs. Successful management of many of the most important stocks of tropical river fishes requires conceptual models of how fish movement influences food web structure and dynamics.     

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.     

Comparative use of side and main channels by small‐bodied fish in a large,unimpounded river

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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.     

Flood pulse trophic dynamics of larval fishes in a restored arid-land, river-floodplain, Middle Rio Grande, Los Lunas, New Mexico

Rio Grande water is intensively managed and regulated by international and interstate compacts, Native American treaties, local water rights, and federal, state, and local agencies. Legislation and engineering projects in the early twentieth century brought about water impoundment projects and channelization of the Rio Grande which led to the eventual loss of floodplain habitats. In particular, current water management practices in the Middle Rio Grande (MRG) have altered the natural flood regime altering the riparian community and floodplain dynamics which may be causing the demise of many fish species by altering food web processes. The Rio Grande silvery minnow (Hybognathus amarus), a federally endangered species, has been classified as an herbivore, detritivore, or carnivore. During low flow conditions H. amarus is primarily an algivore; however, during flood conditions, hydrodynamic scouring reduces or eliminates benthic algal food sources. The objective of this study was to identify and characterize food resources and trophic interactions for H. amarus on a restored floodplain during an extended flood-pulse release from reservoirs using stable isotope analyses (δ¹³C and δ¹⁵N) and paleolimnology techniques. Results from stable isotope ratios indicate that H. amarus obtained carbon primarily from chironomids while aquatic invertebrates (including chironomids) obtained their carbon from macrophytes. Results from the GLIMMIX procedure indicate that the range of isotopic signatures for prey items was much broader at parallel habitats (i.e. floodplain flow parallel to main stem flow) than perpendicular (i.e. floodplain flow perpendicular to main stem flow) or leeward habitats (i.e. leeward sides of island where flow was near zero) indicating a wider selection of food resources. This study suggests that increased duration of floodplain inundation in the MRG provides vital habitats for spawning, nursery, and recruitment of threatened and endangered fish species. A combination of allochthonous and autochthonous resources best describes the nutrient and energy transfers for the Los Lunas, NM restored floodplain.     

Ephemeral floodplain habitats provide best growth conditions for juvenile Chinook salmon in a California river

We reared juvenile Chinook salmon for two consecutive flood seasons within various habitats of the Cosumnes River and its floodplain to compare fish growth in river and floodplain habitats. Fish were placed in enclosures during times when wild salmon would naturally be rearing in floodplain habitats. We found significant differences in growth rates between salmon reared in floodplain and river enclosures. Salmon reared in seasonally inundated habitats with annual terrestrial vegetation experienced higher growth rates than those reared in a perennial pond on the floodplain. Growth of fish in the non-tidal river upstream of the floodplain varied with flow in the river. When flows were high, there was little growth and high mortality, but when the flows were low and clear, the fish grew rapidly. Fish displayed very poor growth in tidally influenced river habitat below the floodplain, a habitat type to which juveniles are commonly displaced during high flow events due to a lack of channel complexity in the main-stem river. Overall, ephemeral floodplain habitats supported higher growth rates for juvenile Chinook salmon than more permanent habitats in either the floodplain or river. Variable responses in both growth and mortality, however, indicate the importance of providing habitat complexity for juvenile salmon in floodplain reaches of streams, so fish can find optimal places for rearing under different flow conditions.     

Variability of fish diets between dry and flood periods in an arid zone floodplain river

The fish assemblages of an arid zone floodplain river, Cooper Creek, Queensland, Australia, were sampled during two dry periods in isolated waterholes and on the inundated floodplain during the early and late phase of a major flood event. Diets were described for nine native species and compared within and between dry and flood periods. In the dry season, when fishes were restricted to waterholes, diets were characteristically simple with narrow diet breadths. Movement onto the floodplain during flooding clearly increased feeding opportunities, with greater diet breadths evident in all species. Despite obvious potential for terrestrial inputs, diets tended to be dominated by aquatic resources in both the waterholes and on the floodplain. Stomach fullness, however, varied little between dry season waterhole and floodplain samples. Fishes appeared to feed on potentially lower value resources such as detritus and calanoid copepods during the dry season, when waterholes were isolated and food resources were limited. They were then able to capitalize on the 'boom' of aquatic production and more diverse food resources associated with episodic flood events.     

Piscivore Responses to Enhancement of the Channelized Kissimmee River,Florida, U.S.A.

Evaluation of the success of ecosystem restoration projects requires identification of appropriate ecological metrics. Comparison of reconstructed food webs (or subsets thereof) from restored and non‐restored habitats may be a valuable tool to evaluate restoration success because food webs help identify critical predator–prey relationships, keystone species, relative importance of direct and indirect trophic interactions, and other aspects of ecological function. We compared the diets of apex predatory fishes collected from enhanced and non‐enhanced portions of the channelized Kissimmee River, Florida, USA to determine whether food web structure responded to experimental hydrologic manipulations. Diets were reconstructed for black crappie (Pomoxis nigromaculatus), bowfin (Amia calva), chain pickerel (Esox niger), Florida gar (Lepisosteus platyrhincus), largemouth bass (Micropterus salmoides), and warmouth (Lepomis gulosus) collected from enhanced and non‐enhanced portions of the Kissimmee River. Prey eaten by apex predatory fishes in the enhanced portion of the Kissimmee River were quantitatively and qualitatively different from prey eaten in non‐enhanced portions of the river. Predators in the enhanced portion of the river had fewer empty stomachs, more prey items per individual, more prey types per individual, more fish prey per individual, greater overall richness of prey, and a multivariate suite of prey distinct from predators in non‐enhanced portions of the river. Results from hydrologic manipulations suggest that large‐scale restoration of hydrologic linkages between the main channel and floodplain habitats will positively affect food web structure and ecosystem function in the Kissimmee River.     

Spatial Variation in δ15N and δ13C Isotopes in the San Juan River, New Mexico and Utah: Implications for the Conservation of Native Fishes

Synopsis Spatial patterns of resource use by small-bodied fishes in the San Juan River were examined using stable isotopes. Using δ¹⁵N of fishes as an index of trophic position, our data suggest both native and non-native fishes primarily consumed macro-invertebrates. The δ¹³C of these fishes further suggested a detritus-based food web, from which most species fed on chironomids in low-velocity habitats. A two-way ANOVA revealed a significant interaction between trophic level of fish species and longitudinal position in the river. This interaction was primarily attributed to a decline in trophic level of non-native red shiner Cyprinella lutrensis, relative to other species, in upstream reaches of the river. In addition, ANCOVA results suggest trophic position of fishes was dependent on channel type (primary vs. secondary), as there was less variability in resource use in secondary channels. These data provided a spatial framework of trophic interactions that can be used to predict the outcome of management actions. Overall, we confirmed high overlap in resource used between native and non-native fishes. However, spatial variation in trophic interactions both longitudinally and laterally in the river present a challenge to resource managers attempting to managing entire river systems.     

Physical structure of habitat network differently affects migration patterns of native and invasive fishes in Lake Biwa and its tributary lagoons: stable isotope approach

A good understanding of how migratory animals use their habitat network is expected to provide important insights for the prediction of population dynamics at both local and regional scales. We focused on how the physical structure of a habitat network could affect fish migration between Lake Biwa and its tributary lagoons. Although the lagoons provide suitable breeding and nursery grounds for native fishes, it is a matter of concern that they can also be a hotbed of invasive exotic fishes. Here, we assessed the migration patterns of native crucian carps (Carassius spp.) and exotic largemouth bass (Micropterus salmoides) and bluegill sunfish (Lepomis macrochirus) using their carbon and nitrogen stable isotopes as migration tracers. As there were marked differences in the carbon isotope ratios of basal food webs between the main lake and its tributary lagoons, stable isotopic signatures of individual fishes collected from each lagoon enabled us to judge whether they were residents of the lagoon or recent immigrants from the main lake. The analysis revealed that native and invasive fishes showed different migration patterns across a variety of lagoons. Exotic fishes frequently immigrated from the main lake to the lagoon as the distance of the channel connecting these two habitats was short. For native crucian carps, in contrast, their migrations were unaffected by the channel distance but were promoted by narrow channels. Physical barriers of weirs and dense vegetation within the channel obstructed their migrations. Such ecological information on migration behavior will be vital to plan designs for habitat restoration to conserve native fishes.     

Flood regime,dam regulation and fish in the Upper Paraná River: effects on assemblage attributes,reproduction and recruitment

The flood regime is the most important force determining seasonality in neotropical rivers. In the Upper Paran River floodplain, it is the primary factor influencing biological processes. The aim of this paper is to summarize information on the influence of dam-controlled floods on some fish assemblage attributes, reproduction and recruitment in the Upper Paran River floodplain, providing preliminary guidelines for dam operation upstream. Fish were collected in different habitats of the Upper Paran River floodplain (river, channels and lagoons) in the period from 1986 to 2001. The high water period in the Paran River usually occurs from November/December to April/May. Annual variation in the hydrograph affects species with distinct life history strategies differently, and influences the composition and structure of fish assemblages. Large floods were associated with higher species richness. Frequencies of individuals with ripe and partially spent gonads, which indicate spawning, were higher during the period of increasing water level. Dependence on floods seems to be lowest in sedentary species that develop parental care, and highest in large migratory species that spawn in the upper stretches of the basin and use flooded areas as nurseries. Migratory fishes were favored by annual floods that lasted more than 75 days, with longer floods yielding larger populations. The occurrence of high water levels at the beginning of summer is fundamental to the spawning success of migratory species. However, the flood may be less important for recruitment of juveniles if it is of short duration. Dam operation upstream (releasing more water during the raining season) has potential to promote greater floods with appropriate duration improving recruitment, particularly for migratory species.     

The influence of floodplain habitat on the quantity and quality of riverine phytoplankton carbon produced during the flood season in San Francisco Estuary

Primary productivity, community respiration, chlorophyll a concentration, phytoplankton species composition, and environmental factors were compared in the Yolo Bypass floodplain and adjacent Sacramento River in order to determine if passage of Sacramento River through floodplain habitat enhanced the quantity and quality of phytoplankton carbon available to the aquatic food web and how primary productivity and phytoplankton species composition in these habitats were affected by environmental conditions during the flood season. Greater net primary productivity of Sacramento River water in the floodplain than the main river channel was associated with more frequent autotrophy and a higher P:R ratio, chlorophyll a concentration, and phytoplankton growth efficiency (α^B). Total irradiance and water temperature in the euphotic zone were positively correlated with net primary productivity in winter and early spring but negatively correlated with net primary productivity in the late spring and early summer in the floodplain. In contrast, net primary productivity was correlated with chlorophyll a concentration and streamflow in the Sacramento River. The flood pulse cycle was important for floodplain production because it facilitated the accumulation of chlorophyll a and wide diameter diatom and green algal cells during the drain phase. High chlorophyll a concentration and diatom and green algal biomass enabled the floodplain to export 14–37% of the combined floodplain plus river load of total, diatom and green algal biomass and wide diameter cells to the estuary downstream, even though it had only 3% of the river streamflow. The study suggested the quantity and quality of riverine phytoplankton biomass available to the aquatic food web could be enhanced by passing river water through a floodplain during the flood season.     

Habitat and resource use by larval and juvenile fishes in an arid-land river (Rio Grande, New Mexico)

1. Studies of mesic temperate and tropical rivers suggest an important role for floodplain habitats as nursery areas for larval and juvenile fishes. In arid‐land rivers the extent and duration of flooding is diminished and habitats and resources used by larval fishes are poorly known. Our study documented habitat and resource use of larval fishes in the Rio Grande, New Mexico, an arid‐land river. 2. Spatial and temporal distribution of larval and juvenile fishes and their inferred microhabitat preferences were studied during spring, summer and autumn, 2003. Stable carbon (¹³C : ¹²C) and nitrogen (¹⁵N : ¹⁴N) isotope ratios were measured to identify nutrient sources and characterise trophic positions of young‐of‐year fishes in this system. 3. Some fishes recruited during high flows (in spring), whereas others recruited during low‐flow periods in late summer. Regardless of the timing of reproduction, microhabitats with lower current velocity and higher temperature appeared to serve as vital nursery grounds for Rio Grande fishes. Ephemeral backwaters and disconnected side channels held the highest abundance and diversity of larvae and juveniles. 4. Stable isotope analyses revealed that fish larvae obtained carbon predominately from algal production in early summer, but used organic carbon derived from emergent macrophytes as river discharge decreased in mid‐summer. This shift may have been facilitated by microinvertebrate prey that grazed down edible algae and then switched to macrophytes in mid‐summer. Nitrogen isotope ratios did not differ among species or early life stages, suggesting that larval and juvenile fishes use similar food resources, especially when restricted to isolated pools in summer.     

Flood disturbance affects macroinvertebrate food chain length in an alluvial river floodplain

1. Characterising food-web responses to environmental factors could greatly improve our understanding of environment-biota relationships, and especially in floodplains where trophic interactions can be particularly important during phases of hydrological disconnection. The effects of floodplain hydrology and environmental attributes on structural aspects of biotic assemblages have been extensively studied, but responses at the functional level remain largely unknown.
2. Here, we characterised a central aspect of food-web architecture, the food chain length, as the maximum trophic position within 24 macroinvertebrate communities of parafluvial habitats in the Maggia river floodplain, in Switzerland. We investigated how the food chain length changed with different levels of habitat size, primary productivity and disturbance, the three factors potentially affecting food chain length in both theoretical and empirical studies.
3. We found that food chain length was lower in frequently flooded habitats and immediately after a flood. We also showed that trophic omnivory, where predators fed at lower trophic levels after flooding, and in more frequently flooded habitats, may explain these changes.
4. These findings show that trophic omnivory may explain how predators resist disturbance and are maintained in highly dynamic landscapes. More importantly, given that trophic omnivory may overall weaken trophic linkages and thus increase food-web stability, this suggests that it could be a key mechanism in sustaining biodiversity in river floodplains.