Flood disturbance, algal productivity, and interannual variation in food chain length

The length of a river food chain changed from year to year, shifting with the hydrologic regime. During drought years, grazers suppressed algae across a nutrient gradient, while predators were functionally unimportant. Following flood disturbance, predators suppressed grazers, releasing algae. These results suggest that hydrologic regime, rather than productivity, determines the functional length of this river food chain. Within years, algae and grazer biomass responded to an experimental productivity gradient in patterns predicted by simple trophic models that assume efficient energy transfer. Understanding differences among species within trophic levels, however, was crucial in delineating the controlling interactions.     

Tracing Mississippi River influences in estuarine food webs of coastal Louisiana

The Breton Sound estuary in southern Louisiana receives large amounts of Mississippi River water via a controlled diversion structure at the upstream end of the estuary. We used stable isotopes to trace spatial and seasonal responses of the downstream food web to winter and spring introductions of river water. Analysis of δ¹³C, δ¹⁵N, and δ³⁴S in the common local consumers such as grass shrimp (Palaemonetes sp.), barnacles (Balanus sp.), and small plankton-feeding fish (bay anchovies, Anchoa mitchilli) showed that the diversion was associated with two of the five major source regimes that were supporting food webs: a river regime near the diversion and a river-influenced productive marsh regime farther away from the diversion. Mixing models identified a third river-influenced source regime at the marine end of the estuary where major natural discharge from the Bird’s Foot Delta wraps around into estuarine waters. The remaining two source regimes represented typical estuarine conditions: local freshwater sources especially from precipitation and a brackish source regime representing higher salinity marine influences. Overall, the Mississippi River diversion accounted for 75% of food web support in the upper estuary and 25% in the middle estuary, with influence strongest along known flow pathways and closest to the diversion. Isotopes also traced seasonal changes in river contributions, and indicated increased plant community productivity along the major flow path of diversion water. In the Breton Sound estuary, bottom–up forcing of food webs is strongly linked to river introductions and discharge, occurring in spatial and temporal patterns predictable from known river input regimes and known hydrologic circulation patterns.     

Food webs in river networks

Food webs and river drainages are both hierarchical networks and complex adaptive systems. How does living within the second affect the first? Longitudinal gradients in productivity, disturbance regimes and habitat structure down rivers have long interested ecologists, but their effects on food web structure and dynamics are just beginning to be explored. Even less is known about how network structure per se influences river and riparian food webs and their members. We offer some preliminary observations and hypotheses about these interactions, emphasizing observations on upstream–downstream changes in food web structure and controls, and introducing some ideas and predictions about the unexplored question of food web responses to some of the network properties of river drainages.     

Avian Community Responses to Variability in River Hydrology

River flow is a major driver of morphological structure and community dynamics in riverine-floodplain ecosystems. Flow influences in-stream communities through changes in water velocity, depth, temperature, turbidity and nutrient fluxes, and perturbations in the organisation of lower trophic levels are cascaded through the food web, resulting in shifts in food availability for consumer species. River birds are sensitive to spatial and phenological mismatches with aquatic prey following flow disturbances; however, the role of flow as a determinant of riparian ecological structure remains poorly known. This knowledge is crucial to help to predict if, and how, riparian communities will be influenced by climate-induced changes in river flow characterised by more extreme high (i.e. flood) and/or low (i.e. drought) flow events. Here, we combine national-scale datasets of river bird surveys and river flow archives to understand how hydrological disturbance has affected the distribution of riparian species at higher trophic levels. Data were analysed for 71 river locations using a Generalized Additive Model framework and a model averaging procedure. Species had complex but biologically interpretable associations with hydrological indices, with species’ responses consistent with their ecology, indicating that hydrological-disturbance has implications for higher trophic levels in riparian food webs. Our quantitative analysis of river flow-bird relationships demonstrates the potential vulnerability of riparian species to the impacts of changing flow variability and represents an important contribution in helping to understand how bird communities might respond to a climate change-induced increase in the intensity of floods and droughts. Moreover, the success in relating parameters of river flow variability to species’ distributions highlights the need to include river flow data in climate change impact models of species’ distributions.     

Seasonal and inter-annual variations of gas exchange in thirteen woody species along a climatic gradient in the Mediterranean island of Mallorca

We studied the influence of summer drought and winter temperatures on seasonal and spatial variations of light-saturated net photosynthesis and stomatal conductance in Mediterranean woody species. We measured variations in leaf gas exchange over 3 years in 13 Mediterranean trees and shrubs, located at four different sites along a climate gradient of temperature and precipitation in the island of Mallorca (West Mediterranean basin). Net photosynthesis and stomatal conductance were at a maximum during spring, autumn or winter and at a minimum during summer in most sites, species and years. Nevertheless, important spatial, temporal and species-specific variations were observed. Summer drought limitation to gas exchange was greatest in the dry part of the transect, where many species showed their maximum gas exchange rate during winter. In contrast, winter temperatures limited gas exchange of many species at the wet and cool end of the transect, while summer depression of gas exchange was shorter and less pronounced. These results suggest that the effect on carbon fixation and productivity by the predicted future increase of aridity in the Mediterranean basin will depend on whether gas exchange is mostly limited by summer drought or by low winter temperatures.     

Relationship between flow regime and fish abundances in a gravel‐bed river, New Zealand

The key elements of the flow regime of the Waipara River on the east coast of New Zealand, that affected fish abundances were the timing of floods and the magnitude and duration of low flows. Generally, fish abundances were highest in early summer, and lowest at the beginning of winter. Spring floods opened the river mouth, allowing recruitment of diadromous fish species, and non‐diadromous fish species spawned after the floods in spring or early summer. Reductions in fish abundances over summer and autumn were consistent with the magnitude and duration of low flows, with significant reductions in the year of lowest flow and little change in abundance in the year when low flows were highest. Variations in fish abundances during periods of low flow were consistent with the amount of instream habitat available, such that abundances of species with high velocity preferences decreased during periods of low flow, whereas abundances of species with low velocity preferences increased.     

Is water temperature an adequate predictor of recruitment success in cyprinid fish populations in lowland rivers?

SUMMARY 1. Higher than average ambient water temperature in the first year of life may be responsible for strong cohorts of adult cyprinid fish. Whilst temperature explains much of the variation in year‐class strength (YCS), however, it is not the only influential factor as high temperature does not inevitably yield strong year‐classes. Furthermore, years in which a strong year‐class is prevalent in one species do not necessarily result in strong year‐classes in other coexisting species, suggesting other biotic and abiotic factors are important in regulating recruitment success. 2. The relationships between water temperature, river discharge, the position of the Gulf Stream, 0‐group fish growth and recruitment success (YCS) were examined in three cyprinid fish species in an English lowland river, using a 15‐year data set. 3. Mean length of 0‐group fish at the end of the summer was positively correlated with water temperature (cumulative degree‐days >12 °C) and negatively correlated with river discharge (cumulative discharge‐days above basal discharge rate). Water temperature was negatively correlated with river discharge. 4. YCS was positively correlated with mean 0‐group fish length at the end of the summer and with the position of the North Wall of the Gulf Stream. 5. ’Critical periods’ (i.e. periods in the first summer of life when fish may be more susceptible to increases in river discharge) were difficult to discern because of interannual variations in river discharge relative to the timing of fish hatching. YCS of roach and chub was most strongly correlated with discharge in the period from June to September inclusive, while YCS of dace was most significantly correlated with discharge in August. 6. River discharge (rather than water temperature) may be the key factor in determining YCS, either directly (through discharge‐induced mortality) or indirectly (via reduced growth at lower water temperatures, discharge‐associated increases in energy expenditure or reduced food availability). It could be that, in effect, water temperature determines potential YCS while discharge determines realised YCS.     

Productivity,Disturbance and Ecosystem Size Have No Influence on Food Chain Length in Seasonally Connected Rivers

The food web is one of the oldest and most central organising concepts in ecology and for decades, food chain length has been hypothesised to be controlled by productivity, disturbance, and/or ecosystem size; each of which may be mediated by the functional trophic role of the top predator. We characterised aquatic food webs using carbon and nitrogen stable isotopes from 66 river and floodplain sites across the wet-dry tropics of northern Australia to determine the relative importance of productivity (indicated by nutrient concentrations), disturbance (indicated by hydrological isolation) and ecosystem size, and how they may be affected by food web architecture. We show that variation in food chain length was unrelated to these classic environmental determinants, and unrelated to the trophic role of the top predator. This finding is a striking exception to the literature and is the first published example of food chain length being unaffected by any of these determinants. We suggest the distinctive seasonal hydrology of northern Australia allows the movement of fish predators, linking isolated food webs and potentially creating a regional food web that overrides local effects of productivity, disturbance and ecosystem size. This finding supports ecological theory suggesting that mobile consumers promote more stable food webs. It also illustrates how food webs, and energy transfer, may function in the absence of the human modifications to landscape hydrological connectivity that are ubiquitous in more populated regions.     

Water discharge affects Atlantic salmon Salmo salar smolt production: a 27 year study in the River Orkla,Norway

A model that explains 48% of the annual variation in Atlantic salmon Salmo salar smolt production in the River Orkla, Norway, has been established. This variation could be explained by egg deposition, minimum daily discharge during the previous winter and minimum weekly discharge during the summer 3 years before smolt migration. All coefficients in the model were positive, which indicates that more eggs and higher minimum discharge levels during the winter before smolt migration and the summer after hatching benefit smolt production. Hence, when the spawning target of the river is reached, the minimum levels of river discharge, in both winter and summer, are the main bottlenecks for the parr survival, and hence for smolt production. The River Orkla was developed for hydropower production in the early 1980s by the construction of four reservoirs upstream of the river stretch accessible to S. salar. Although no water has been removed from the catchment, the dynamics of water flow has been altered, mainly by increasing discharges during winter and reducing spring floods. In spite of the higher than natural winter discharges, minimum winter discharge is still a determinant of smolt production. Hence, in regulated rivers, the maintenance of discharges to ensure that they are as high as possible during dry periods is an important means of securing high S. salar smolt production.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Landscape-Scale Hydrologic Characteristics Differentiate Patterns of Carbon Flow in Large-River Food Webs

Efforts to conserve, restore, or otherwise manage large rivers and the services they provide are hindered by limited understanding of the functional dynamics of these systems. This shortcoming is especially evident with regard to trophic structure and energy flow. We used natural abundances of carbon and nitrogen isotopes to examine patterns of material flow in ten large-river food webs characterized by different landscape-scale hydrologic characteristics (low-gradient river, high-gradient river, river stretches downstream of reservoirs, and reservoirs), and tested predictions from three ecosystem concepts commonly applied to large-rivers: The River Continuum Concept, The Flood Pulse Concept and the Riverine Productivity Model. Carbon derived from aquatic C₃ plants and phytoplankton were the dominant energy sources supporting secondary consumers across the ten large-river food webs examined, but relative contributions differed significantly among landscape types. For low-gradient river food webs, aquatic C₃ plants were the principal carbon source, contributing as much as 80% of carbon assimilated by top consumers, with phytoplankton secondarily important. The estimated relative importance of phytoplankton was greatest for food webs of reservoirs and river stretches downriver from impoundments, although aquatic C₃ plants contributed similar amounts in both landscape types. Highest 99th percentile source contribution estimates for C₄ plants and filamentous algae (both approximately 40%) were observed for high-gradient river food webs. Our results for low-gradient rivers supported predictions of the Flood Pulse Concept, whereas results for the three other landscape types supported the Riverine Productivity Model to varying degrees. Incorporation of landscape-scale hydrologic or geomorphic characteristics, such as river slope or floodplain width, may promote integration of fluvial ecosystem concepts. Expanding these models to include hydrologically impacted landscapes should lead to a more holistic understanding of ecosystem processes in large-river systems.     

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.     

Phenology of Mediterranean woody plants from NE Spain: Synchrony,seasonality, and relationships among phenophases

Plant phenological studies in the Mediterranean have traditionally focused on the limits that summer drought exerts over plant seasonal behaviour. However, Mediterranean areas also occur across extensive regions under cooler climates. Here, we analyse phenological data of Mediterranean woody species from winter-cold sites to address the following questions: what is the most limiting period of the year for phenological development? How, and how synchronously, do different phenophases adjust to the bimodal limitation of summer drought and winter cold? How ample is the suit of phenological strategies that are successful for Mediterranean phanaerophytes from cool areas?     

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.     

Assessment of river health: accounting for perturbation pathways in physical and ecological space

1. A full understanding of changes to river ecosystem structure and functioning along the continuum from relatively pristine to profoundly perturbed requires knowledge of physical, chemical and ecological properties at many spatial and temporal scales. Perturbations may span broad spatio-temporal scales or be spatially and temporally discrete. 2. The relevant scale for study depends on the manner in which perturbations move through two kinds of pathways – through the physical space of the catchment hierarchy and through the ecological space of river food webs. Different kinds of perturbation (e.g. inputs of sediment, nitrogen or phosphorus, changes to shading or discharge) vary in the degree and manner in which they are propagated downstream and through food webs. 3. The fundamental importance of disturbance regimes and refugia to river health is now clear and managers need to take this into account when devising river management schemes. 4. A comparison of the physics, chemistry and ecology of streams in catchments of native tussock and exotic pasture grassland in New Zealand serves to emphasize (i) the extreme complexity of interacting factors resulting from land use change, (ii) the importance of disturbance regime (not encapsulated in biotic indices) and biogeographic patterns (usually unmeasured) in determining ecosystem structure and functioning, and (iii) the lack of concordance between any single index of health and various fundamental features of ecosystem functioning. 5. Practical considerations limit most evaluations of river health to a small suite of indices, though it is important that researchers continue to evaluate the spatial, temporal and biological limitations of these indices. The vagaries of history and geography (extinction and colonization trajectories in relation to natural disturbance regimes) and the complexity of interacting physical, chemical and ecological responses to perturbation suggest that multi-scale, multi-temporal studies of river function offer the best opportunity to evaluate river health.     

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.     

Summer drought disturbance on plant detritus decomposition processes in three River Tirso (Sardinia, Italy) sub-basins

Here, we study the variation patterns of detritus decomposition along a river continuum in a Mediterranean type river basin, and the influence of summer drought disturbance on this pattern. The study was carried out in three 4th order sub-basins (hereafter referred to as Assolo, Laconi and Olzai) of the river Tirso basin (Sardinia, ITALY), with one study site per stream order in each sub-basin (4 stream orders × 3 sub-basins = 12 study sites). The three sub-basins were selected according to their exposure to summer drought, Assolo being the most exposed and Laconi the least exposed. Reed (Phragmites australis (Cav.) Steudel) detritus decomposition was studied during the fall and spring periods, utilising the leaf packs technique with 3 sampling dates and 8 replicate packs per sample in each period. At all sites, the stream width was recorded fortnightly for one year. Overall, the reed leaf packs underwent rapid decomposition in the river Tirso basin (k=0.0193 d^?1), but very high variation was observed among leaf pack decomposition rates (C.V. = 112.7%). Seasons, sub-basins, and stream orders accounted for 88% of the total variance observed (Three Way ANOVA). The decomposition rate was significantly higher in spring than in fall, in the Laconi sub-basin than in the Olzai and Assolo sub-basins, and in 4th order streams than in 2nd and 3rd. Decomposition rates were also higher at relatively undisturbed sites than at the study sites exposed to complete summer desiccation (e.g., 2nd, 3rd, 4th in the Olzai sub-basin; 1st, 2nd, 3rd in the Assolo sub-basin). The strongest effects of summer drought disturbance occurred at the lowest order stream branches (i.e., 1st and 2nd order) suggesting that the resilience of detritus decomposition processes to the disturbance arising from Mediterranean type climates increases with stream order.     

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.     

Simulating food web dynamics along a gradient: quantifying human influence

Realistically parameterized and dynamically simulated food-webs are useful tool to explore the importance of the functional diversity of ecosystems, and in particular relations between the dynamics of species and the whole community. We present a stochastic dynamical food web simulation for the Kelian River (Borneo). The food web was constructed for six different locations, arrayed along a gradient of increasing human perturbation (mostly resulting from gold mining activities) along the river. Along the river, the relative importance of grazers, filterers and shredders decreases with increasing disturbance downstream, while predators become more dominant in governing eco-dynamics. Human activity led to increased turbidity and sedimentation which adversely impacts primary productivity. Since the main difference between the study sites was not the composition of the food webs (structure is quite similar) but the strengths of interactions and the abundance of the trophic groups, a dynamical simulation approach seemed to be useful to better explain human influence. In the pristine river (study site 1), when comparing a structural version of our model with the dynamical model we found that structurally central groups such as omnivores and carnivores were not the most important ones dynamically. Instead, primary consumers such as invertebrate grazers and shredders generated a greater dynamical response. Based on the dynamically most important groups, bottom-up control is replaced by the predominant top-down control regime as distance downstream and human disturbance increased. An important finding, potentially explaining the poor structure to dynamics relationship, is that indirect effects are at least as important as direct ones during the simulations. We suggest that our approach and this simulation framework could serve systems-based conservation efforts. Quantitative indicators on the relative importance of trophic groups and the mechanistic modeling of eco-dynamics could greatly contribute to understanding various aspects of functional diversity.