The fauna of dynamic riverine landscapes

1.  Riverine landscapes are heterogeneous in space (complex mosaic of habitat types) and time (expansion and contraction cycles, landscape legacies). They are inhabited by a diverse and abundant fauna of aquatic, terrestrial and amphibious species.
2.  Faunal distribution patterns are determined by interactive processes that reflect the landscape mosaic and complex environmental gradients. The life cycles of many riverine species rely upon a shifting landscape mosaic and other species have become adapted to exploit the characteristically high turn-over of habitats.
3.  The complex landscape structure provides a diversity of habitats that sustains various successional stages of faunal assemblages. A dynamic riverine landscape sustains biodiversity by providing a variety of refugia and through ecological feedbacks from the organisms themselves (ecosystem engineering).
4.  The migration of many species, aquatic and terrestrial, is tightly coupled with the temporal and spatial dynamics of the shifting landscape mosaic. Alternation of landscape use by terrestrial and aquatic fauna corresponds to the rise and fall of the flood. Complex ecological processes inherent to intact riverine landscapes are reflected in their biodiversity, with important implications for the restoration and management of river corridors.     

Food web structure in riverine landscapes

1. Most research on freshwater (and other) food webs has focused on apparently discrete communities, in well-defined habitats at small spatial and temporal scales, whereas in reality food webs are embedded in complex landscapes, such as river corridors. Food web linkages across such landscapes may be crucial for ecological pattern and process, however. Here, we consider the importance of large scale influences upon lotic food webs across the three spatial dimensions and through time.
2. We assess the roles of biotic factors (e.g. predation, competition) and physical habitat features (e.g. geology, land-use, habitat fragmentation) in moulding food web structure at the landscape scale. As examples, external subsidies to lotic communities of nutrients, detritus and prey vary along the river corridor, and food web links are made and broken across the land–water interface with the rise and fall of the flood.
3. We identify several avenues of potentially fruitful research, particularly the need to quantify energy flow and population dynamics. Stoichiometric analysis of changes in C : N : P nutrient ratios over large spatial gradients (e.g. from river source to mouth, in forested versus agricultural catchments), offers a novel method of uniting energy flow and population dynamics to provide a more holistic view of riverine food webs from a landscape perspective. Macroecological approaches can be used to examine large-scale patterns in riverine food webs (e.g. trophic rank and species–area relationships). New multivariate statistical techniques can be used to examine community responses to environmental gradients and to assign traits to individual species (e.g. body-size, functional feeding group), to unravel the organisation and trophic structure of riverine food webs.     

Island-dominated landscapes of large floodplain rivers, a European perspective

1. The landscapes of large floodplain rivers are characterised by heterogeneous environments related to the interplay of flood flows, sediment transport and vegetation dynamics.
2. The large rivers of Europe, and probably most rivers throughout the forest biomes, were characterised by islands but over the period of major human interference, many have become dominated by incision and narrowing so that they are now characterised by single-thread and relatively simple channel forms.
3. Vegetation plays an active role in developing heterogeneous channel forms through (a) biotic processes such as seed dispersal, vegetative regeneration and succession and (b) abiotic effects such as increasing flow resistance inducing sedimentation, and decreasing bank erodibility.
4. In particular, accumulations of living driftwood (cf. dead driftwood accumulations and dispersed seedlings) accelerate sedimentation and island development.
5. River reaches with vegetated islands have a high habitat diversity.
6. The natural influences of flood disturbance, wood accumulation, vegetation growth, island development and tree die-off, cause island-dominated reaches to undergo cycles of island growth and decay that are related to cycles of aquatic habitat diversification and simplification.     

Riverine landscape diversity

1. This review is presented as a broad synthesis of riverine landscape diversity, beginning with an account of the variety of landscape elements contained within river corridors. Landscape dynamics within river corridors are then examined in the context of landscape evolution, ecological succession and turnover rates of landscape elements. This is followed by an overview of the role of connectivity and ends with a riverine landscape perspective of biodiversity. 2. River corridors in the natural state are characterised by a diverse array of landscape elements, including surface waters (a gradient of lotic and lentic waterbodies), the fluvial stygoscape (alluvial aquifers), riparian systems (alluvial forests, marshes, meadows) and geomorphic features (bars and islands, ridges and swales, levees and terraces, fans and deltas, fringing floodplains, wood debris deposits and channel networks). 3. Fluvial action (erosion, transport, deposition) is the predominant agent of landscape evolution and also constitutes the natural disturbance regime primarily responsible for sustaining a high level of landscape diversity in river corridors. Although individual landscape features may exhibit high turnover, largely as a function of the interactions between fluvial dynamics and successional phenomena, their relative abundance in the river corridor tends to remain constant over ecological time. 4. Hydrological connectivity, the exchange of matter, energy and biota via the aqueous medium, plays a major though poorly understood role in sustaining riverine landscape diversity. Rigorous investigations of connectivity in diverse river systems should provide considerable insight into landscape‐level functional processes. 5. The species pool in riverine landscapes is derived from terrestrial and aquatic communities inhabiting diverse lotic, lentic, riparian and groundwater habitats arrayed across spatio‐temporal gradients. Natural disturbance regimes are responsible for both expanding the resource gradient in riverine landscapes as well as for constraining competitive exclusion. 6. Riverine landscapes provide an ideal setting for investigating how complex interactions between disturbance and productivity structure species diversity patterns.     

Local and ecoregion effects on fish assemblage structure in tributaries of the Rio Paraíba do Sul, Brazil

1.  We examined the effects of physical and chemical habitat variables and ecoregions on species occurrence and fish assemblage structure in streams of the Paraíba do Sul basin, in southeast Brazil.
2.  Fish and environmental data were collected from 42 sites on 26 first to fourth order streams (1 : 50 000 map scale) in three ecoregions. The sites occurred in one valley and two plateau ecoregions at altitudes of 40–1080 m and distances of 0.1–188 km from the main channel of the Rio Paraíba do Sul. Physical habitat (substratum, riparian cover, habitat types) and water quality (dissolved oxygen, pH, temperature and conductivity) variables were measured at each sampling site.
3.  A total of 2684 individuals in 16 families and 59 species were recorded.
4.  Ecoregion was a better predictor of the fish assemblage than the other environmental variables, according to the differences between the mean within-class and mean between-class similarities in assemblage data.
5.  Differing landscape characteristics were associated with differing local variables and thereby with differing fish assemblage structures. Riffles, shrub, grass, dissolved oxygen, conductivity and temperature were closely related to fish assemblage structure.
6.  Fish assemblages in sites far from the main river and at higher altitudes also differed from those near the Paraíba do Sul main channel, presumably as a result of differences in connectivity, covarying environmental factors and anthropogenic influence.
7.  These results reinforce the importance of understanding how stream communities are influenced by processes and patterns operating at local and regional scales, which will aid water resource managers to target those factors in their management and rehabilitation efforts.     

Global patterns of diversity among the specialist birds of riverine landscapes

1. Despite the growing view that biodiversity provides a unifying theme in river ecology, global perspectives on richness in riverine landscapes are limited. As a result, there is little theory or quantitative data on features that might have influenced global patterns in riverine richness, nor are there clear indications of which riverine landscapes are important to conservation at the global scale. As conspicuous elements of the vertebrate fauna of riverine landscapes, we mapped the global distributions of all of the world's specialist riverine birds and assessed their richness in relation to latitude, altitude, primary productivity and geomorphological complexity (surface configuration). 2. Specialist riverine birds, typical of high‐energy riverine landscapes and dependent wholly or partly on production from river ecosystems, occur in 16 families. They are represented by an estimated 60 species divided equally between the passerines and non‐passerines. Major radiation has occurred among different families on different continents, indicating that birds have evolved several times into the niches provided by riverine landscapes. 3. Continental richness varies from four species in Europe to 28 in Asia, with richness on the latter continent disproportionately larger than would be expected from a random distribution with respect to land area. Richness is greatest in mountainous regions at latitudes of 20–40°N in the riverine landscapes of the Himalayan mountains, where 13 species overlap in range. 4. Family, genus and species richness in specialist riverine birds all increase significantly with productivity and surface configuration (i.e. relief). However, family richness was the best single predictor of the numbers of species or genera. In keeping with the effect of surface configuration, river‐bird richness peaks globally at 1300–1400 m altitude, and most species occur typically on small, fast rivers where they feed predominantly on invertebrates. Increased lengths of such streams in areas of high relief and rainfall might have been responsible for species–area effects. 5. We propose the hypothesis that the diversity in channel forms and habitats in riverine landscapes, in addition to high temperature and primary productivity, have been prerequisites to the development of global patterns in the richness of specialist riverine organisms. We advocate tests of this hypothesis in other taxonomic groups. We draw attention, however, to the challenges of categorically defining riverine organisms in such tests because (i) rivers grade into many other habitat types across several different ecotones and (ii) `terrestrialisation' processes in riverine landscapes means that they offer habitat for organisms whose evolutionary origins are not exclusively riverine.     

Seasonal changes in habitat suitability for adult shovelnose sturgeon in the lower Mississippi River

Shovelnose sturgeon Scaphirhynchus platorynchus are a large‐river fish distributed throughout the Mississippi River basin, including the lower 1,533 km of the Mississippi River where riverine habitat has been and continues to be modified for navigation and is a potential site for development of instream hydrokinetic electric power generation. Information about habitat use and preference is essential to future conservation efforts. Shovelnose sturgeon have previously been found to select particular habitat types, and these selected habitats vary seasonally; although these past analyses do not consider the selected habitats in a landscape context. We used ecological niche factor analysis (ENFA) that uses distributions of telemetry locations and environmental variables to model habitat suitability in a landscape context. We recorded 333 locations of shovelnose sturgeon during July–December 2013 that included periods of relatively high and low river stages. The ENFA analysis indicated high‐suitability locations were in or near deep water during both high and low river stages. During high river stages, high‐suitability locations were near island tip habitat, deep water, and steep bottom slope and far from main channel habitat. During low stages, high‐suitability locations were in or near deep water and main channel habitat and far from secondary channel and wing dike habitats. This landscape‐scale analysis supports seasonal shifts in habitat use and provides insights that can be used to inform habitat conservation and management to benefit shovelnose sturgeon in the lower Mississippi River and possibly other large rivers.     

Movement patterns of the bush cricket Platycleis albopunctata in different types of habitat: matrix is not always matrix

Abstract.  1. Inter-patch movement is usually assumed to be homogeneous across a landscape. As the intervening area between suitable patches is usually richly textured, it cannot be assumed to be uniform in real landscapes.
2. In an experimental mark-and-resight study, the movement behaviour of the bush cricket Platycleis albopunctata in four habitat types as well as at the border between two of these habitat types was observed.
3. Analysis of recapture data indicated differences in mortality risk (or emigration rates) between habitat types.
4. When released at the border between suitable habitat and a crop field, P. albopunctata did not show a consistent preference for the suitable habitat. This suggests that the crop field is at least temporarily attractive for P. albopunctata .
5. Movement in suitable habitat was not always different from movement in the matrix, and movement between different types of matrix also differed.
6. The results indicate that the movement behaviour of P. albopunctata is influenced not only by suitability for breeding but also by structural resistance and other factors (e.g. food availability or habitat-specific mortality risk).     

Riverine landscapes: taking landscape ecology into the water

1. Landscape ecology deals with the influence of spatial pattern on ecological processes. It considers the ecological consequences of where things are located in space, where they are relative to other things, and how these relationships and their consequences are contingent on the characteristics of the surrounding landscape mosaic at multiple scales in time and space. Traditionally, landscape ecologists have focused their attention on terrestrial ecosystems, and rivers and streams have been considered either as elements of landscape mosaics or as units that are linked to the terrestrial landscape by flows across boundaries or ecotones. Less often, the heterogeneity that exists within a river or stream has been viewed as a `riverscape' in its own right.
2. Landscape ecology can be unified about six central themes: (1) patches differ in quality (2) patch boundaries affect flows, (3) patch context matters, (4) connectivity is critical, (5) organisms are important, and (6) the importance of scale. Although riverine systems differ from terrestrial systems by virtue of the strong physical force of hydrology and the inherent connectivity provided by water flow, all of these themes apply equally to aquatic and terrestrial ecosystems, and to the linkages between the two.
3. Landscape ecology therefore has important insights to offer to the study of riverine ecosystems, but these systems may also provide excellent opportunities for developing and testing landscape ecological theory. The principles and approaches of landscape ecology should be extended to include freshwater systems; it is time to take the `land' out of landscape ecology.     

The use of Bayesian networks to guide investments in flow and catchment restoration for impaired river ecosystems

1.  The provision of environmental flows and the removal of barriers to water flow are high priorities for restoration where changes to flow regimes have caused degradation of riverine ecosystems. Nevertheless, flow regulation is often accompanied by changes in catchment and riparian land-use, which also can have major impacts on river health via local habitat degradation or modification of stream energy regimes.
2.  The challenges are determining the relative importance of flow, land-use and other impacts as well as deciding where to focus restoration effort. As a consequence, flow, catchment and riparian restoration efforts are often addressed in isolation. River managers need decision support tools to assess which flow and catchment interventions are most likely to succeed and, importantly, which are cost-effective.
3.  Bayesian networks (BNs) can be used as a decision support tool for considering the influence of multiple stressors on aquatic ecosystems and the relative benefits of various restoration options. We provide simple illustrative examples of how BNs can address specific river restoration goals and assist with the prioritisation of flow and catchment restoration options. This includes the use of cost and utility functions to assist decision makers in their choice of potential management interventions.
4.  A BN approach facilitates the development of conceptual models of likely cause and effect relationships between flow regime, land-use and river conditions and provides an interactive tool to explore the relative benefits of various restoration options. When combined with information on the costs and expected benefits of intervention, one can derive recommendations about the best restoration option to adopt given the network structure and the associated cost and utility functions.     

Riparian vegetation diversity along regulated rivers: contribution of novel and relict habitats

1. The creation and maintenance of spatial and temporal heterogeneity by rivers flowing through floodplain landscapes has been disrupted worldwide by dams and water diversions. Large reservoirs ( novel ecosystems ) now separate and isolate remnant floodplains ( relict ecosystems ). From above, these appear as a string of beads, with beads of different sizes and string connections of varying lengths.
2. Numerous studies have documented or forecast sharp declines in riparian biodiversity in relict ecosystems downstream from dams. Concurrently, novel ecosystems containing species and communities of the former predam ecosystems have arisen along all regulated rivers. These result from the creation of new environments caused by upper reservoir sedimentation, tributary sedimentation and the formation of reservoir shorelines.
3. The contribution of novel habitats to the overall biodiversity of regulated rivers has been poorly studied. Novel ecosystems may become relatively more important in supporting riverine biodiversity if relict ecosystems are not restored to predam levels. The Missouri River of the north-central U.S.A. is used to illustrate existing conditions on a large, regulated river system with a mixture of relict and novel ecosystems.     

Ontogenetic patterns of habitat use by fishes within the main channel of an Australian floodplain river

The ontogenetic patterns of habitat use by a community of fishes in the main channel of the Broken River, an Australian lowland river, was investigated. Stratified sampling was conducted fortnightly across six habitat types throughout the spring‐summer period within the main channel. As predicted by the 'low flow recruitment hypothesis', backwaters and still littoral habitats were important nursery habitats for most species. These habitats were found to be used by some species throughout all stages of their life cycle, while other species showed clear ontogenetic shifts in habitat preference. Only one species, Murray cod Maccullochella peelii peelii , was never found in backwaters. This study confirms the significance of main channel habitats in the rearing of larvae of some riverine fish species, and emphasizes the importance of considering the habitat requirements of all stages of a fish's life cycle in the management and restoration of rivers and streams.     

Microfaunal communities in three lowland rivers under differing flow regimes

Microfaunal samples were collected from within the channels of three rivers in north eastern Victoria, Australia (the Murray, Ovens and Broken Rivers) as a component of a study examining the effects of flow on the biota of lowland rivers in Australia. Samples were collected from the water column of the river channel and slackwaters and from the layer of water immediately above the bottom sediment of the slackwaters. There was no connectivity between the river channel and the floodplain wetlands for all three rivers during the sampling period. Substantial numbers of microfauna were resident in the slackwaters of all three rivers, with the greatest densities occurring close to the bottom sediment, with densities often exceeding 1000 animals l⁻¹ whereas in the plankton samples densities were usually less than 500 animals l⁻¹. The presence of large and diverse microfaunal communities and the lack of connectivity between the river channel and associated floodplain wetland indicate that these communities are capable of persisting and recruiting within riverine channel slackwaters.     

Large wood and fluvial processes

1. Large wood forms an important component of woodland river ecosystems. The relationship between large wood and the physical characteristics of river systems varies greatly with changes in the tree species of the marginal woodland, the climatic and hydrological regime, the fluvial geomorphological setting and the river and woodland management context. 2. Research on large wood and fluvial processes over the last 25 years has focussed on three main themes: the effects of wood on flow hydraulics; on the transfer of mineral and organic sediment; and on the geomorphology of river channels. 3. Analogies between wood and mineral sediment transfer processes (supply, mobility and river characteristics that affect retention) are found useful as a framework for synthesising current knowledge on large wood in rivers. 4. An important property of wood is its size when scaled to the size of the river channel. ′Small′ channels are defined as those whose width is less than the majority of wood pieces (e.g. width < median wood piece length). `Medium' channels have widths greater than the size of most wood pieces (e.g. width < upper quartile wood piece length), and `Large' channels are wider than the length of all of the wood pieces delivered to them. 5. A conceptual framework defined here for evaluating the storage and dynamics of wood in rivers ranks the relative importance of hydrological characteristics (flow regime, sediment transport regime), wood characteristics (piece size, buoyancy, morphological complexity) and geomorphological characteristics (channel width, geomorphological style) in `Small', `Medium' and `Large' rivers. 6. Wood pieces are large in comparison with river size in `small' rivers, therefore they tend to remain close to where they are delivered to the river and provide important structures in the stream, controlling rather than responding to the hydrological and sediment transfer characteristics of the river. 7. For `Medium' rivers, the combination of wood length and form becomes critical to the stability of wood within the channel. Wood accumulations form as a result of smaller or more mobile wood pieces accumulating behind key pieces. Wood transport is governed mainly by the flow regime and the buoyancy of the wood. Even quite large wood pieces may require partial burial to give them stability, so enhancing the importance of the sediment transport regime. 8. Wood dynamics in `Large' rivers vary with the geometry of the channel (slope and channel pattern), which controls the delivery, mobility and breakage of wood, and also the characteristics of the riparian zone, from where the greatest volume of wood is introduced. Wood retention depends on the channel pattern and the distribution of flow velocity. A large amount is stored at the channel margins. The greater the contact between the active channel and the forested floodplain and islands, the greater the quantity of wood that is stored.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

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

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Collapse of an avifauna: climate change appears to exacerbate habitat loss and degradation

Aim  We characterized changes in reporting rates and abundances of bird species over a period of severe rainfall deficiency and increasing average temperatures. We also measured flowering in eucalypts, which support large numbers of nectarivores characteristic of the region.
Location  A 30,000-km² region of northern Victoria, Australia, consisting of limited amounts of remnant native woodlands embedded in largely agricultural landscapes.
Methods  There were three sets of monitoring studies, pitched at regional (survey programmes in 1995–97, 2004–05 and 2006–08), landscape (2002–03 and 2006–07) and site (1997–2008 continuously) scales. Bird survey techniques used a standard 2-ha, 20-min count method. We used Bayesian analyses of reporting rates to document statistically changes in the avifauna through time at each spatial scale.
Results  Bird populations in the largest remnants of native vegetation (up to 40,000 ha), some of which have been declared as national parks in the past decade, experienced similar declines to those in heavily cleared landscapes. All categories of birds (guilds based on foraging substrate, diet, nest site; relative mobility; geographical distributions) were affected similarly. We detected virtually no bird breeding in the latest survey periods. Eucalypt flowering has declined significantly over the past 12 years of drought.
Main conclusions  Declines in the largest woodland remnants commensurate with those in cleared landscapes suggest that reserve systems may not be relied upon to sustain species under climate change. We attribute population declines to low breeding success due to reduced food. Resilience of bird populations in this woodland system might be increased by active management to enhance habitat quality in existing vegetation and restoration of woodland in the more fertile parts of landscapes.     

Community structure and composition of microfaunal egg bank assemblages in riverine and floodplain sediments

Dormancy may be an important aspect influencing the ecology of riverine microfauna, yet fundamental knowledge concerning riverine egg bank communities is still scant compared with that for communities in floodplain habitats. We investigated the microfaunal egg bank communities in slackwater habitats of an Australian floodplain river, and compared them with the communities occurring in nearby floodplain wetlands. This was achieved by taking replicate sediment cores from paired examples of each habitat and later incubating the resting stages within these sediment cores. Results from the study indicated that the egg bank communities in each habitat differed in both composition and structure, with only 12 of the 31 taxa recorded being common to both habitat types. This suggests that in addition to supporting microfaunal persistence in the main channel, riverine egg bank communities represent an important source of microfaunal diversity together with floodplain egg bank communities in river–floodplain systems.     

The hydrogeomorphic approach to functional assessment of riparian wetlands: evaluating impacts and mitigation on river floodplains in the U.S.A.

1. The ’hydrogeomorphic‘ approach to functional assessment of wetlands (HGM) was developed as a synthetic mechanism for compensatory mitigation of wetlands lost or damaged by human activities. The HGM approach is based on: (a) classification of wetlands by geomorphic origin and hydrographic regime (b) assessment models that associate variables as indicators of function, and (c) comparison to reference wetlands that represent the range of conditions that may be expected in a particular region. In this paper, we apply HGM to riparian wetlands of alluvial rivers. 2. In the HGM classification, riverine wetlands are characterized by formative fluvial processes that occur mainly on flood plains. The dominant water sources are overbank flooding from the channel or subsurface hyporheic flows. Examples of riverine wetlands in the U.S.A. are: bottomland hardwood forests that typify the low gradient, fine texture substratum of the south-eastern coastal plain and the alluvial flood plains that typify the high gradient, coarse texture substratum of western montane rivers. 3. Assessment (logic) models for each of fourteen alluvial wetland functions are described. Each model is a composite of two to seven wetland variables that are independently scored in relation to a reference data set developed for alluvial rivers in the western U.S.A. Scores are summarized by a ’functional capacity index‘ (FCI), which is multiplied by the area of the project site to produce a dimensionless ’functional capacity unit‘ (FCU). When HGM is properly used, compensatory mitigation is based on the FCUs lost that must be returned to the riverine landscape under statutory authority. 4. The HGM approach also provides a framework for long-term monitoring of mitigation success or failure and, if failing, a focus on topical remediation. 5. We conclude that HGM is a robust and easy method for protecting riparian wetlands, which are critically important components of alluvial river landscapes.