1. Riparian vegetation composition, structure and abundance are governed to a large degree by river flow regime and flow-mediated fluvial processes. Streamflow regime exerts selective pressures on riparian vegetation, resulting in adaptations (trait syndromes) to specific flow attributes. Widespread modification of flow regimes by humans has resulted in extensive alteration of riparian vegetation communities. Some of the negative effects of altered flow regimes on vegetation may be reversed by restoring components of the natural flow regime. 2. Models have been developed that quantitatively relate components of the flow regime to attributes of riparian vegetation at the individual, population and community levels. Predictive models range from simple statistical relationships, to more complex stochastic matrix population models and dynamic simulation models. Of the dozens of predictive models reviewed here, most treat one or a few species, have many simplifying assumptions such as stable channel form, and do not specify the time-scale of response. In many cases, these models are very effective in developing alternative streamflow management plans for specific river reaches or segments but are not directly transferable to other rivers or other regions. 3. A primary goal in riparian ecology is to develop general frameworks for prediction of vegetation response to changing environmental conditions. The development of riparian vegetation-flow response guilds offers a framework for transferring information from rivers where flow standards have been developed to maintain desirable vegetation attributes, to rivers with little or no existing information. 4. We propose to organise riparian plants into non-phylogenetic groupings of species with shared traits that are related to components of hydrologic regime: life history, reproductive strategy, morphology, adaptations to fluvial disturbance and adaptations to water availability. Plants from any river or region may be grouped into these guilds and related to hydrologic attributes of a specific class of river using probabilistic response curves. 5. Probabilistic models based on riparian response guilds enable prediction of the likelihood of change in each of the response guilds given projected changes in flow, and facilitate examination of trade-offs and risks associated with various flow management strategies. Riparian response guilds can be decomposed to the species level for individual projects or used to develop flow management guidelines for regional water management plans.
Temporal environmental fluctuations, such as seasonality, exert strong controls on biodiversity. While the effects of seasonality are well known, the predictability of fluctuations across years may influence seasonality in ways that are less well understood. The ability of a habitat to support unique, non-nested assemblages of species at different times of the year should depend on both seasonality (occurrence of events at specific periods of the year) and predictability (the reliability of event recurrence) of characteristic ecological conditions. Drawing on tools from wavelet analysis and information theory, we developed a framework for quantifying both seasonality and predictability of habitats, and applied this using global long-term rainfall data. Our analysis predicted that temporal beta diversity should be maximized in highly predictable and highly seasonal climates, and that low degrees of seasonality, predictability, or both would lower diversity in characteristic ways. Using stream invertebrate communities as a case study, we demonstrated that temporal species diversity, as exhibited by community turnover, was determined by a balance between temporal environmental variability (seasonality) and the reliability of this variability (predictability). Communities in highly seasonal mediterranean environments exhibited strong oscillations in community structure, with turnover from one unique community type to another across seasons, whereas communities in aseasonal New Zealand environments fluctuated randomly. Understanding the influence of seasonal and other temporal scales of environmental oscillations on diversity is not complete without a clear understanding of their predictability, and our framework provides tools for examining these trends at a variety of temporal scales, seasonal and beyond. Given the uncertainty of future climates, seasonality and predictability are critical considerations for both basic science and management of ecosystems (e.g., dam operations, bioassessment) spanning gradients of climatic variability.
Abstract1. River networks are hierarchical dendritic habitats embedded within the terrestrial landscape, with varying connectivity between sites depending on their positions along the network. This physical organisation influences the dispersal of organisms, which ultimately affects metacommunity dynamics and biodiversity patterns.2. We provide a conceptual synthesis of the role of river networks in structuring metacommunities in relation to dispersal processes in riverine ecosystems. We explore where the river network best explains observed metacommunity structure compared to other measurements of physical connectivity. We mostly focus on invertebrates, but also consider other taxonomic groups, including microbes, fishes, plants, and amphibians.3. Synthesising studies that compared multiple spatial distance metrics, we found that the importance of the river network itself in explaining metacommunity patterns depended on a variety of factors, including dispersal mode (aquatic versus aerial versus terrestrial) and landscape type (arid versus mesic), as well as location-specific factors, such as network connectivity, land use, topographic heterogeneity, and biotic interactions. The river network appears to be less important for strong aerial dispersers and insects in arid systems than for other groups and biomes, but there is considerable variability. Borrowing from other literature, particularly landscape genetics, we developed a conceptual model that predicts that the explanatory power of the river network peaks in mesic systems for obligate aquatic dispersers. 4.We propose directions of future avenues of research, including the use of manipulative field and laboratory experiments that test metacommunity theory in river networks. While field and laboratory experiments have their own benefits and drawbacks (e.g. reality, control, cost), both are powerful approaches for understanding the mechanisms structuring metacommunities, by teasing apart dispersal and niche-related factors.5. Finally, improving our knowledge of dispersal in river networks will benefit from expanding the breadth of cost-distance modelling to better infer dispersal from observational data; an improved understanding of life-history strategies rather than relying on independent traits; exploring individual-level variation in dispersal through detailed genetic studies; detailed studies on fine-scale environmental
Aim Meta-community structure is a function of both local (site-specific) and regional (landscape-level) ecological factors, and the relative importance of each may be mediated by the dispersal ability of organisms. Here, we used aquatic invertebrate communities to investigate the relationship between local and regional factors in explaining distance decay relationships (DDRs) in fragmented dendritic stream networks.Location Dryland streams distributed within a 400-km 2 section of the San Pedro River basin, south-eastern Arizona, USA.Methods We combined fine-scale local information (flow and habitat characteristics) with regional-scale information to explain DDR patterns in community composition of aquatic invertebrate species with a wide range of dispersal abilities. We used a novel application of a landscape resistance modelling approach (originally developed for landscape genetic studies) that simultaneously assessed the importance of local and regional ecological factors as well as dispersal ability of organisms.Results We found evidence that both local and regional factors influenced aquatic invertebrate DDRs in dryland stream networks, and the importance of each factor depended on the dispersal capacities of the organisms. Local and weak dispersers were more affected by site-specific factors, intermediate dispersers by landscape-level factors, and strong dispersers showed no discernable pattern. This resulted in a strongly hump-shaped relationship between dispersal ability and landscape-level factors, where only moderate dispersers showed evidence of DDRs. Unlike most other studies of dendritic networks, our results suggest that overland pathways, using perennial refugia as stepping-stones, might be the main dispersal route in fragmented stream networks.Main conclusions We suggest that using a combination of landscape and local distance measures can help to unravel meta-community patterns in dendritic systems. Our findings have important conservation implications, such as the need to manage river systems for organisms that span a wide variety of dispersal abilities and local ecological requirements. Our results also highlight the need to preserve perennial refugia in fragmented networks, as they may ensure the viability of aquatic meta-communities by facilitating dispersal.
Summary 1. Temporary streams comprise a large proportion of the total length of most stream networks, and the great majority of arid‐land stream networks, so it is important to understand their contribution to biotic diversity at both local and landscape scales. 2. In late winter 2010, we sampled invertebrate assemblages in 12 reaches of a large arid‐land stream network (including perennial and intermittent headwaters, intermittent middle reaches and perennial rivers) in south‐east Arizona, U.S.A. Intermittent reaches had then been flowing for c. 60 days, following a dry period of more than 450 days. We sampled a subset of the perennial study reaches three more times between 2009 and 2011. Since intermittent reaches were dry during these additional sampling periods, we used assemblage data from two other intermittent streams in the study network (sampled in 2004–05 and 2010) to explore interannual variability in intermittent stream assemblage composition. 3. Invertebrate richness was lowest in intermittent reaches, despite their often being connected to species‐rich perennial reaches. The assemblages of these intermittent reaches were not simply a subset of the species in perennial streams, but rather were dominated by a suite of stoneflies, blackflies and midges with adaptations to intermittency (e.g. egg and/or larval diapause). On average, 86% of individuals in these samples were specialists or exclusive to intermittent streams. Predators were 7–14 times more abundant in perennial than in intermittent reaches. 4. Despite being separated by long distances (12–25 km) and having very different physical characteristics, the assemblages of perennial headwaters and rivers were more similar to one another than to intervening intermittent reaches, emphasising the prime importance of local hydrology in this system. 5. The duration and recurrence intervals of dry periods, and the relative importance of dispersal from perennial refuges, probably influence the magnitude of biological differences between neighbouring perennial and temporary streams. Although perennial headwaters supported the highest diversity of invertebrates, intermittent reaches supported a number of unique or locally rare species and as such contribute to regional species diversity and should be included in conservation planning.
Riparian cottonwood (Populus deltoides) forests form the one of the most extensive deciduous forest ecosystems in arid regions of the western United States. However, cottonwood populations are threatened by flow alteration and channel degradation caused by dams, water diversions, and groundwater pumping. We developed a stochastic, densitydependent, population model to (1) consolidate information concerning cottonwood population dynamics in a conceptual and analytical framework, (2) determine whether complex forest stand dynamics can be predicted from basic cottonwood vital rates and river hydrology, and (3) aid in planning prescribed floods by projecting how altered flow regimes might affect populations. The model describes how annual variation in the hydrograph affects cottonwood mortality (via floods and droughts) and recruitment (via scouring of new habitat and seedling establishment). Using parameter values for the undammed Yampa River in Colorado, we found that abundances of seedlings and younger trees followed a boom-bust cycle driven by high flood mortalities while reproductive adult abundance followed a less erratic 5-15-yr periodicity driven by multiyear sequences of flows favorable to stand recruitment. Conversely, chance occurrences of multiple drought years eliminated cottonwood from up to 50% of available habitat, providing opportunities for competing plant species to establish. By simulating flow alterations on the Yampa ranging from channelization (many floods/droughts) to damming (few floods/droughts), the model suggested that mature cottonwood forest should be most abundant near the observed natural flow regime. Model analysis also suggested that flow regimes with high flood frequencies result in stable (albeit small) population sizes, while stable flows result in highly variable population sizes prone to local extinction.
Summary Climate change is expected to intensify drought in many regions, but ecological impacts on stream communities are poorly understood. Many arid‐land streams are characterised by predictable seasonal cycles of wetting and drying, to which species are adapted, but unpredictable supraseasonal droughts may constitute extreme events that challenge resident biota. In this article, we synthesise research conducted in arid‐land streams of the Madrean Sky Islands (MSI) in Arizona, U.S.A, to evaluate the resistance and resilience of invertebrate communities to drying disturbances caused by normal seasonal drying and severe supraseasonal drought. We also highlight how spatial context (e.g. distance to perennial refuges) influences recovery patterns. Invertebrate community structure changes predictably as habitat contraction progresses from loss of lateral connectivity to complete drying of MSI streams. When drying events are predictable (e.g. seasonal drying), post‐drought community recovery is often rapid, since most MSI taxa possess life history traits conferring high resistance and/or resilience to stream drying. Extreme supraseasonal droughts, in contrast, cause unprecedented transitions from perennial to intermittent flow in some MSI streams. While species richness may recover quickly following this flow regime shift, marked turnover in community structure can occur and may delay or preclude recovery to pre‐drought conditions. In such cases, short‐lived (<1 year) strong dispersers replace relatively long‐lived (≥1 year) weak dispersers. As habitat isolation increases, the potential for community recovery from extreme drought decreases. Many MSI aquatic species are threatened by extreme drought. Extinctions of endemic aquatic species due to habitat drying have already been observed in nearby deserts. Further studies are urgently needed to identify drought‐sensitive species and understand how the loss of such species may affect stream ecosystem functioning.
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