Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver-response relationships to develop conceptual models across ecosystems.

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed 'beta diversity') is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.

Consequences of hydrological alteration for beta diversity of fish assemblages at multiple spatial scales.

Effects of dam operation and extraction of water from rivers on spatial variation in hydrological regimes, and consequences for freshwater biodiversity, are widely predicted but seldom assessed empirically. Evidence of linkages between hydrology and beta diversity contributes to water-management decisions to support landscape-scale biodiversity and avoid inadvertently contributing to further biodiversity decline. Using six lowland rivers in Australia's Murray - Darling Basin that formed a gradient of hydrological alteration, we examined (1) spatial variation in hydrology under modelled scenarios of low water-resource development and flow modification by dams and extraction, (2) how beta diversity of fish among and within rivers was associated with spatial hydrological variation and whether patterns of overall beta diversity differed between native and non-native species, and (3) the associations of spatial and environmental variables and both recent and long-term hydrology with beta diversity. Spatial variation in hydrology among rivers was higher under the modified scenario than under the low-development scenario yet change in the magnitude of within-river (longitudinal) variation was inconsistent between rivers. Beta diversity among rivers was significantly associated with spatial variation in hydrology only in certain circumstances (native species assemblages in specific years). Within-river beta diversity varied among rivers yet was unrelated to longitudinal variation in modified hydrological regimes. Patterns of beta diversity did not differ appreciably if non-native species were included in or excluded from analyses. These findings contradict predictions adopted in ecohydrological science that water resource development homogenises hydrological regimes, in turn causing biotic homogenisation in lowland rivers.

Assessing Effects of Flow Regulation and an Experimental Flow Pulse on Population Size Structure of Riverine Fish with Contrasting Biological Characteristics.

Despite effects of dams and water extraction on river hydrology and consequences of aquatic ecosystems being broadly appreciated, empirical evidence is lacking for many regions (e.g. subtropics). Evidence is necessary to determine (i) the circumstances where environmental flows are necessary to protect or improve ecological processes and (ii) what hydrological events are required to achieve those ecological characteristics. Here, temporal variation in the size structure of two small-bodied fish species with contrasting ecological characteristics (Australian smelt, Cox's gudgeon) was compared between two pairs of unregulated and regulated rivers in subtropical Australia. Frequency of in-channel flow pulses in each regulated river was lower compared to paired unregulated rivers. An experimental flow pulse was delivered to one regulated river to assess the ecological outcomes of in-channel flow pulses and inform future decisions about the use of environmental water allocations. Temporal changes in the population size structure of both species were similar between unregulated and regulated rivers and showed no response to the experimental pulse. While the experimental flow briefly suppressed in-stream temperature, changes in temperature were not beyond the thresholds at which spawning occurs or the thermal tolerances for either species. Similar population structure between unregulated and regulated rivers can be attributed to the magnitude of flow regulation being insufficient to alter the physico-chemical conditions, habitat and trophic mechanisms supporting population dynamics during the study period. This suggests current regulated hydrology is protective of local populations of Australian smelt and Cox's gudgeon in these study rivers.

Scaling biodiversity responses to hydrological regimes.

Of all ecosystems, freshwaters support the most dynamic and highly concentrated biodiversity on Earth. These attributes of freshwater biodiversity along with increasing demand for water mean that these systems serve as significant models to understand drivers of global biodiversity change. Freshwater biodiversity changes are often attributed to hydrological alteration by water-resource development and climate change owing to the role of the hydrological regime of rivers, wetlands and floodplains affecting patterns of biodiversity. However, a major gap remains in conceptualising how the hydrological regime determines patterns in biodiversity's multiple spatial components and facets (taxonomic, functional and phylogenetic). We synthesised primary evidence of freshwater biodiversity responses to natural hydrological regimes to determine how distinct ecohydrological mechanisms affect freshwater biodiversity at local, landscape and regional spatial scales. Hydrological connectivity influences local and landscape biodiversity, yet responses vary depending on spatial scale. Biodiversity at local scales is generally positively associated with increasing connectivity whereas landscape-scale biodiversity is greater with increasing fragmentation among locations. The effects of hydrological disturbance on freshwater biodiversity are variable at separate spatial scales and depend on disturbance frequency and history and organism characteristics. The role of hydrology in determining habitat for freshwater biodiversity also depends on spatial scaling. At local scales, persistence, stability and size of habitat each contribute to patterns of freshwater biodiversity yet the responses are variable across the organism groups that constitute overall freshwater biodiversity. We present a conceptual model to unite the effects of different ecohydrological mechanisms on freshwater biodiversity across spatial scales, and develop four principles for applying a multi-scaled understanding of freshwater biodiversity responses to hydrological regimes. The protection and restoration of freshwater biodiversity is both a fundamental justification and a central goal of environmental water allocation worldwide. Clearer integration of concepts of spatial scaling in the context of understanding impacts of hydrological regimes on biodiversity will increase uptake of evidence into environmental flow implementation, identify suitable biodiversity targets responsive to hydrological change or restoration, and identify and manage risks of environmental flows contributing to biodiversity decline.

A framework for evaluating food-web responses to hydrological manipulations in riverine systems.

Environmental flows are used to restore elements of the hydrological regime altered by human use of water. One of the primary justifications and purposes for environmental flows is the maintenance of target species populations but, paradoxically, there has been little emphasis on incorporating the food-web and trophic dynamics that determine population-level responses into the monitoring and evaluation of environmental flow programs. We develop a generic framework for incorporating trophic dynamics into monitoring programs to identify the food-web linkages between hydrological regimes and population-level objectives of environmental flows. These linkages form the basis for objective setting, ecological targets and indicator selection that are necessary for planning monitoring programs with a rigorous scientific basis. Because there are multiple facets of trophic dynamics that influence energy production and transfer through food webs, the specific objectives of environmental flows need to be defined during the development of monitoring programs. A multitude of analytical methods exist that each quantify distinct aspects of food webs (e.g. energy production, prey selection, energy assimilation), but no single method can provide a basis for holistic understanding of food webs. Our paper critiques a range of analytical methods for quantifying attributes of food webs to inform the setting, monitoring and evaluation of trophic outcomes of environmental flows and advance the conceptual understanding of trophic dynamics in river-floodplain systems.

Modelling food-web mediated effects of hydrological variability and environmental flows.

Environmental flows are designed to enhance aquatic ecosystems through a variety of mechanisms; however, to date most attention has been paid to the effects on habitat quality and life-history triggers, especially for fish and vegetation. The effects of environmental flows on food webs have so far received little attention, despite food-web thinking being fundamental to understanding of river ecosystems. Understanding environmental flows in a food-web context can help scientists and policy-makers better understand and manage outcomes of flow alteration and restoration. In this paper, we consider mechanisms by which flow variability can influence and alter food webs, and place these within a conceptual and numerical modelling framework. We also review the strengths and weaknesses of various approaches to modelling the effects of hydrological management on food webs. Although classic bioenergetic models such as Ecopath with Ecosim capture many of the key features required, other approaches, such as biogeochemical ecosystem modelling, end-to-end modelling, population dynamic models, individual-based models, graph theory models, and stock assessment models are also relevant. In many cases, a combination of approaches will be useful. We identify current challenges and new directions in modelling food-web responses to hydrological variability and environmental flow management. These include better integration of food-web and hydraulic models, taking physiologically-based approaches to food quality effects, and better representation of variations in space and time that may create ecosystem control points.

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish.

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context-dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non-native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.

Can species traits predict the susceptibility of riverine fish to water resource development? An Australian case study.

Water resource developments alter riverine environments by disrupting longitudinal connectivity, transforming lotic habitats, and modifying in-stream hydraulic conditions. Effective management of anthropogenic disturbances therefore requires an understanding of the range of potential ecosystem effects and the inherent traits symptomatic of elevated vulnerability to disturbance. Using 42 riverine fish native to South Eastern Australia as a case study, we quantified six morphological, behavioral, and life-history traits to classify species into groups reflecting potential differences in their response to ecosystem changes as a result of water resource development. Classification analysis identified five strategies based on fish life-history dispersal requirements, climbing potential, and habitat preference. These strategies in turn highlight the potential species at risk from the separate impacts of water resource development and inform management decisions to mitigate those risks. Swimming ability did not contribute to distinguishing species into functional groups, likely due to methodological inconsistencies in quantifying swimming performance that may ultimately hinder the ability of fish passage facilities to function within the physical capabilities of species at risk of habitat fragmentation. This study improves our ability to predict the performance of groups of species at risk from the multiple environmental changes imposed by humans and goes beyond broad-scale dispersal requirements as a predictor of individual species response.

Cattle-derived microbial input to source water catchments: An experimental assessment of stream crossing modification.

The provision of safe drinking water is a global issue, and animal production is recognized as a significant potential origin of human infectious pathogenic microorganisms within source water catchments. On-farm management can be used to mitigate livestock-derived microbial pollution in source water catchments to reduce the risk of contamination to potable water supplies. We applied a modified Before-After Control Impact (BACI) design to test if restricting the access of livestock to direct contact with streams prevented longitudinal increases in the concentrations of faecal indicator bacteria and suspended solids. Significant longitudinal increases in pollutant concentrations were detected between upstream and downstream reaches of the control crossing, whereas such increases were not detected at the treatment crossing. Therefore, while the crossing upgrade was effective in preventing cattle-derived point source pollution by between 112 and 158%, diffuse source pollution to water supplies from livestock is not ameliorated by this intervention alone. Our findings indicate that stream crossings that prevent direct contact between livestock and waterways provide a simple method for reducing pollutant loads in source water catchments, which ultimately minimises the likelihood of pathogenic microorganisms passing through source water catchments and the drinking water supply system. The efficacy of the catchment as a primary barrier to pathogenic risks to drinking water supplies would be improved with the integration of management interventions that minimise direct contact between livestock and waterways, combined with the mitigation of diffuse sources of livestock-derived faecal matter from farmland runoff to the aquatic environment.

Spatial and temporal patterns in fish assemblages following an artificially extended floodplain inundation event, northern Murray-Darling Basin, Australia.

Water extraction from dryland rivers is often associated with declines in the health of river and floodplain ecosystems due to reduced flooding frequency and extent of floodplain inundation. Following moderate flooding in early 2008 in the Narran River, Murray-Darling Basin, Australia, 10,423 ML of water was purchased from agricultural water users and delivered to the river to prolong inundation of its terminal lake system to improve the recruitment success of colonial waterbirds that had started breeding in response to the initial flooding. This study examined the spatial and temporal patterns of fish assemblages in river and floodplain habitats over eight months following flooding to assess the possible ecological benefits of flood extension. Although the abundances of most fish species were greater in river channel habitats, the fish assemblage used floodplain habitats when inundated. Young-of-the-year (4-12 months age) golden perch (Macquaria ambigua) and bony bream (Nematalosa erebi) were consistently sampled in floodplain sites when inundated, suggesting that the floodplain provides rearing habitat for these species. Significant differences in the abundances of fish populations between reaches upstream and downstream of a weir in the main river channel indicates that the effectiveness of the environmental water release was limited by restricted connectivity within the broader catchment. Although the seasonal timing of flood extension may have coincided with sub-optimal primary production, the use of the environmental water purchase is likely to have promoted recruitment of fish populations by providing greater access to floodplain nursery habitats, thereby improving the ability to persist during years of little or no flow.