Ground‐dwelling arthropod communities across braided river landscape mosaics: a Mediterranean perspective

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Unifying research on the fragmentation of terrestrial and aquatic habitats: patches,connectivity and the matrix in riverscapes

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Contrasting metacommunity structure and beta diversity in an aquatic‐floodplain system

Habitat connectivity and dispersal interact to structure metacommunities, but few studies have examined these patterns jointly for organisms across the aquatic–terrestrial ecotone. We assessed metacommunity structure and beta diversity patterns of instream benthic invertebrates, riparian carabid beetles (Order: Coleoptera; Family: Carabidae) and riparian spiders (Order: Araneae) at fifteen sites in a river‐floodplain system. Sampling took place over a three‐year period (2010–2012) in the Rhine‐Main‐Observatory LTER site on the Kinzig River, central Germany. This allowed disentangling the combined influence, and temporal variability, of habitat connectivity (i.e. between aquatic and terrestrial) and dispersal ability (i.e. between spiders and beetles, and aerial and aquatic dispersing invertebrates) on the dominant paradigms structuring these metacommunities. We found mostly consistent differences in the manner that metacommunities were structured between groups, with lower levels of variability explained for beetles compared to the other groups. Beetles were consistently structured more by turnover than nestedness components, with greater beta diversity than expected by chance and a minor spatial compared to environmental signal emerging with variance partitioning. Conversely, spiders and benthic invertebrates had lower beta diversity and greater nestedness than null expectation, and a clearer spatial signal controlling metacommunity structure. Our results suggest varying levels of mass effects and species sorting shape river‐floodplain metacommunities, depending on habitat connectivity and dispersal ability. That is, greater connectivity and lower fragmentation along the river compared to the terrestrial zone promoted mass effects, and differences in overall dispersal ability and mode (i.e. active and passive) for instream and riparian communities shifted paradigms between mass effects and species sorting.     

Broad‐scale patterns of invertebrate richness and community composition in temporary rivers: effects of flow intermittence

Temporary rivers are increasingly common freshwater ecosystems, but there have been no global syntheses of their community patterns. In this study, we examined the responses of aquatic invertebrate communities to flow intermittence in 14 rivers from multiple biogeographic regions covering a wide range of flow intermittence and spatial arrangements of perennial and temporary reaches. Hydrological data were used to describe flow intermittence (FI, the proportion of the year without surface water) gradients. Linear mixed‐effects models were used to examine the relationships between FI and community structure and composition. We also tested if communities at the most temporary sites were nested subsets of communities at the least temporary and perennial sites. Taxon richness decreased as FI increased and invertebrate communities became dominated by ubiquitous taxa. The number of resilient taxa (with high dispersal capacities) decreased with increased FI, whereas the number of resistant taxa (with adaptations to desiccation) was not related to FI. River‐specific and river‐averaged model comparisons indicated most FI‐community relationships did not differ statistically among rivers. Community nestedness along FI gradients was detected in most rivers and there was little or no influence of the spatial arrangement of perennial and temporary reaches. These results indicate that FI is a primary driver of aquatic communities in temporary rivers, regardless of the biogeographic species pool. Community responses are largely due to resilience rather than resistance mechanisms. However, contrary to our expectations, resilience was not strongly influenced by spatial fragmentation patterns, suggesting that colonist sources other than adjacent perennial reaches were important.     

How does temporal variation in habitat connectivity influence metapopulation dynamics?

Metapopulation persistence depends on connectivity between habitat patches. While emphasis has been placed on the spatial dynamics of connectivity, much less has been placed on its short‐term temporal dynamics. In many terrestrial and aquatic ecosystems, however, transient (short‐term) changes in connectivity occur as habitat patches are connected and disconnected due, for example, to climatic or hydrological variability. We evaluated the implications of transient connectivity using a network‐based metapopulation model and a series of scenarios representing temporal changes in connectivity. The transient loss of connectivity can influence metapopulation persistence, and more strongly autocorrelated temporal dynamics affect metapopulation persistence more severely. Given that many ecosystems experience short‐term and temporary loss of habitat connectivity, it is important that these dynamics are adequately represented in metapopulation models; failing to do so may yield overly optimistic‐estimates of metapopulation persistence in fragmented landscapes.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.     

Mediterranean-climate streams and rivers: geographically separated but ecologically comparable freshwater systems

Streams and rivers in mediterranean-climate regions (med-rivers in med-regions) are ecologically unique, with flow regimes reflecting precipitation patterns. Although timing of drying and flooding is predictable, seasonal and annual intensity of these events is not. Sequential flooding and drying, coupled with anthropogenic influences make these med-rivers among the most stressed riverine habitat worldwide. Med-rivers are hotspots for biodiversity in all med-regions. Species in med-rivers require different, often opposing adaptive mechanisms to survive drought and flood conditions or recover from them. Thus, metacommunities undergo seasonal differences, reflecting cycles of river fragmentation and connectivity, which also affect ecosystem functioning. River conservation and management is challenging, and trade-offs between environmental and human uses are complex, especially under future climate change scenarios. This overview of a Special Issue on med-rivers synthesizes information presented in 21 articles covering the five med-regions worldwide: Mediterranean Basin, coastal California, central Chile, Cape region of South Africa, and southwest and southern Australia. Research programs to increase basic knowledge in less-developed med-regions should be prioritized to achieve increased abilities to better manage med-rivers.     

Drying as a primary hydrological determinant of biodiversity in river systems: a broad‐scale analysis

Determining and understanding relationships between biodiversity and hydrology is a critical goal in ecology, particularly given biodiversity in the freshwater realm is in crisis. Despite the prevalence of rivers experiencing natural drying disturbances (which we collectively refer to as intermittent rivers), and projections of increased frequency and duration of drying events, the importance of drying relative to other flow‐related determinants of river biodiversity remains understudied. We assessed the influence of drying on alpha‐ and beta‐diversity using discharge and macroinvertebrate data collated from Australia and southwest Europe over broad spatial and temporal scales, providing information on current and past drying events, and combining a wide variety of flow metrics. We found clear evidence that drying acts as a strong environmental filter and is a primary hydrological determinant of alpha‐diversity; even when considering both intermittent and perennial rivers, drying‐event conditions were its most important predictors. Macroinvertebrate richness declined with increasing durations of drying over the long‐term (Australia) and recent (Australia and southwest Europe) history of river discharge, and with decreasing predictability of event timing (Australia). Our analysis also revealed that: responses can be taxon specific due to variation in traits of resistance and resilience to drying; some taxa may respond just as or more strongly to variation in other discharge components (e.g. high‐ or low‐flow events) than to drying; and together these phenomena may result in differing community‐level responses within and across regions. Patterns of beta‐diversity across the wide biogeographical range of our study suggested that convergent and divergent niche‐selection processes may act in combination on aquatic communities of rivers experiencing drying disturbances. However, strong ability to disperse by flight (not by water) weakened beta‐diversity patterning among rivers. Our findings can be used to improve understanding of biodiversity organisation in disturbed systems, notably in those with dendritic features, including intermittent rivers.     

Low river flow alters the biomass and population structure of a riparian predatory invertebrate

1. Low flows in rivers are predicted to increase in extent and severity in many areas in the future, yet the consequent impacts of river drying on terrestrial communities via (i) changes to riparian microclimatic conditions and (ii) the identity and abundance of emerging aquatic insects available to riparian predators have not been quantified. 2. We investigated the influence of low river flow on a riparian fishing spider, Dolomedes aquaticus, in five New Zealand rivers containing permanently flowing and drying reaches and, in one river, along a longitudinal drying gradient. 3. The biomass of aquatic insects, potential prey for D. aquaticus, declined with low river flows while the abundance of potential terrestrial prey remained similar at all sites. In the replicate rivers, and along the longitudinal drying gradient, spider biomass was lower, and size classes were skewed towards more small and fewer large spiders, in drying sites. A desiccation experiment in the laboratory indicated high sensitivity of the spiders, with prey presence increasing spider survival. 4. Differences in the spatial distribution, biomass and population size structure of spiders were observed along the longitudinal drying gradient and disappeared within 16 days of the water returning to all sites. 5. In total, low river flow affected the biomass of D. aquaticus, as well as their size class structure and spatial distribution. This indicates that low river flows have the potential to affect adjacent terrestrial ecosystems.     

Benthic and hyporheic invertebrate assemblages along a flow intermittence gradient: effects of duration of dry events

1. A large proportion of the total river length on Earth comprises rivers that are temporary in nature. However, the effects of periodical dry events have received far less attention from ecologists than those of floods and low flows. 2. This study concomitantly examined the effects of flow intermittence on invertebrates from the streambed surface and from a depth of 30 cm in the hyporheic zone. Invertebrates were collected during 3 years in the Albarine River, France, before and after summer dry events from 18 sites (seven were perennial) distributed along a longitudinal flow intermittence gradient. 3. I predicted benthic and hyporheic density and taxonomic richness to decrease, and assemblage composition to shift from desiccation‐sensitive to desiccation‐resistant taxa with increased dry event duration. Second, I predicted benthic and hyporheic assemblages from sites that dried for longer periods to be nested subsets of assemblages from sites that dried for shorter periods. Last, I predicted a convergence in benthic and hyporheic assemblage composition with increasing duration of dry events, resulting from increased vertical migration of benthic taxa into the hyporheic sediments to cope with dry events. 4. Increased dry event duration in the Albarine River led to a decrease in both benthic and hyporheic density and taxonomic richness. Invertebrate assemblage composition shifted along the gradient of increasing flow intermittence, but broad taxonomic overlap between perennial and temporary reaches and nestedness patterns indicated that these shifts were because of the loss of taxa susceptible to drying rather than selection for desiccation‐resistant specialists. 5. Assemblage composition between benthic and hyporheic invertebrates diverged with increasing dry event duration, suggesting that the hyporheic zone did not act as a refuge during dry events in this river. 6. Quantitative studies on the relationships between ecology and intermittence are still rare but are needed to predict the consequences of future changes in flow intermittence. The relationships found in this study should be tested across a wide range of temporary rivers to better evaluate the generality of these findings.     

Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.

     

Hyporheic invertebrates as bioindicators of ecological health in temporary rivers: A meta-analysis

Worldwide, many rivers cease flow and dry either naturally or owing to human activities such as water extraction. However, even when surface water is absent, diverse assemblages of aquatic invertebrates inhabit the saturated sediments below the river bed (hyporheic zone). In the absence of surface water or flow, biota of this zone may be sampled as an alternative to surface water-based ecological assessments. The potential of hyporheic invertebrates as ecological indicators of river health, however, is largely unexplored. We analysed hyporheic taxa lists from the international literature on temporary rivers to assess compositional similarity among broad-scale regions and sampling conditions, including the presence or absence of surface waters and flow, and the regional effect of hydrological phase (dry channel, non-flowing waters, surface flow) on richness. We hypothesised that if consistent patterns were found, then effects of human disturbances in temporary rivers may be assessable using hyporheic bioindicators. Assemblages differed geographically and by climate, but hydrological phase did not have a strong effect at the global scale. However, hyporheic assemblage composition within regions varied along a gradient of higher richness during wetter phases. This indicates that within geographic regions, hyporheic responses to surface drying are predictable and, by extension, hyporheic invertebrates are potentially useful ecological indicators of temporary river health. With many rivers now experiencing, or predicted to experience, lower flows and longer dry phases owing to climate change, the development of ecological assessment methods specific to flow intermittency is a priority. We advocate expanded monitoring of hyporheic zones in temporary rivers and recommend hyporheic invertebrates as potential bioindicators to complement surface water assessments.     

Within-river gene flow in the hellbender (Cryptobranchus alleganiensis) and implications for restorative release

Understanding how populations are genetically and demographically connected is beneficial for species management, since gene flow and dispersal contribute to genetic diversity and population persistence. For hellbenders (Cryptobranchus alleganiensis), an aquatic salamander species experiencing dramatic declines in population size, fine-scale (i.e. within river) patterns of genetic diversity and gene flow are not well understood. Previous findings indicate that hellbenders are habitat specialists that exhibit extreme site fidelity and low vagility, suggesting that gene flow is restricted among the several, discrete habitat patches within a river. Using 15 polymorphic microsatellite loci and 497 hellbender samples from four Missouri rivers, we assessed fine-scale patterns of genetic diversity in order to infer population connectivity and aid in population management. Results indicate moderate levels of genetic variation (H_O = 0.66–0.78) with little differentiation among habitat patches (avg. F_ST = 0.002) and no evidence of isolation by distance. Our data suggest that hellbender gene flow has been extensive even among habitat patches separated by distances greater than >100 km. These results are useful for hellbender management, especially in terms of making informed decisions regarding restorative releases of captively propagated individuals.     

Diverse invertebrate fauna using dry sediment as a refuge in semi-arid and temperate Australian rivers

Dormant aquatic invertebrates can remain viable in riverbed sediment during dry phases, forming a source for recolonisation during wet periods. Regional differences in capacity for invertebrates to survive drying in this way are poorly understood, but may indicate regional differences in vulnerability to altered flow regimes. We compared diversity of invertebrates in dry sediment from intermittent rivers in temperate and semi-arid Australia after 4–8 weeks of drying. We predicted adaptations of semi-arid biota to severe and unpredictable drying would make dry sediment a more significant recolonisation source, with higher relative diversity when compared with temperate rivers. Emerging aquatic invertebrate assemblages were compared to those sampled in nearby pools, as a common drying refuge. Relative taxa richness in rehydrated sediments was higher in the semi-arid region (83 ± 16% of pool taxa) than the temperate (47 ± 6% of pool taxa), despite lower overall richness (24 taxa in semi-arid, 32 taxa in temperate). Semi-arid rivers had greater potential for dry riverbeds to act as a source for recolonisation, given high relative diversity and abundance in dry sediment, combined with the frequent absence of alternative refuges. However, dry riverbeds in both regions provided a significant short-term refuge for aquatic invertebrates.     

The boundaries of river systems: the metazoan perspective

1. This overview of metazoans associated with the riparian/groundwater interface focuses on the fauna inhabiting substratum interstices within the stream bed and in alluvial aquifers beneath the floodplain. The objective is to integrate knowledge of habitat conditions and ecology of the interstitial fauna into a broad spatiotemporal perspective of lotic ecosystems. 2. Most aquatic metazoans of terrestrial ancestry, secondarily aquatic forms including insects and water mites (Hydracarina), are largely confined to surface waters (epigean), most of the time penetrating only the superficial interstices of the stream bed. 3. Primary aquatic metazoans include crustaceans and other groups whose entire evolutionary histories took place in water. Some species are epigean, whereas other members of the primary aquatic fauna are true subterranean forms (hypogean ) , residing deep within the stream bed and in alluvial aquifers some distance laterally from the channel. 4. The hypogean/epigean affinities of interstitial animals are reflected in repetitive gradients of species distribution patterns along vertical (depth within the stream bed), longitudinal (riffle/pool), and lateral (across the floodplain) spatial dimensions, as well as along recovery trajectories following floods (temporal dimension). 5. Fluvial dynamics and sediment characteristics interact to determine hydraulic conductivity, oxygen levels, pore space, particle size heterogeneity, organic content and other habitat conditions within the interstitial milieu. 6. Multidimensional environmental gradients occur at various scales across riparian/groundwater boundary zones. The spatiotemporal variability of hydrogeomorphological processes plays an important role in determining habitat heterogeneity, habitat stability, and connectivity between habitat patches, thereby structuring biodiversity patterns across the riverine landscape. 7. The erosive action of flooding maintains a diversity of hydrarch and riparian successional stages in alluvial floodplains. The patchy distribution patterns of interstitial communities at the floodplain scale reflect, in part, the spatial heterogeneity engendered by successional processes. 8. Interstitial metazoans engage in passive and active movements between surface waters and ground waters, between aquatic and riparian habitats, and between different habitat types within the lotic system. Some of these are extensive migrations that involve significant exchange of organic matter and energy between ecosystem compartments. 9. The generally high resilience of lotic ecosystems to disturbance is attributable, in part, to high spatiotemporal heterogeneity. Habitat patches less affected by a particular perturbation may serve as ’refugia ‘; from which survivors recolonize more severely affected areas. Mechanisms of refugium use may also occur within habitats, as, for example, through ontogenetic shifts in microhabitat use. Rigorous investigations of interstitial habitats as refugia should lead to a clearer understanding of the roles of disturbance and stochasticity in lotic ecosystems. 10. Development of realistic ’whole river ‘; food webs have been constrained by the exclusion of interstitial metazoans, which may in fact contribute the majority of energy flow in lotic ecosystems. A related problem is failure to include groundwater/riparian habitats as integral components of alluvial rivers. A conceptual model is presented that integrates groundwater and riparian systems into riverine food webs and that reflects the spatiotemporal complexity of the physical system and connectivity between different components. 11. Interstitial metazoans also serve as ’ecosystem engineers, ‘; by influencing the availability of resouces to other species and by modifying habitat conditions within the sediment. For example, by grazing on biofilm, interstitial animals may markedly stimulate bacterial growth rates and nutrient dynamics. 12. Although there has been a recent surge of interest in the role of interstitial animals in running waters, the knowledge gaps are vast. For example, basic environmental requirements of the majority of groundwater metazoans remain uninvestigated. Virtually nothing is known regarding the role of biotic interactions in structuring faunal distribution patterns across groundwater/riparian boundary zones. Interstitial metazoans may contribute significantly to the total productivity and energy flow of the biosphere, but such data are not available. Nor are sufficient data available to determine the contribution of groundwater animals to estimates of global biodiversity. 13. Effective ecosystem management must include groundwater/riparian ecotones and interstitial metazoans in monitoring and restoration efforts. Evidence suggests that a ’connected ‘; groundwater/riparian system provides natural pollution control, prevents clogging of sediment interstices and maintains high levels of habitat heterogeneity and successional stage diversity. River protection and restoration should maintain or re-establish at least a portion of the natural fluvial dynamics that sustains the ecological integrity of the entire riverine–floodplain–aquifer ecosystem. Keywords: groundwater/riparian ecotones, hyporheic habitat, epigean, hypogean, interstitial fauna, biodiversity, food webs     

Reduced breeding success of Pied Flycatchers Ficedula hypoleuca along regulated rivers

Most large rivers in northern Sweden are regulated to produce hydropower, with subsequent effects on flow dynamics and aquatic insect communities. Several studies have shown that aquatic and terrestrial systems are intimately connected via the export of emergent aquatic insects, but few have assessed how human modifications of aquatic habitats may influence this connection. We compared breeding success of the insectivorous Pied Flycatcher Ficedula hypoleuca in near‐riparian upland forests along two regulated and two free‐flowing large rivers in northern Sweden over 3 years. The regulated rivers showed lower aquatic insect export to the surroundings, as a consequence of regulation‐induced loss of suitable aquatic insect habitats. Survival of Pied Flycatcher nestlings was 10–15% greater along the free‐flowing rivers. Females breeding near the free‐flowing rivers also started egg‐laying earlier and with greater synchrony than those at the regulated rivers, and showed a smaller decrease in weight during breeding than did females along the regulated rivers. However, there were no differences in occupation rate, clutch size or number of successfully hatched juveniles between regulated and free‐flowing rivers. As regulated rivers showed lower abundance of flying aquatic insects, which may also reduce the abundance of terrestrial invertebrate prey, regulation‐induced changes in the export of emergent aquatic insects may explain both directly and indirectly the observed reduction in Pied Flycatcher breeding success along regulated rivers. Large‐scale river regulation may therefore impair the breeding success of insectivorous birds through impacts on prey availability.     

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

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Response of wetland plant communities to inundation within floodplain landscapes

Summary Many floodplain wetlands in south‐eastern Australia have become isolated from the main river channel as a consequence of reduced high flows and associated flood events following river regulation. In the Central Murray region of south‐eastern Australia, many temporary wetlands would have received water once every five years or so, with large floods maintaining floodplain connectivity every decade, under natural conditions. Now, the River Murray is highly regulated and many of these wetland areas have not been flooded for periods of up to 30 years. Consequently, these wetlands are becoming degraded and the biodiversity of the area is in decline. From 2001–2003, 21 Black Box depression wetlands in the Central Murray region were each watered once. Plant communities in each wetland were monitored for changes in abundance (assessed as percentage cover) before and during the wetting and drying phases. Wetlands were watered during spring or early summer with the length of inundation ranging from 6 to 19 weeks. After watering, the percentage cover of native plant taxa and native plant functional groups in most wetlands increased. In general, there was a decrease in the percentage number of terrestrial plants present and an increase in the percentage cover of aquatic plants. Introduced species were a minor component. Although these wetlands are all located in the Central Murray region, individual wetlands developed plant communities that contained taxa specific to individual wetlands despite initial similarities. These results indicate that wetland plant biodiversity within the landscape can be promoted and maintained by ensuring there is a diversity of wetlands with varying flood regimes within the landscape.     

Terrestrial origin of bacterial communities in complex boreal freshwater networks

Bacteria inhabiting boreal freshwaters are part of metacommunities where local assemblages are often linked by the flow of water in the landscape, yet the resulting spatial structure and the boundaries of the network metacommunity have never been explored. Here, we reconstruct the spatial structure of the bacterial metacommunity in a complex boreal aquatic network by determining the taxonomic composition of bacterial communities along the entire terrestrial/aquatic continuum, including soil and soilwaters, headwater streams, large rivers and lakes. We show that the network metacommunity has a directional spatial structure driven by a common terrestrial origin of aquatic communities, which are numerically dominated by taxa recruited from soils. Local community assembly is driven by variations along the hydrological continuum in the balance between mass effects and species sorting of terrestrial taxa, and seems further influenced by priority effects related to the spatial sequence of entry of soil bacteria into the network.     

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.