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.     

Vulnerability of stream biota to climate change in mediterranean climate regions: a synthesis of ecological responses and conservation challenges

Freshwater species worldwide are experiencing dramatic declines partly attributable to ongoing climate change. It is expected that the future effects of climate change could be particularly severe in mediterranean climate (med-) regions, which host many endemic species already under great stress from the high level of human development. In this article, we review the climate and climate-induced changes in streams of med-regions and the responses of stream biota, focusing on both observed and anticipated ecological responses. We also discuss current knowledge gaps and conservation challenges. Expected climate alterations have already been observed in the last decades, and include: increased annual average air temperatures; decreased annual average precipitation; hydrologic alterations; and an increase in frequency, intensity and duration of extreme events, such as floods, droughts and fires. Recent observations, which are concordant with forecasts built, show stream biota of med-regions when facing climate changes tend to be displaced towards higher elevations and upper latitudes, communities tend to change their composition and homogenize, while some life-history traits seem to provide biota with resilience and resistance to adapt to the new conditions (as being short-lived, small, and resistant to low streamflow and desiccation). Nevertheless, such responses may be insufficient to cope with current and future environmental changes. Accurate forecasts of biotic changes and possible adaptations are difficult to obtain in med-regions mainly because of the difficulty of distinguishing disturbances due to natural variability from the effects of climate change, particularly regarding hydrology. Long-term studies are needed to disentangle such variability and improve knowledge regarding the ecological responses and the detection of early warning signals to climate change. Investments should focus on taxa beyond fish and macroinvertebrates, and in covering the less studied regions of Chile and South Africa. Scientists, policy makers and water managers must be involved in the climate change dialogue because the freshwater conservation concerns are huge.     

Ecological status assessment in mediterranean rivers: complexities and challenges in developing tools for assessing ecological status and defining reference conditions

Rivers in mediterranean regions are subject to hydrological extremes. They range from highly stable, perennial ground- or snow-fed systems to highly ephemeral, unpredictable ones in semi-arid environments. Spatial and temporal complexity inherent in these systems presents challenges for ecological status assessment and defining reference conditions, particularly as many areas have been extensively transformed through anthropogenic activities. Temporal variability driven by sequential and predictable, seasonal events of flooding and drying accentuates the need to take season and/or hydrological period into account. Intermittent streams, which are common in mediterranean regions (med-regions) and which have aquatic communities distinct from perennial streams, are often not incorporated in bioassessment and present distinct challenges. Med-regions are also known for their high biodiversity and rates of endemism, as well as large numbers of introduced species. Med-regions are expected to be among the most affected by global climate change and, in these systems, climate change is an additional driver influencing ecosystems that are already stressed. From this review it is evident that an understanding of responses of indices, metrics, and models to climate change in comparison to existing stresses, and the development of thermally specific bioassessment tools are needed for this region.     

Disturbance and the role of refuges in mediterranean climate streams

Refuges protect plant and animal populations from disturbance. Knowledge of refuges from disturbance in mediterranean climate rivers (med-rivers) has increased the last decade. We review disturbance processes and their relationship to refuges in streams in mediterranean climate regions (med-regions). Med-river fauna show high endemicity and their populations are often exposed to disturbance; hence the critical importance of refuges during (both seasonal and supraseasonal) disturbances. Disturbance pressures are increasing in med-regions, in particular from climatic change, salinisation, sedimentation, water extraction, hydropower generation, supraseasonal drought, and wildfire. Med-rivers show annual cycles of constrained precipitation and predictable seasonal drying, causing the biota to depend on seasonal refuges, in particular, those that are spatially predictable. This creates a spatial and temporal mosaic of inundation that determines habitat extent and refuge function. Refuges of sufficient size and duration to maintain populations, such as perennially flowing reaches, sustain biodiversity and may harbour relict populations, particularly during increasing aridification, where little other suitable habitat remains in landscapes. Therefore, disturbances that threaten perennial flows potentially cascade disproportionately to reduce regional scale biodiversity in med-regions. Conservation approaches for med-river systems need to conserve both refuges and refuge connectivity, reduce the impact of anthropogenic disturbances and sustain predictable, seasonal flow patterns.     

Riparian vegetation research in Mediterranean-climate regions: common patterns, ecological processes, and considerations for management

Riparian corridors in Mediterranean-climate regions (med-regions) are resource-rich habitats within water-limited, larger landscapes. However, little is known about how their plant communities compare functionally and compositionally across med-regions. In recent decades, research on these ecosystems has expanded in both geographic scope and disciplinary depth. We reviewed 286 riparian-vegetation studies across the five med-regions, and identified common themes, including: (1) high levels of plant biodiversity, structural complexity, and cross-region species introductions; (2) strong physical controls on plant demographics and community structure; and (3) intensive human impacts. European and Californian ecosystems were the most represented among the studies reviewed, but Australia, South Africa, and Chile had the greatest proportional increases in articles published since 2000. All med-regions support distinct riparian flora, although many genera have invaded across regions. Plant species in all regions are adapted to multiple abiotic stressors, including dynamic flooding and sediment regimes, seasonal water shortage, and fire. The most severe human impacts are from land-use conversion to agriculture, streamflow regulation, nutrient enrichment, and climate change. Current knowledge gaps and subjects for future research include cumulative impacts to small, ephemeral streams and large, regulated rivers, as well as understudied ecosystems in North Africa, the western Mediterranean basin, and Chile.     

Pollution in mediterranean-climate rivers

This review examines information generated over the past decade on the pollution of rivers in regions with a mediterranean-type climate (med-climate). Pollution has clearly increased in the last 100 years and is correlated with the development of industry, agriculture and human population. Important efforts have been made in some med-climate countries in order to characterise the chemical status of rivers. In addition, the number of chemical substances detectable in mediterranean-climate rivers (med-rivers), as well as the limits of detection, have improved from the development of better analytical methods. New substances detected in rivers are gaining attention. We discuss available knowledge regarding real and potential effects of pollutants on the biota and ecosystems in med-rivers, taking into account natural environmental characteristics of these rivers. The extreme seasonal conditions in med-rivers add to the potential risk because these characteristics can enhance pollutant effects. Efforts and policies to prevent or reduce pollution effects on med-rivers are linked to the knowledge about pollution pressures associated with the degree of economic development. Aquatic communities in med-rivers are more sensitive to pollutants because they are exposed to strong natural and human stressors.     

Maintaining and restoring hydrologic habitat connectivity in mediterranean streams: an integrated modeling framework

Hydrologic alterations designed to provide a stable water supply and to prevent flooding are commonly used in mediterranean-climate river (med-rivers) basins, and these alterations have led to habitat loss and significant declines in aquatic biodiversity. Often the health of freshwater ecosystems depends on maintaining and recovering hydrologic habitat connectivity, which includes structural components related to the physical landscape, functionality of flow dynamics, and an understanding of species habitat requirements for movement, reproduction, and survival. To advance our understanding of hydrologic habitat connectivity and benefits of habitat restoration alternatives we provide: (1) a review of recent perspectives on hydrologic connectivity, including quantitative methods; and (2) a modeling framework to quantify the effects of restoration on hydrologic habitat connectivity. We then illustrate this approach through a case study on lateral hydrologic habitat connectivity that results from channel restoration scenarios using scenarios with different historic and climate-change flows to restore fish floodplain habitat in a med-river, the San Joaquin River, California. Case study results show that in addition to the channel alterations, higher flows are required to recover significant flooded habitat area, especially given reductions in flows expected under climate change. These types of studies will help the planning for restoration of hydrologic habitat connectivity in med-rivers, a critical step for mediterranean species recovery.     

Aquifer and surface-water ostracodes in Quaternary paleowetland deposits of southern Nevada,USA

Despite the importance of small tropical streams for maintaining freshwater biodiversity and providing essential ecosystem services to humans, relatively few studies have investigated multiple-stressor effects of climate and land-use change on these ecosystems, and how these effects may interact. To illustrate these knowledge gaps, we reviewed the current state of knowledge regarding the ecological impacts of climate change and catchment land use on small tropical streams. We consider the effects of predicted changes in streamflow dynamics and water temperatures on water chemistry, habitat structure, aquatic biota, and ecosystem processes. We highlight the pervasive individual effects of climate and land-use change on algal, macroinvertebrate, and fish communities, and in stream metabolism and decomposition processes. We also discuss potential responses of tropical streams in a multiple-stressor scenario, considering higher temperatures and shifts in hydrological dynamics. Finally, we identify six key knowledge gaps in the ecology of low-order tropical streams and indicate future research directions that may improve catchment management in the tropics to help alleviate climate-change impacts and biodiversity losses.     

An attack on two fronts: predicting how changes in land use and climate affect the distribution of stream macroinvertebrates

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Impacts of co-occurring environmental changes on Alaskan stream fishes

1. Freshwater fishes are now facing unprecedented environmental changes across their northern ranges, especially due to rapid warming occurring at higher latitudes. However, empirical research that examines co-occurring environmental effects on northern fish communities remains limited.
2. We used fish community data from 1587 Alaskan stream sites to examine the potential combined and interacting effects of climate change, current weather, habitat, land use, and fire on two community-level metrics (species richness, relative abundance), and on the distributions of three Alaskan fish species.
3. Our models were 71–76% accurate in predicting the distribution of Alaskan stream fishes using a combination of climate and habitat variables. In contrast to other freshwater ecosystems that are most threatened by land use pressures, we did not detect any evidence for the potential stress of anthropogenic land use or fire on stream fishes.
4. Warming temperatures increased overall community richness and abundance but produced differing responses at the species level. Juvenile salmon presence was positively associated with several climate variables including warmer spring and autumn temperatures and wetter summers. In comparison, warmer seasonal temperatures contributed to declines for northern-adapted species such as Arctic grayling and Dolly Varden.
5. This study highlights the overarching role of current and changing climate in regulating northern stream fish biodiversity. Although many fish species may benefit from climate change across their northern ranges, localised declines are likely to occur and may prove detrimental for communities with limited fishing portfolios. Climate change adaptation and mitigation strategies customised for rapidly changing northern ecosystems will play an essential role in preserving ecologically unique northern species.

     

Resource subsidies between stream and terrestrial ecosystems under global change

Streams and adjacent terrestrial ecosystems are characterized by permeable boundaries that are crossed by resource subsidies. Although the importance of these subsidies for riverine ecosystems is increasingly recognized, little is known about how they may be influenced by global environmental change. Drawing from available evidence, in this review we propose a conceptual framework to evaluate the effects of global change on the quality and spatiotemporal dynamics of stream–terrestrial subsidies. We illustrate how changes to hydrological and temperature regimes, atmospheric CO₂ concentration, land use and the distribution of nonindigenous species can influence subsidy fluxes by affecting the biology and ecology of donor and recipient systems and the physical characteristics of stream–riparian boundaries. Climate‐driven changes in the physiology and phenology of organisms with complex life cycles will influence their development time, body size and emergence patterns, with consequences for adjacent terrestrial consumers. Also, novel species interactions can modify subsidy dynamics via complex bottom‐up and top‐down effects. Given the seasonality and pulsed nature of subsidies, alterations of the temporal and spatial synchrony of resource availability to consumers across ecosystems are likely to result in ecological mismatches that can scale up from individual responses, to communities, to ecosystems. Similarly, altered hydrology, temperature, CO₂ concentration and land use will modify the recruitment and quality of riparian vegetation, the timing of leaf abscission and the establishment of invasive riparian species. Along with morphological changes to stream–terrestrial boundaries, these will alter the use and fluxes of allochthonous subsidies associated with stream ecosystems. Future research should aim to understand how subsidy dynamics will be affected by key drivers of global change, including agricultural intensification, increasing water use and biotic homogenization. Our conceptual framework based on the match–mismatch between donor and recipient organisms may facilitate understanding of the multiple effects of global change and aid in the development of future research questions.     

景观对河流生态系统的影响

从景观的视角研究河流生态系统,是目前河流生态学中受到广泛关注且发展迅速的内容之一。流域内多尺度景观强烈影响河流的理化及生物特征,已成为共识,但有关量度、整合景观与河流生态系统二者之间联系的理论体系与方法的建立尚处于起步阶段。对景观组成与空间格局影响河流生态系统的机制与途径进行了系统总结,提出了该领域研究中的主要难题,即如何识别景观中人为因素和自然因素的协变现象,如何衡量多空间尺度景观对河流生态系统的交互影响,如何理解景观阈值的不确定性。为克服当前研究面临的困难与挑战,填补已有知识的不足,提出今后研究的重点方向:河流景观分类系统的改进;更具代表性时空数据的采集;新型景观指标的开发与应用;微观尺度数据与宏观尺度数据的整合等。     

Synergistic and antagonistic interactions of future land use and climate change on river fish assemblages

River ecosystems are threatened by future changes in land use and climatic conditions. However, little is known of the influence of interactions of these two dominant global drivers of change on ecosystems. Does the interaction amplify (synergistic interaction) or buffer (antagonistic interaction) the impacts and does their interaction effect differ in magnitude, direction and spatial extent compared to single independent pressures. In this study, we model the impact of single and interacting effects of land use and climate change on the spatial distribution of 33 fish species in the Elbe River. The varying effects were modeled using step‐wise boosted regression trees based on 250 m raster grid cells. Species‐specific models were built for both ‘moderate’ and ‘extreme’ future land use and climate change scenarios to assess synergistic, additive and antagonistic interaction effects on species losses, species gains and diversity indices and to quantify their spatial distribution within the Elbe River network. Our results revealed species richness is predicted to increase by 0.7–2.9 species by 2050 across the entire river network. Changes in species richness are likely to be spatially variable with significant changes predicted for 56–85% of the river network. Antagonistic interactions would dominate species losses and gains in up to 75% of the river network. In contrast, synergistic and additive effects would occur in only 20% and 16% of the river network, respectively. The magnitude of the interaction was negatively correlated with the magnitudes of the single independent effects of land use and climate change. Evidence is provided to show that future land use and climate change effects are highly interactive resulting in species range shifts that would be spatially variable in size and characteristic. These findings emphasize the importance of adaptive river management and the design of spatially connected conservation areas to compensate for these high species turnovers and range shifts.     

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.     

The effect of agriculture on the seasonal dynamics and functional diversity of benthic biofilm in tropical headwater streams

Tropical streams are one of the most endangered ecosystems in the world due to the constant pressures from human activities. Among these activities, agriculture represents a land use that is crucial for human development but also a key driver of stream degradation and biodiversity decline in the tropics. Against this background, we investigated indirect effects of agriculture (alterations in stream flow and nutrient availability) and climate characteristics (water temperature) on benthic biofilm communities in tropical streams (São Paulo State, Brazil). Three first‐order streams draining catchments dominated by agricultural land use (sugarcane for bioenergy production, pasture) with some remaining riparian forest were studied for 1 year. We focused on the relationships of benthic biofilm biomass, algal biomass, diatom community, and functional structure with streamflow dynamics, nitrate concentrations, and water temperature. Our results indicate that these biological responses were mainly mediated by flow and water temperature and not by resource availability in the studied headwater streams. This result could be explained by the heavy rains and elevated runoff generation in these tropical catchments under agricultural influence, which might override the known effects of nutrient enrichment on benthic biofilm communities. Considering forecast climate and land‐use changes in tropical streams, our findings may suggest potential shifts in benthic biofilm communities, with functional consequences for aquatic food webs in these environments. Abstract in Brazilian Portuguese is available with online material.     

Predicting Plant Diversity Patterns in Madagascar: Understanding the Effects of Climate and Land Cover Change in a Biodiversity Hotspot

Climate and land cover change are driving a major reorganization of terrestrial biotic communities in tropical ecosystems. In an effort to understand how biodiversity patterns in the tropics will respond to individual and combined effects of these two drivers of environmental change, we use species distribution models (SDMs) calibrated for recent climate and land cover variables and projected to future scenarios to predict changes in diversity patterns in Madagascar. We collected occurrence records for 828 plant genera and 2186 plant species. We developed three scenarios, (i.e., climate only, land cover only and combined climate-land cover) based on recent and future climate and land cover variables. We used this modelling framework to investigate how the impacts of changes to climate and land cover influenced biodiversity across ecoregions and elevation bands. There were large-scale climate- and land cover-driven changes in plant biodiversity across Madagascar, including both losses and gains in diversity. The sharpest declines in biodiversity were projected for the eastern escarpment and high elevation ecosystems. Sharp declines in diversity were driven by the combined climate-land cover scenarios; however, there were subtle, region-specific differences in model outputs for each scenario, where certain regions experienced relatively higher species loss under climate or land cover only models. We strongly caution that predicted future gains in plant diversity will depend on the development and maintenance of dispersal pathways that connect current and future suitable habitats. The forecast for Madagascar’s plant diversity in the face of future environmental change is worrying: regional diversity will continue to decrease in response to the combined effects of climate and land cover change, with habitats such as ericoid thickets and eastern lowland and sub-humid forests particularly vulnerable into the future.     

Taxonomic and biological trait differences of stream macroinvertebrate communities between mediterranean and temperate regions: implications for future climatic scenarios

Streams in mediterranean regions have highly seasonal discharge patterns, with predictable torrential floods and severe droughts. In contrast, discharge is less variable in temperate regions and intermittent flow conditions are uncommon. Hydroclimatic models predict that climate change would increase frequency and severity of floods and droughts across Europe, thus increasing the proportion of streams with mediterranean characteristics in actually temperate areas. Correspondingly, understanding actual ecological differences between mediterranean and temperate streams may help to anticipate large‐scale ecological impacts of climate change. Given that large‐scale factors determine local community composition, we hypothesized that climatic differences between mediterranean and temperate regions should affect the taxonomic and biological trait composition in streams. We assembled the abundance of stream macroinvertebrate genera of 265 sites each from the Mediterranean Basin and from temperate Europe and linked these abundances to published information on 61 categories of 11 biological traits reflecting the potential of resilience from and resistance to disturbances. Although regional taxonomic richness was higher in the mediterranean than in the temperate region, local taxonomic richness and diversity did not significantly differ between regions. Local trait richness and diversity were significantly higher in the mediterranean region. Both local taxonomic and trait‐community composition differed between regions, but the former varied much more than the latter, highlighting that climate change could produce large changes in the taxonomic but rather weak changes in the trait composition. The mediterranean region was characterized by macroinvertebrates with higher dispersion and colonization capabilities, suggesting that species loss in the temperate region, by extinction or northward emigration of taxa, would be compensated for by immigration of southern mediterranean taxa. Thus, climate change would likely have stronger implications for the local conservation of taxa than for the trait composition of stream macroinvertebrate communities.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Evaluation of Gangetic dolphin habitat suitability under hydroclimatic changes using a coupled hydrological-hydrodynamic approach

The aquatic ecosystems are an integral part of the global environment and play a critical role in hydrogeological processes in the rivers. In recent decades, the external stressors on the aquatic species have significantly increased due to hydrologic alterations, human activities, and anthropogenic changes to their natural habitat. Global climate change has led to rivers' hydrological flow regime shifts, leading to unsuitable habitat conditions. It is, therefore, crucial to assess the potential implications of climate change on habitat suitability to ensure the long-term sustainability of freshwater species. In this direction, we investigated the association between endangered Gangetic dolphin populations and climate-driven hydrologic flow regime alterations in the Kulsi river, India. We developed a coupled eco-hydraulic framework comprising hydrological and hydrodynamic modeling to study the impact of past and projected climate change scenarios on the habitat suitability of target species. The framework was tested on a 15-km stretch of the Kulsi River, where the dolphin population has significantly declined in recent years. The temporal changes in the Weighted Usable Area (WUA) were analyzed using flow parameters and habitat suitability curves. Our findings suggest that the dolphin population decline coincided with a decrease in WUA, indicating a strong association between flow regimes and habitat suitability. Under climate change scenarios, multi-model climate projections and hydrological-hydrodynamic simulations show a rising trend in precipitation and streamflow in the basin, with substantial uncertainty. Higher flow depth and velocity would enhance WUA (habitat suitability). Still, the proposed river development projects in upstream regions could pose a serious threat to fragile dolphin communities by changing the seasonal flow patterns. The findings of this study can be included in conservation action plans and flow regulations strategies in upstream projects to ensure the long-term survival of endangered species.     

Environmental determinants of woody and herbaceous riparian vegetation patterns in a semi-arid mediterranean basin

What environmental variables determine riparian vegetation patterns? Are there differences between woody and herbaceous species? To answer these questions, we first explored the composition and richness patterns of both riparian woody and herbaceous species in a semi-arid mediterranean basin. Then, we assessed the environmental factors (climate, geology, topography, hydrogeomorphology and land use) that best explain these patterns. We used the following methodological approaches: clustering analyses, distance-based linear models, generalised linear models and hierarchical partitioning procedures. Valley shape, drought duration, river habitat heterogeneity, water conductivity and agricultural land use were the most important variables explaining variation in species composition for both groups. Woody riparian richness was mainly influenced by flow conditions and valley shape, whereas herbaceous one was more dependent on substrate features. Thus, although some differences in the importance of individual variables were observed, we found a notable congruence in the composition and species richness of both groups and also in the main types of variables explaining these patterns (hydrogeomorphology and land use, especially agriculture). Our results show that both communities could be treated in a holistic way, since they respond similarly to the strong natural and anthropogenic environmental gradients present in mediterranean basins.