The fauna of dynamic riverine landscapes

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

Riverine landscape diversity

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

Riverine landscape dynamics and ecological risk assessment

1. The aim of ecological risk assessments is to evaluate the likelihood that ecosystems are adversely affected by human‐induced disturbance that brings the ecosystem into a new dynamic equilibrium with a simpler structure and lower potential energy. The risk probability depends on the threshold capacity of the system (resistance) and on the capacity of the system to return to a state of equilibrium (resilience). 2. There are two complementary approaches to assessing ecological risks of riverine landscape dynamics. The reductionist approach aims at identifying risk to the ecosystem on the basis of accumulated data on simple stressor–effect relationships. The holistic approach aims at taking the whole ecosystem performance into account, which implies meso‐scale analysis. 3. Landscape patterns and their dynamics represent the physical framework of processes determining the ecosystem's equilibrium. Assessing risks of landscape dynamics to riverine ecosystems implies addressing complex interactions of system components (e.g. population dynamics and biogeochemical cycles) occurring at multiple scales of space and time. 4. One of the most important steps in ecological risk assessment is to establish clear assessment endpoints (e.g. vital ecosystem and landscape attributes). Their formulation must recognise that riverine ecosystems are dynamic, structurally complex and composed of both deterministic and stochastic components. 5. Remote sensing (geo)statistics and geographical information systems are primary tools for quantifying spatial and temporal components of riverine ecosystem and landscape attributes. 6. The difficulty to experiment at the riverine landscape level means that ecological risk management is heavily dependent on models. Current models are targeted towards simulating ecological risk at levels ranging from single species to habitats, food webs and meta‐populations to ecosystems and entire riverine landscapes, with some including socio‐economic considerations.     

Aquatic invertebrates in riverine landscapes

1. Riverine systems consist of a mosaic of patches and habitats linked by diverse processes and supporting highly complex communities. Invertebrates show a high taxonomic and functional diversity in riverine systems and are in several ways important components of these systems. Their distribution patterns, movements and effects on ecological flows, testify to their importance in various landscape ecological processes. This paper reviews the invertebrate literature with respect to patterns and processes in the riverine landscape. 2. The distribution of invertebrates in riverine habitats is governed by a number of factors that typically act at different scales. Hence, the local community structure can be seen as the result of a continuous sorting process through environmental filters ranging from regional or catchment‐wide processes, involving speciation, geological history and climate, to the small‐scale characteristics of individual patches, such as local predation risk, substratum porosity and current velocity. 3. Dispersal is an important process driving invertebrate distribution, linking different ecological systems across boundaries. Dispersal occurs within the aquatic habitat as well as into the terrestrial surrounding, and also over land to other waterbodies. New genetic techniques have contributed significantly to the understanding of aquatic invertebrate dispersal and revealed the importance of factors such as physical barriers, synchrony of emergence and taxonomic affiliation. 4. Invertebrates affect the cycling of nutrients and carbon by being a crucial intermediate link between primary producers, detritus pools or primary consumers, and predators higher up in the trophic hierarchy. Suspension feeders increase the retention of carbon. The subsidies of aquatic invertebrates to the terrestrial ecosystem have been shown to be important, as are reciprocal processes such as the supply of terrestrial invertebrates that fall into the water. 5. Future studies are needed both to advance theoretical aspects of landscape ecology pertaining to the invertebrates in riverine systems and to intensify the experimental testing of hypotheses, for example with respect to the scaling of processes and to linkages between the terrestrial and aquatic systems. Another promising avenue is to take advantage of naturally steep environmental gradients, and of systems disturbed by humans, such as regulated rivers. By comparison with unimpaired reference sites, the mechanisms involved might be identified. The use of `natural' experiments, especially where environmental gradients are steep, is another technique with great potential.     

Metamorphosis in river ecology: from reaches to macrosystems

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A method for spatial freshwater conservation prioritization

1. Freshwater ecosystems are amongst the most threatened and poorly protected globally. They continue to be degraded through habitat loss, pollution and invading species and conservation measures are urgently needed to halt declining trends in their biodiversity and integrity.
2. During the past decade a suite of decision support tools and computational approaches have been developed for efficient and targeted conservation action in terrestrial or marine ecosystems. These methods may be poorly suited for planning in freshwater systems because connectivity in terrestrial and marine systems is typically modelled in a way unsuitable for rivers, where connectivity has a strong directional component.
3. We modify the conservation prioritization method and software, zonation , to account for connectivity in a manner better suited to freshwater ecosystems. Prioritization was performed using subcatchment/catchment-based planning units and connectivity was modified to have directional upstream and downstream components consistent with the ecology of our target species.
4. We demonstrate this modified method for rivers and streams in the southern North Island of New Zealand. Data included predicted occupancy from boosted regression tree models of species distributions for 18 fish species. The study area covered 2.1 million hectares and included 394 first- to fourth order catchment or subcatchment planning units.
5. Realistic modelling of connectivity had a major influence on the areas proposed for conservation. If connectivity was ignored, recommended conservation areas were very fragmented. By contrast, when connectivity was modelled, high priority conservation targets consisted of entire river basins or headwater subcatchments.
6. The proposed method serves as a starting point for the implementation of reserve selection methods in river ecosystems.     

景观生态学与退化生态系统恢复

退化生态系统的恢复是一项艰巨任务,它需要考虑到所要恢复的退化生态系统的结构,多样性和其动态的整体性和长期性。现在对于退化生态系统恢复研究已经要使生态学家们关注受损生态系统的理论和实际问题。退化生态系统恢复所面临的挑战是理解和利用生态演替理论来完成并加速恢复进程。恢复的主要目标是建立一个自维持的,由不同的群落或生态系统组成的能够满足不同需要如生物保护和粮食生产需要的景观。景观生态学关注于大的空间尺度的生态学问题。景观生态学研究方法可以为退化生态系统恢复实践提供指导。在解决退化生态系统的恢复问题时,景观生态学的方法在理论和实践上是有效的。景观生态学中的核心概念和其一般原理斑块形状、生态系统间相互作用、镶嵌系列等都同退化生态系统的恢复有着密切的关系。如恢复地点的选择和适当的恢复要素的空间配置。在评价退化生态系统的恢复是否取得成功,利用景观生态学也具有重要的意义。景观生态学理论如景观格局与景观异质性理论,干扰理论和尺度理论都能够指导退化生态系统的恢复实践。同样地,退化生态系统的恢复可以为景观生态学的研究提供非常恰当的实验场。寓景观生态学思想于退化生态系统恢复过程是一种新的有效途径。     

Habitat quality and connectivity in agricultural landscapes: The role of land use systems at various scales in time

Connectivity is a key concept of landscape ecology as it relates to flows and movements of organisms as driven by landscape structure. More and more aspects of landscape heterogeneity are considered in measuring connectivity, as the diversity of crops in agricultural landscapes. In this paper, we explored the value of considering changes and cumulated effects of connectivity over time. As an example, we analysed connectivity among patches influenced by maize over 7 years in an agricultural landscape in Brittany, France.Clear temporal patterns appeared: maize is concentrated in certain parts of the landscape, but over the period the whole area, 70% of the landscape, used for maize was connected. Instead of discrete patches, maize may produce large clusters allowing movement from patch to patch from year to year. This reinforces the importance of understanding land use allocation rules within farms and landscapes to evaluate the ecological effects of agriculture.     

景观生态学在近海资源环境中的应用——论海洋景观生态学的发展

长期以来,景观生态学研究主要集中于陆地景观生态研究,并在土地利用、植被退化等方面取得了长足的进展,而在海洋领域涉足颇少.论述了景观生态学在海洋赤潮景观、海洋溢油景观、海域使用景观、滨海湿地景观、海岛景观和海洋环境污染景观等方面的应用前景,并就海洋景观生态学发展几点认识进行了深入探讨,如海洋景观的均质性与异质性、海洋景观格局与生态过程的关系、边缘效应与海岸带、海洋景观评价与评估、海洋景观模型等,目的是为景观生态学在海洋资源环境中的应用、海洋景观生态学的发展探索新的方向.     

景观生态恢复与重建是区域生态安全格局构建的关键途径

生态恢复与重建是跨尺度、多等级的问题,其主要表现层次应是生态系统（生物群落）、景观,甚至区域,而不能仅仅局限于生态系统。景观的恢复与重建是针对景观退化而言,景观退化从表现形式上可分为景观结构退化与景观功能退化。景观结构退化即景观破碎化,是指景观中各生态系统之间的各种功能联系断裂或连接度(connectivity)减少的现象;而鲜受重视的景观聚集（aggregation）在很多情况下同样具有造成景观退化的负面效应。景观功能退化是指与前一状态相比,由于景观异质性的改变导致景观的稳定性与服务功能等的衰退现象。景观恢复是指恢复原生生态系统间被人类活动终止或破坏的相互联系;景观生态建设应以景观单元空间结构的调整和重新构建为基本手段,包括调整原有的景观格局,引进新的景观组分等,以改善受胁或受损生态系统的功能,提高其基本生产力和稳定性,将人类活动对于景观演化的影响导入良性循环。二者的综合,统称为景观生态恢复与重建,是构建安全的区域生态格局的关键途径。其目标是建立一种由结构合理、功能高效、关系协调的模式生态系统（model ecosystem）组成的模式景观（model landscape）,以实现生态系统健康、生态格局安全和景观服务功能持续,以3S(RS,GPS,GIS)技术为支撑的GAP(ageographic approach to protect biological diversity)分析将为大尺度景观恢复的诊断、评价、规划提供重要的手段。景观中某些关键性点、位置或关系的破坏对整个生态安全具有毁灭性的后果,研究景观层次上的生态恢复模式及恢复技术、选择恢复的关键位置、构筑生态安全格局已成为景观生态学家关注的焦点。     

景观生态学中生态连接度研究进展

生态连接度对生物迁移扩散、基因流动、干扰扩散等生态过程具有重要作用,是目前景观生态学研究的热点内容.生态连接度是测度景观对于资源斑块间运动的促进或者阻碍作用程度的指标.它主要基于渗透和图论两大理论,通过实验、模型模拟以及指数等量化方法描述区域景观结构和功能的变化,广泛应用在自然景观和城市景观格局优化中,对生物多样性保护以及城市开放空间规划具有重要作用.介绍生态连接度的理论基础、评价方法,应用以及主要结论,并对景观生态学中生态连接度的未来研究方向进行展望,以促进生态连接度研究的进一步发展.     

基于图论的景观图表达、分析及应用

景观结构和空间格局一直是景观生态学的核心问题,图论的应用为景观格局的分析提供了一种研究框架,基于图论的景观图逐渐被应用于生物多样性保护的连通性建模和景观规划的决策支持研究,景观图的表达、分析和应用已成为保护生物学和景观生态学研究的热点。本文首先介绍了景观图的图论基础,在Scopus数据库的基础上,检索了1993—2019年在标题、摘要和关键词中出现 “landscape graph”、“connectivity”和“network”词汇的257篇已发表的期刊论文。从年发文量、来源期刊、研究区域、研究机构、景观类型等方面分析了该领域的研究进展和发展趋势。分析发现,2017年之前,发表的期刊论文数量整体呈上升趋势,2017年之后年发文量逐年下降;主要研究力量集中在美国、西班牙、法国、加拿大和中国,发文量占到86.8%。大部分研究成果发表在“Landscape Ecology”、“Landscape and Urban Planning”和“Biological Conservation”期刊上。在研究内容上,景观图表达主要包括点的定义、边的度量和景观的模拟3方面,景观图分析研究包括分析指数、景观图划分两方面。我们重点关注了景观图在生物多样性保护、景观(生态网络)规划和管理、景观影响评价等科学与实践中的应用。基于图论的景观图通过帮助理解景观连通性变化、动物行为和栖息地保护,影响着保护科学和规划实践者。图论对保护科学和规划的影响来自于丰富的理论基础和成熟的研究方法,基于图论的景观图为景观结构和格局的生态学理解提供了跳板,并将继续成为全球研究人员和实践者的重要工具。     

Hydrological connectivity for riverine fish: measurement challenges and research opportunities

1. In this review, we first summarize how hydrologic connectivity has been studied for riverine fish capable of moving long distances, and then identify research opportunities that have clear conservation significance. Migratory species, such as anadromous salmonids, are good model organisms for understanding ecological connectivity in rivers because the spatial scale over which movements occur among freshwater habitats is large enough to be easily observed with available techniques; they are often economically or culturally valuable with habitats that can be easily fragmented by human activities; and they integrate landscape conditions from multiple surrounding catchment(s) with in‐river conditions. Studies have focussed on three themes: (i) relatively stable connections (connections controlled by processes that act over broad spatio‐temporal scales >1000 km² and >100 years); (ii) dynamic connections (connections controlled by processes acting over fine to moderate spatio‐temporal scales ～1–1000 km² and <1–100 years); and (iii) anthropogenic influences on hydrologic connectivity, including actions that disrupt or enhance natural connections experienced by fish. 2. We outline eight challenges to understanding the role of connectivity in riverine fish ecology, organized under three foci: (i) addressing the constraints of river structure; (ii) embracing temporal complexity in hydrologic connectivity; and (iii) managing connectivity for riverine fishes. Challenges include the spatial structure of stream networks, the force and direction of flow, scale‐dependence of connectivity, shifting boundaries, complexity of behaviour and life histories and quantifying anthropogenic influence on connectivity and aligning management goals. As we discuss each challenge, we summarize relevant approaches in the literature and provide additional suggestions for improving research and management of connectivity for riverine fishes. 3. Specifically, we suggest that rapid advances are possible in the following arenas: (i) incorporating network structure and river discharge into analyses; (ii) increasing explicit consideration of temporal complexity and fish behaviour in the scope of analyses; and (iii) parsing degrees of human and natural influences on connectivity and defining acceptable alterations. Multiscale analyses are most likely to identify dominant patterns of connections and disconnections, and the appropriate scale at which to focus conservation activities.     

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.     

景观生态网络研究进展

作为生态学重要的概念与方法,生态网络是景观生态学研究的热点问题,也是耦合景观结构、生态过程和功能的重要途径。景观生态网络对于保护生物多样性、维持生态平衡、增加景观连接度具有重要意义。从景观生态网络的相关理论、研究进展、研究方法模型等进行分析,并对其应用前景进行展望,主要介绍了传统景观格局分析、网络分析、模型模拟等方法的适用性与特点,并分析了景观生态网络在城市景观格局优化、自然保护区规划、生物多样性保护、土地规划等领域的应用,最后提出了研究的主要问题。     

Restoration strategies for river floodplains along large lowland rivers in Europe

1. Most temperate rivers are heavily regulated and characterised by incised channels, aggradated floodplains and modified hydroperiods. As a consequence, former extensive aquatic/terrestrial transition zones lack most of their basic ecological functions.
2. Along large rivers in Europe and North America, various floodplain restoration or rehabilitation projects have been planned or realised in recent years. However, restoration ecology is still in its infancy and the literature pertinent to river restoration is rather fragmented. (Semi-) aquatic components of floodplains, including secondary channels, disconnected and temporary waters as well as marshes, have received little attention, despite their significant contribution to biological diversity.
3. Many rehabilitation projects were planned or realised without prior knowledge of their potential for success or failure, although, these projects greatly contributed to our present understanding of river–floodplain systems.
4. River rehabilitation benefits from a consideration of river ecosystem concepts in quantitative terms, comparison with reference conditions, historical or others, and the establishment of interdisciplinary partnerships.
5. We present examples from two large European rivers, the Danube and the Rhine, in which the role of aquatic connectivity has been extensively studied. The Danube delta with its diversity of floodplain lakes across an immense transversal gradient (up to 10 km) serves as a reference system for restoration projects along lowland sections of large rivers such as the Rhine in the Netherlands.     

Landscape Ecotoxicology and Assessment of Risk at Multiple Scales

Traditional approaches to ecotoxicology and ecological risk assessment frequently have ignored the complexities arising due to the spatial heterogeneity of natural systems. In recent years, however, ecologists have become increasingly aware of the influence of spatial organization on ecological processes. Landscape ecology provides a conceptual and theoretical framework for the analysis of spatial patterns, the characterization of spatial aspects of ecosystem function, and the understanding of landscape dynamics. Incorporating the insights of landscape ecology into ecotoxicology will enhance our ability to understand and ultimately predict the effects of toxic substances in ecological systems. Ecological risk assessments need to explicitly consider multiple spatial scales, accounting for heterogeneity within contaminated areas and for the larger landscape context. A simple simulation model is presented to illustrate the effects of spatial heterogeneity by linking an individual-based toxicokinetic model with a spatially distributed metapopulation model. Dispersal of organisms between contaminated and uncontaminated patches creates a situation where risk analysis must consider a spatial extent broader than the toxicant-contaminated area. In general, the addition of a toxicant to a source patch (i.e., a net exporter of individuals) will have a greater impact than the same toxicant addition to a sink patch (i.e., a net importer of individuals).     

Diversity in riverine metacommunities: a network perspective

The influence of spatial processes on diversity and community dynamics is generally recognized in ecology and also applied to conservation projects involving forest and grassland ecosystems. Riverine ecosystems, however, have been for a long time viewed from a local or linear perspective, even though the treelike branching of river networks is universal. River networks (so-called dendritic networks) are not only structured in a hierarchic way, but the dendritic landscape structure and physical flows often dictate distance and directionality of dispersal. Theoretical models suggest that the specific riverine network structure directly affects diversity patterns. Recent experimental and comparative data are supporting this idea. Here, I provide an introduction on theoretical findings suggesting that genetic diversity, heterozygosity and species richness are higher in dendritic systems compared to linear or two-dimensional lattice landscapes. The characteristic diversity patterns can be explained in a network perspective, which also offers universal metrics to better understand and protect riverine diversity. I show how appropriate metrics describing network centrality and dispersal distances are superior to classic measures still applied in aquatic ecology, such as Strahler order or Euclidian distance. Finally, knowledge gaps and future directions of research are identified. The network perspective employed here may help to generalize findings on riverine biodiversity research and can be applied to conservation and river restoration projects.