We propose an integrated, heuristic model of lotic biocomplexity across spatiotemporal scales from headwaters to large rivers. This riverine ecosystem synthesis (RES) provides a framework for understanding both broad, often discontinuous patterns along longitudinal and lateral dimensions of river networks and local ecological patterns across various temporal and smaller spatial scales. Rather than posing a completely new model, we arrange a conceptual marriage of eco-geomorphology (ecological aspects of fluvial geomorphology) with a terrestrial landscape model describing hierarchical patch dynamics. We modify five components of this terrestrial model for lotic ecosystems: (1) nested, discontinuous hierarchies of patch mosaics; (2) ecosystem dynamics as a composite of intra-and inter-patch dynamics; (3) linked patterns and processes; (4) dominance of non-equilibrial and stochastic processes; and (5) formation of a quasi-equilibrial, metastable state. Our conceptual model blends our perspectives on biocomplexity with aspects of aquatic models proposed from 1980-2004.Contrasting with a common view of rivers as continuous, longitudinal gradients in physical conditions, the RES portrays rivers as downstream arrays of large hydrogeomorphic patches (e.g. constricted, braided and floodplain channel areas) formed by catchment geomorphology and climate. The longitudinal distribution of these patches, which are identifiable using standard geomorphic techniques, varies amongst rivers and is difficult to forecast above ecoregional scales. Some types of hydrogeomorphic patches may reoccur along this downstream passage. Unique ecological 'functional process zones' are formed by individual types of hydrogeomorphic patches because of physiochemical habitat differences which affect ecosystem structure and function.The RES currently includes 14 tenets predicting how patterns of individual species distributions, community regulation, lotic ecosystem processes, and floodplain interactions will vary over spatiotemporal scales, especially as they relate to the functional process zones formed by hydrogeomorphic differences in the river network. Copyright # 2006 John Wiley & Sons, Ltd.key words: eco-geomorphology; floodplains; flood pulse concept; headwaters; hierarchical patch dynamics; hydrogeomorphic patches; inshore retention concept; large rivers; river continuum concept; riverine productivity model HISTORICAL PERSPECTIVES AND PHILOSOPHICAL FOUNDATION Understanding the ecological structure and function of natural and/or altered lotic ecosystems is a common goal of many stream and river ecologists. This has spurred development of numerous conceptual models, shaped empirical research and funding, and occasionally altered government policies on river conservation, management and rehabilitation. Formation of conceptual theories can expand our knowledge of factors regulating river networks as long as popular theories are viewed as the 'latest best approximations' rather than iron-clad truths and if ecologists seek to test theorie...

Australian dryland rivers are acknowledged as being among the most variable and unpredictable in the world in terms of their flow regimes. Although renowned for their spectacular floods over vast and complex floodplains, rivers exist for much of the time as discrete waterholes, which are important refugia for aquatic biota. Recent work has shown that waterholes are filled by surface flows and there is little evidence of groundwater contributions. The permanence of these refugia is largely determined by waterhole morphology and evaporative loss, and some waterholes can clearly persist for 2 years or more without surface flow connection. As a consequence, the spatial distribution of refugia for aquatic biota is determined not only by the physical template but also by the duration of dry spells and the timing of flow events. Flow variability also has a major influence on aquatic production in these systems and ultimately influences food availability for fish and other consumers. During dry spells, aquatic food webs in waterholes are largely supported by algal production in the shallow littoral zone. At the other extreme, during floods, the boom of aquatic production on floodplains provides an immense food resource. However, there are many occasions when in-channel flows (flow pulses) result in periods where neither of these sources of production is likely to be available. Although such flow pulses are essential for the physical persistence and connectivity of waterholes, we propose that they may lead to food limitation and stress for populations of fish and other consumers. Water resource development in dryland rivers often leads to an increase in the frequency and duration of flow pulses, due to reduced floods and elevated base flows. This increase in the 'bits in between' natural boom or bust conditions may help to explain the observed decline in ecosystem health in dryland river systems with significant water resource development.

Spatial and temporal variation in fish-assemblage structure within isolated waterholes on the floodplains of Cooper Creek, Australia, was studied during the 2001 dry season, a period of natural drought in this arid-zone river. Spatial variation in fish-assemblage structure and the abundance of five species in disconnected waterholes early in the dry season (April 2001) were related to the extent of floodplain inundation 14 months previously, and to the interconnectedness of waterholes and waterhole habitat structure. As the dry season progressed, waterhole volumes decreased owing to evaporative water loss and structural habitat elements (anabranches, bars, boulders) became exposed. Marked changes in fish assemblage structure between the early (April) and late (September) dry season were related to habitat loss but not to water chemistry. Interactions between flow and habitat across a nested hierarchy of spatial scales (the floodplain, the waterhole and habitat patches within waterholes) were crucial to the persistence of fish assemblages through the 2001 dry season. We conclude that the magnitude, timing and frequency of floodplain inundation and natural variations in waterhole volume must be maintained if we wish to sustain the distinctive habitats and fish assemblages of this arid-zone floodplain river.

Summary 1. Traditionally the assessment of river water quality has been based solely on the measurement of physical, chemical and some biological characteristics. While these measurements may be efficient for regulating effluent discharges and protecting humans, they are not very useful for large‐scale management of catchments or for assessing whether river ecosystems are being protected. 2. Measurements of aquatic biota, to identify structural or functional integrity of ecosystems, have recently gained acceptance for river assessment. Empirical evidence from studies of river ecosystems under stress suggests that a small group of biological ecosystem‐level indicators can assess river condition. However, physical and chemical features of the environment affect these indicators, the structure and function of which may be changed by human activities. 3. The term ‘river health’, applied to the assessment of river condition, is often seen as being analogous with human health, giving many a sense of understanding. Unfortunately, the meaning of ‘river health’ remains obscure. It is not clear what aspects of river health sets of ecosystem‐level indicators actually identify, nor how physical, chemical and biological characteristics may be integrated into measures rather than just observations of cause and effect. 4. Increased examination of relationships between environmental variables that affect aquatic biota, such as habitat structure, flow regime, energy sources, water quality and biotic interactions and biological condition, are required in the study of river health.

No abstract