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<br /> <br />402 <br /> <br />J, A. STANFORD ET At. <br /> <br />As a consequence of reservoir storage of peak flows for flood control, navigation, irrigation and hydro- <br />power production, baseflows increase substantially and often fluctuate so erratically that aquatic biota can- <br />not survive in shallow, near-shore habitats. The varial zone, shown in Figure I, constricts owing to loss of <br />peak flows and is depopulated by cyclic dewatering and rewatering that occurs on weekly, daily or even <br />hourly schedules (Cushman, 1985; 10urdonnais and Hauer, 1993). In stark contrast, a natural river pulses <br />water on to often expansive floodplains within a range of variation that allows a diversity of aquatic and <br />riparian biota to exist in multiple successional stages in a complex array of habitats. Persistent shallow or <br />slack water habitats are especially important for the survival of early life history stages of fishes that cannot <br />survive in the strong currents of the channel thalweg. Storage of bedload in the reservoir and constant clear <br />water flushing downstream artificially depletes gravel and finer sediments in the tailwaters, causing armour- <br />ing of the bed with large cobble and boulder substratum (Simons, 1979), Large rocks eroded from the can- <br />yon walls and coarse bedload from tributaries jam the channel and increase the size of rapids over time, <br />because peak flows are insufficient to scour and transport the largest materials downstream (Dolan et aI., <br />1978). Channel constrictions and habitat simplification occur as the channel downcuts and riparian vegeta- <br />tion invades to the top of the varial zone in aggraded reaches, owing to loss of upstream sediment supply and <br />loss of scouring flood flows (Johnson, 1994; Church, 1995). <br />The general conclusion is that regulation creates a discontinuum of environmental conditions and severs the <br />connectivity of channel, groundwater, floodplain and upland components of the catchment ecosystem; habi- <br />tats for riverine biota become spatially homogenous, limited to the permanently wetted portion of the channel <br />thalweg that is dominated by conditions dictated by operations of upstream storage reservoirs (Figure 5), <br />Indeed. serial construction of low-head dams has converted virtually all the mainstems of the largest rivers <br />in USA, Europe, Sweden and Finland into shallow reservoir habitat that is neither truly lacustrine nor riverine, <br />2. Native biodiversity decreases and non-native species proliferate Native biodiversity almost always <br />decreases after regulation (Minckley and Deacon, 1991; Ward and Stanford, 1991; Moyle and Leidy, <br />1992: Stevens et ai" in press), as conceptualized in Figure 5 compared with Figure 2, Vital core <br />populations may be extirpated and satellite populations may become increasingly isolated by regulation <br />schemes. Moreover, for anadromous species of fish mortality resulting from passage through dams and <br />reservoirs on the mainstem may be selective for certain of the geographically diverse populations that use <br />the mainstem as a common migratory pathway, thereby reducing biodiversity and increasing the <br />probability of metapopulation extinction (Harrison and Quinn, 1989; Reiman and McIntyre, 1993), <br />Altered temperature patterns and continual export of very fine organic matter and dissolved nutrients, <br />coupled with simplification of the channel, stabilization of bottom substratum and loss of floodplain inun- <br />dation. promotes environmental conditions to which native species are poorly adapted, opening opportu- <br />nities for non-native plants and animals to establish robust populations (Stanford and Ward, 1986; Li et <br />at.. 1987; Pflieger and Grace, 1987; Bain et ai" 1988; Shannon et al., 1994). In some cases one or a few native <br />species are more abundant than they were before regulation (e.g. Poe et aI" 1991), But, the most pervasive <br />result of habitat change produced by regulation is the proliferation of non-native species, Non-native inver- <br />tebrates and fishes are consistently more abundant in regulated compared with unregulated river reaches (Li <br />et aI" 1987; Bain et ai" 1988). Native riparian plants cannot exist on dewatered floodplains, which opens <br />niches for exotic, dryland plants, Moreover, owing to loss of scouring flows, exotic and some native riparian <br />plants choke the periodically saturated area of the shoreline above the narrowed varial zone and exotic <br />hydrophytes usually invade and quickly dominate shallow water habitats (Decamps and Tabacchi, 1994; <br />Johnson, 1994). Explicit reasons for non-native proliferation in regulated rivers vary, but, in general, <br />non-natives are simply better competitors in the homogenous habitats of regulated rivers, plus the fact <br />that a wide array of non-natives have been purposely introduced into regulated rivers, <br />], Biophysical conditions reset pr.edictably in relation to influences of tributaries and as distance downstream <br />from the dam increases, The serial discontinuity concept (SDC) (Ward and Stanford, 1983, 1985b) explicitly <br />acknowledges the inherent connectivity of the river continuum and predicts that the conditions described <br />above will ameliorate downstream as a natural consequence of the biophysical energetics of rivers, The <br />spatial rate at which reset occurs and its manifestation relative to position within the river continuum <br />(Figure 5) is related to the limnological attributes (depth, volume, water retention time, trophic state) of <br />