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<br />J, A. STANFORD ET At.
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<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
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