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INSTREAM FLOWS TO ASSIST THE RECOVERY OF ENDANGERED FISHES 25 <br />zoobenthos communities are more productive <br />on cobble bars than sand, but substratum size <br />on river bars is highly variable as a function of <br />the dynamic sediment transport and deposi- <br />tion processes that occur as the river fluctuates <br />between peak and base flows (Fig. 2). Another <br />example, though not well documented, is the <br />propensity for high benthic and planktonic pro- <br />duction in subchannels (backwaters) and flood- <br />plain wetlands that were (predam) seasonally <br />flooded. These different, yet interactive, space <br />and time scales that produce natural biophysi- <br />cal variation are the essence of the ecosystem <br />in which the endangered fishes evolved and <br />must be documented thoroughly. <br />4. Studies in the Upper Colorado River Basin <br />indicate that flow regulation, specifically re- <br />duction of the amplitude between peak and <br />base flows, is a likely contributor to the decline <br />of the native fishes, but the cause and effect <br />relationship is not simple. For example, years <br />of regulated flows, coupled with construction of <br />revetments, seem to have reduced the avail- <br />ability of backwaters and wetlands as nursery <br />habitats that support larval and juvenile <br />squawfish. Although extremely high flows <br />seem to be associated with weak cohorts of <br />Colorado River squawfish and humpback chub, <br />occasional extreme flooding needed to main- <br />tain channel morphology and channel-flood- <br />plain interactions probably is critical for long- <br />term survival of the fishes. Indeed, the only <br />recent incident of successful recruitment of <br />adult razorback sucker occurred when high <br />flows reconnected riparian gravel pits to the <br />mainstem Colorado River. On the other hand, <br />squawfish recruitment can occur over a wide <br />range of spring flows, and squawfish spawning <br />may be much less site-specific than is indicated <br />by the literature, or a wide range of preferred <br />spawning conditions exists on the spawning <br />bars where squawfish are routinely found (e.g., <br />Cleopatra's couch bar on the Yampa, Three <br />Fords on the Green). Presence of nonnative <br />predators and reduced complexity of habitats <br />needed by the different life history stages of the <br />endangered fishes (due to severing of channel- <br />floodplain connections and encroachment of ri- <br />parian vegetation into the channel) further <br />confound determination of cause and effect. <br />The fundamental problem with respect to pro- <br />vision of flows to recover the endangered fishes <br />is balancing the many interactive effects in a <br />manner that will favor the native fishes over <br />the long term (i.e., decades). <br />5. The life histories of the endangered fishes, as <br />well as those of zoobenthos that also have been <br />studied in detail, are either directly or indi- <br />rectly controlled by flow magnitude and timing <br />and the relation between flow and tempera- <br />ture. However, relationships between flow, <br />channel configuration, and thermal heteroge- <br />neity (cf., Ward 1984) have not been well inte- <br />grated conceptually or empirically or in the <br />context of the various life history stages of the <br />fishes. A squawfish life history energetics <br />model, for example, would be very helpful in <br />this regard. <br />6. Stream regulation has introduced serial discon- <br />tinuities (i.e., downstream extension of coldwa- <br />ter or rhithron environments) within the river <br />continua of the Upper Colorado River Basin. <br />The location and persistence of these disconti- <br />nuities are directly related to flow and largely <br />determine where the endangered and other <br />native fishes can achieve a positive life history <br />energy balance (i.e., complete the life history <br />with net recruitment of young at or above mini- <br />mum viable population size). Remember, these <br />fishes are adapted to potamon conditions, and <br />the length of the potamon zone has decreased <br />as a consequence of the downstream extension <br />(discontinuity) of the rhithron zone through <br />regulation of flow from the deep storage reser- <br />voirs. The concept of ecosystem "resets" and <br />discontinuities (sensu Ward and Stanford <br />1983), coupled with the notion that connected <br />channel and floodplain (backwaters, wetlands) <br />components of the riverscape are seasonally <br />pulsed by flooding (Ward 1989), robustly inte- <br />grates the myriad biophysical processes that <br />are influenced by stream regulation. Strong <br />inferences about how a river ecosystem may <br />respond to alternative flow management ac- <br />tions must be derived in this ecosystem con- <br />text. The downstream shift in the position of <br />the rhithron-potamon transition is an ecosys- <br />tem-level measure of change wrought by regu- <br />lation and should be used to adjust flows to <br />maximize conditions known to be favorable to <br />potamon (e.g., endangered fishes) and rhithron <br />(e.g., trout) fisheries. <br />7. Strong food web interactions are probably oc- <br />curring as a consequence of the presence <br />of a wide variety of nonnative fishes, which <br />now dominate fish communities throughout