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<br />2,500 <br />E <br />U <br />`r 2,000 <br />N <br />aNi 1,500 <br />t 1,000 <br />N <br />z <br />a 500 <br />c <br />0 <br />0 <br />co _ <br />Distance from mouth (km) <br />Figure 4. Longitudinal profile of bound- <br />ary shear stress in the McKenzie River <br />for a ten-year recurrence-interval flood. <br />spawning habitat to accommodate <br />the nests (which are known as redds) <br />of the female salmon that return to <br />spawn.' <br />The number of female salmon per <br />redd counted in a reach of the <br />McKenzie for the years 1970-1986 <br />ranges between 4 and 15, and aver- <br />ages approximately 8.5 (Figure 6). <br />If adequate spawning gravels were <br />available, the ratio of female salmon <br />to redds should be approximately <br />1:1. Because on average 8.5 times as <br />many females as redds are counted, <br />it is likely that spawning-gravel limi- <br />tations are resulting in redd super- <br />imposition (i.e., the building of one <br />redd on top of another). <br />Redd superimposition is a com- <br />mon source of density-dependent <br />mortality in salmonids (Hayes 1987, <br />McNeil 1964). When spawning grav- <br />els are saturated, subsequent spawn- <br />ers do not increase the total number <br />of eggs but merely replace those <br />already deposited with their own. If <br />there are sufficient spawners to satu- <br />rate the spawning gravels, the avail- <br />ability of spawning gravels deter- <br />mines the largest number of eggs <br />that can be seeded in a river, which <br />in turn can determine the equilib- <br />rium or average density at which the <br />population fluctuates-the fluctua- <br />tions are generally driven by den- <br />sity-independent mortality. <br />There appears to have been a <br />change in the population dynamics <br />of the salmon in the McKenzie from <br />the period 1945-1960 to 1969-1989 <br />(Figure 7). In addition to smaller <br />fluctuations in 1969-1986, the av- <br />' F. K. Ligon and W. E. Dietrich, 1994, manu- <br />script in preparation. <br />erage population size has decreased <br />approximately 50%, from 17,000 <br />to 8000. One of the two flood-con- <br />trol dams-Cougar Dam, built in <br />1963-eliminated access to a sig- <br />nificant amount of upstream spawn- <br />ing gravels (Howell et al. 1988). <br />The immediate reduction in the av- <br />erage population size after this re- <br />duction in spawning habitat is con- <br />sistent with our hypothesis that <br />spawning gravel limitations in the <br />McKenzie are the primary source of <br />density-dependent mortality. <br />As channel simplification due to <br />the reduced peak flows continues, <br />and further spawning gravels are <br />lost, we predict that the salmon <br />population will continue to decline. <br />The geomorphic evolution of the <br />McKenzie to a predominantly single- <br />thread channel is a gradual process, <br />occurring over decades. Because it <br />happens slowly, it goes unnoticed <br />even by fishing guides who have <br />been floating the river for many <br />years-the river still seems pristine. <br />Considerable research has been <br />conducted on the biology of McKen- <br />zie River salmon, such as examining <br />smolt mortality as a result of water <br />diversions. Variation in such mor- <br />186 BioScience Vol. 4S No. 3 <br />0 50 100 150 <br />Figure 5. This reach reflects the loss of islands between 1967 and 1990 in the <br />McKenzie River. River path is shown in dark blue for 1967 and in light blue for <br />1990. Islands in 1967 are green; there were no islands in 1990.