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..:. <br />~~ - - <br />ECOLOGY 25 <br />le. Both populations and <br />e range of environmental <br />'rsh. Each species has its <br />its habitat (or better, its <br />ality members of a species <br />limits, in habitats that <br />duction (realized niche). <br />s from one stage of its life <br />knge occur in anadromous <br />young in streams but as <br />r, if more subtle, habitat <br />understand the ecological <br />age of its life history have <br />sh species are many, but <br />iient and flow, dissolved <br />re, barriers to movement, <br />populations, and aquatic <br />ctors .as pollution, dams, <br />gated systems, and the <br />e also become important <br />tors, such as temperature <br />t~ution than others, they <br />ct. Since different species <br />different ways to a given <br />it segregation on a broad <br />and niche segregation on <br />ties with similar environ- <br />and interact to subdivide <br />acies that occur together <br />distinct zones, each zone <br />'. The idea of zonation of <br />attempted to classify the <br />Hess (1) the Trout Zone, <br />streams; (2) the Grayling <br />Zone, dominated by grayling and trout, but with a few fast-water minnows present as <br />well, in cool, fairly fast, midelevation streams; (3} the Barbel Zone, dominated by <br />stream cyprinids with a few fishes from the -zones above and below, in slow, <br />wazmwater stretches of stream close to the valley floor; and (4) the Bream Zone, <br />dominated by deep~odied cyprinids and predatory fishes, such as pike and perch, in <br />the warm, sluggish waters of the valley floor. The concept of zonation is useful mostly <br />as a broad descriptive tool, since the exact composition of the fish fauna tends to vary <br />from stream to stream as a result of minor environmental differences and accidents of <br />distribution. Also, sharp boundaries between the zones are rare. The fish fauna changes <br />gradually as the nature of each stream shifts from swift headwaters to slow river. The <br />faunal change is generally less a replacement of one species by another than it is an <br />addition of species as the habitat becomes more complex. Far example, in Europe <br />trout are found not only in the Trout Zone but also in the Grayling and Barbel zones. <br />Despite these limitations, zonation remains a handy way to describe the fish fauna of <br />a region and its relationships to the different types of stream and lake habitats. In fact, <br />fish zones can be described quantiatively, as has been shown for the fishes of the <br />foothills above the San Joaquin Valley (Moyle and Nichols, 1973). There, 80 percent <br />of the stream localities sampled had a fish fauna that could be assigned by computer to <br />one of four fish associations (= zones). The following sections on the fish ecology of <br />each -major California drainage system will, therefore, attempt to describe the fish <br />zones both as they were before civilization and as they are today. They will also, <br />where possible, describe how the species that occur together in these zones subdivide <br />the habitat and its resources. <br />SACRAMENTO-SAN JOAQUIN SYSTEM <br />Despite the enormous size of this system, fish zones are surprisingly similar in its <br />various subsystems. The differences in fish fauna between subsystems indicate the <br />presence of zoogeographical barriers. Hardhead, for example, apparently were never <br />able to invade the Pajaro-Salinas subsystem nor were Sacramento blackfish able to <br />invade the Russian River subsystem. Both systems seem to have suitable habitats for <br />the missing species. Similarly, the slow-water fishes of the Central Valley (splittail, <br />blackfish, Sacramento perch, tule perch) have been unable to get over the falls on the <br />lower Pit River, although suitable habitats abound far them in its upper reaches. <br />Nevertheless, throughout the Sacramento-San Joaquin system five fish zones can be <br />recognized: (1) Rainbow Trout Zone, (2) California Roach Zone, (3) Squawfish- <br />sucker-hardhead Zone, (4) Deep-bodied Fishes Zone, and (5) Estuarine Fishes Zone. <br />Rainbow Trout Zone. This zone is found in -clear headwater streams where the <br />stream gradient is high (usually a total drop of 3 m or more for every km of stream). <br />The water is swift and permanent with more riffles than pools. The water is also cold, <br />seldom exceeding 21 °C, and is saturated with oxygen. The bottom material is <br />predominately cobbles, boulders, and bedrock. The banks are well shaded and <br />frequently undercut. Aquatic plants, submerged or emergent, are few except where the <br />streams flow through boggy alpine meadows. The dominant native fish is rainbow <br />trout but sculpin (usually riffle or Pit sculpin) and speckled dace are likely to be found <br />