Life History, Status, and Distribution of Klamath River Chinook Salmon
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S. D. G. Life History, Status, and Distribution of Klamath River Chinook Salmon By Jafet Andersson INTRODUCTION The Klamath River Basin on the Oregon-California border is a diverse, unusual and important watershed (Figure 1). It is diverse in its geology, climate and vegetation; unusual in its geomorphology, and important from aspects as diverse as supporting endangered fish species, providing for American Indian tribes, and supplying farmers with water for irrigation. Geomorphologically, the basin stretches from high desert in the Upper Basin (above the Iron Gate Dam) characterised by large natural lakes and low relief, to the mountainous Lower Basin characterised by steeper terrain and higher topographic complexity. The Scott River watershed in specific has bedrock dominated upper reaches, a central alluvial valley, a west side with steep-gradient tributaries, an east side with low-gradient tributaries and a steep bedrock gorge just above the junction with the Klamath main stem [refer to (Sanchez, 2003) in this volume for more detail]. The mountainous Lower Basin interacts with the meteorological conditions dominated by Pacific storms to produce the characteristic spatial climate distribution in the region. The further from the ocean, the lower the precipitation as water is successively released (mostly as snow) over the mountains in the west. This pattern influences the densely forested Scott River watershed in that the west side is much more moist than the valley and the east side. The hydrograph is snowmelt dominated with the bulk of the discharge being released in the spring [refer to (Anderson, 2003 and Chambers, 2003) in this volume for more detail]. There are 19 native and 13 alien fish species in the Lower Klamath Basin (Mount and Moyle, 2003). Nearly all native species spend some of their life in salt water while most alien species are entirely freshwater based. Important fishes of the Lower Klamath include the federally recognised threatened coho salmon (Oncorhynchus kisutch), the declining steelhead (O. mykiss) and the largest and most abundant salmonid, the chinook salmon (O. tshawytscha). Due to the importance of anadromous1 fishes for tribal, sport and commercial fisheries there has been extensive support for restoration of these fish populations. Although focus has centred on the threatened coho, long-term holistic management of all native species Page 1 of 21 J. C. M. Andersson May 5, 2003 is needed to prevent additional species of being threatened, and to consider - at times counterintuitive - ecosystem interactions. For this purpose, the natural history of the chinook salmon (with focus on the Lower Klamath Basin above the Trinity River confluence) is reviewed in terms of its life history (the developmental history of an organism from birth to death) and status (the population size and spatial distribution in a temporal perspective). Figure 1. The Klamath River Basin ( source Klamath Basin fish and Water Management Symposium, http://www.humboldt.edu/~extended/klamath/klamathmap.html) 1 Anadromous fish spawn in freshwater but migrate to the sea and spend their adult life in the ocean Page 2 of 21 J. C. M. Andersson May 5, 2003 LIFE HISTORY Chinook salmon (Oncorhynchus tshawytscha) is an anadromous fish with a broad range of life history patterns. In general, it is characterised by a life cycle that begins in freshwater where the eggs are laid, where the alevins emerge and where the fry rear until they metamorphose into smolts (smoltification) in preparation for their ocean life as adults in the sea. The cycle is completed as the spawning adults migrate back into the rivers to lay their eggs (Healey, 1991). In the Klamath River Basin, there are, broadly speaking, two distinct chinook populations: the spring run2 and the fall run. Moyle (2002) indicates that a late fall run may also have existed, but it is either poorly documented or extinct. For management purposes, the National Marine Fisheries Services (NMFS) recognises two populations: the Southern Oregon and Coastal California ESU3 (including the Klamath chinook below the Trinity River confluence) and the Upper Klamath and Trinity Rivers ESU (including both the fall run and the spring run chinook salmon from the Klamath River based on their genetic similarity; Myers et al., 1998). The distinct differences between the spring run and the fall run life histories may, however, merit a managerial differentiation (Moyle, 2002). The spring run differ from the fall run in that the adults enter the river before they are ready to spawn and reside in deep pools for 2-4 months before they spawn whereas the fall run adults spawn closely after they reach their spawning destination (Moyle, 2002). In addition, the spring run juveniles remain in the streams for a year or longer before their seaward migration (from which the term "stream-type" originates), whereas the fall run juveniles are generally less than a year old before they migrate to the sea (from which the term "ocean-type" originates) (Healey, 1991). The spring run juveniles are, furthermore, more territorial due to their larger size resulting from their relatively longer pool residence time in comparison with the fall run juveniles (Moyle, 2002). Below follows a discussion of the different life history stages of chinook salmon with particular focus on general timing and habitat requirements for each stage. Egg An adult female lays between 2,000 and 17,000 eggs (averaging about 9 mm in diameter) depending, in part, on female size and geographic position such that larger females lay more eggs and females at lower latitudes lay fewer eggs (Healey, 1991; Myers et al., 1998).The 2 A run is a large group of fish migrating in order to spawn Page 3 of 21 J. C. M. Andersson May 5, 2003 eggs are laid down in a gravel spawning bed - also known as a redd - at the head of a riffle in several depressions created during a few days of digging by the female. At the time of deposition, usually more than one male release their sperm into the depressions which are subsequently covered by gravel (Allen and Hassler, 1986). Eggs are incubated for about 30 days in the fall and winter before they hatch (Nawa and Frissell, 1993). Figure 2. Chinook salmon egg (source Columbia Environmental Research Centre: http://www.cerc.cr.usgs.gov) Redds vary greatly in water depth, water velocity and depth of gravel overlaying the eggs (e.g. 5-720 cm, 10-189 cms-1, and 10-80 cm respectively), such that meaningful averages are hard to establish (Healey, 1991). Sub-gravel flow does, however, seem to be of overriding importance in the choice of redd site because of its role of bringing oxygen to and removing metabolic waste from the eggs (Allen and Hassler, 1986). It has been suggested that chinook eggs are more sensitive than other Pacific salmons to low sub-gravel flows due to the high surface-to-volume ratio of their relatively large eggs that thus require higher rates of oxygenation (Healey, 1991). Silver et al. (1963) noted that, at least at about 11ºC, oxygen levels during incubation were influencing the size of chinook at hatching even near oxygen saturation. A study conducted at Mill Creek, California indicated that mortality increased rapidly as oxygen concentration decreased (3.9% at 13 ppm and 37.9% at less than 5 ppm respectively) presumably because of decreased water flows through the gravel (Gangmark and Bakkala, 1960). The water flow through the gravel is affected by depth of water above it since more water increases the hydraulic head (the pressure) in the gravel (Allen and Hassler, 3 An ESU is an Evolutionary Significant Unit (Myers et al., 1998) Page 4 of 21 J. C. M. Andersson May 5, 2003 1986). Siltation is also an important determinant of gravel throughflow in that the silt clogs up the gravel and thus decreases the flow through it (Healey, 1991). Egg mortality is influenced by the temperature at which the eggs are incubated such that 3-16ºC constitutes the upper and lower limits for 50% pre-hatch mortality at constant temperatures. At variable temperatures, the mortality is reduced (Healey, 1991). Ideal incubation temperatures are 5.5-12.8 (McCullough, 1999). Hatching time is inversely proportional to temperature of incubation; at 16ºC e.g., eggs hatch after ca 32 days. Other sources of egg mortality include predation by other fish species and by invertebrates (e.g. oligochaete worms) if the eggs are not buried deep enough, shock from scouring and high concentrations of toxic chemicals (Allen and Hassler, 1986). All in all, these and many other factors in the chinook life cycle add up to the 1:1 ratio between adult female parent and adult female spawner in the next generation (Healey, 1991). Alevin Newly hatched chinooks that still have the yolk sac attached are termed alevins (Figure 3). Alevins emerge from the eggs in general between January and March (Trihey & Associates, 1996), but the timing varies from year to year. Alevins remain in the same habitat as the eggs (the redd) until the nutrients in the yolk sac are used up, which takes about 2-4 weeks (Nawa and Frissell, 1993). Figure 3. Chinook alevin (source Columbia Environmental Research Centre: http://www.cerc.cr.usgs.gov) In addition to the habitat requirements of the eggs, alevins need adequate amounts of water in the gravel surrounding them. In the event of both short recurrent, and prolonged single-event dewatering of the redds, development rate decreases and survival plummets. A Page 5 of 21 J. C. M. Andersson May 5, 2003 study found that the survival rate for alevins experiencing recurrent one hour dewaterings or a single six hour dewatering event was a mere 4% (Healey, 1991). Fry & Fingerling When the nutrients of the yolk sac are nearly gone, the chinook starts feeding on its own in the water column, develops neutral buoyancy and social behaviour (Allen and Hassler, 1986).