Its,

Er .11,31

The Biological and Technical Justification

for the

Flow Proposal

of the

Columbia Basin Fish and Wildlife Authority

Prepared jointly by the members of the Columbia Basin Fish and Wildlife Authority 2501 S.W. First Ave., Suite 200 Portland, Oregon 97201-4752

February, 1991 The Biological and Technical Justification

for the

Flow Proposal

of the

Columbia Basin Fish and Wildlife Authority

Prepared jointly by the members of the Columbia Basin Fish and Wildlife Authoritv 2501 S.W. First Ave., Suite 200 Portland, Oregon 97201-4752

Februarv, 1991 TABLE OF CONTENTS

LIST OF FIGURES LIST OF TABLES ...... V

SUMMARY ...... vi 1. BACKGROUND I A. Introduction 1 B. Historical Perspective 2 ...... II. IDENTIFICATION OF THE NEED FOR FLOW 8 A- Juvenile Salmonids 8 1. Migratory Behavior 9 a. Biology ...... 9 b. Control ...... of Migratory Behavior 10 ...... 2. Physiological Development @ ...... 12 a. Delay and Residualism 13 b...... Salinity Preference and Tolerance 15 c. "Windows" BioloQical ...... of Timeliness and Their Management Implications 16 ...... 3. Predation Losses and Flow 18 a. Time ...... Residence Effects on Predation Losses 18 b. Effects ...... Temperature on Predation Losses 19 c...... Water Velocity - 19 ...... 4. Conclusions ...... 20 B. Adult Salmonids 20 I. Migratory Behavior...... I I ')I a. Migratory Tiraina...... - 1 21 b. - ...... Bioenergetics of Spawning ...... the Migration )I c. Spawning and Incubation ...... 21 2. Flow Effects on Passage 23 a...... Reservoir Passage 23 b. Passage ...... 25 c. Reach Passage ...... 26 3. Water Quality Associated with Flows ?6 a. Temperature ...... 26 4. Conclusions ...... 28 C. Species Other 28 1. Biology ...... 29 Spawning...... a. 29 b. Larval Stage...... and Flow ...... 29

i TABLE OF CONTENTS (continued)

Liu

III. FLOW AND WATER VELOCT 31 A...... Water Particle Movement ...... 31 r I. Storage ...... Replacement Method - ...... 31 2. Average Velocity Method ...... 33 B. Smolt Movement 36 1...... Smolt vs. Water Particle Travel Time ...... 36 ...... C. Ways Increased to Meet Water Velocity Needs ...... 39 1. Reservoir Drawdown ...... 39 D. Conclusion ...... 40 IV. JUSTIFICATION OF FLOWS PRESENTED IN THE FLOW PROPOSAL 43 A. Fish Flow Objectives7@@ ...... 43 B. Justification for Flows 43 1. Spring (April I ...... 15) - June ...... 44 2. Summer 16 ...... (June - August 31) 5* 3. Fall ...... (September I - November 30) 4. Winter (December I March 30) ...... - ...... 53 C. Conclusions ...... 55 V. CITED REFERENCES ...... 56 LIST OF FIGURES Page

L Generalized effect of hydroelectric operations on mainstem flows at The Dalles, Oregon 5 ......

2. Travel time of juvenile spring chinook salmon from Lower Granite Dam to John Day Dam and the calculated water particle travel time as related to flows at Ice Harbor Dam 11 ......

3. Mean gill ATPase activity (p moles P, - mg Prot-' - h-1) of hatchery and wild steelhead during the spring migration, 1989 14 .

1 ......

4. Gill microsomal Na', K'- stimulated adenosine activity of yearling steelhead as a function of the time of year and temperature 17 ...... 5. Streatriflow and timing adult chinook of and sockeve salmon runs . 22 ...... 6. Hydrographs of daily average flow at Vernita Bar in the Hanford Reach of the mid-Columbia River (RM 392) during spawning, incubation and emergence of upriver fall chinook salmon ...... 24 7. Water particle travel time, in davs, through: (A) lower Snake River: (B) mid-Columbia River River; and (C) lower Columbia reservoirs 32 ...... 8. Water particle travel time, in days, through: (A) lower Snake River; (B) mid-Columbia River; and (C) lower Columbia River reservoirs ' showing pre-project (free-flowing) and post-project water movement . 34 ...... I ...... 9. Water particle velocity, in miles per hour (mph), for: (A) lower Snake River; (B) mid-Columbia River: and (C) lower Columbia River reservoirs. compared to pre-project water particle velocities 15 ...... @ 10. Water particle travel time, in days, from the head of Lower Granite reservoir to Bonneville Dam at various streamflows, compared to water particle travel time under pre-project river conditions. 37 . . @ ...... 11. Water particle travel time, predicted fish travel time, and observed fish travel time vs flow ...... 38 12. Water particle travel time, predicted fish travel time, and observed fish travet time vs flow for subyearling chinook salmon in John Day reservoirs 40 ...... 13. Water particle travel time, in days, for various strearnflows through Lower

Granite reservoir : 41 ...... 14. T evels of flow augmentation ...... 42

15. Travel Time, in days, of Snake River: (A) yearling chinook and (B) steelhead 46 ...... 16. Survival of Snake River: (A) yearling chinook and (B) steelhead 47 ......

iii LIST OF FIGURES (continued)

17. Travel time, in days, of Snake River: (A) yearling chinook and (B) steelhead 48 ...... 18. Survival of Snake River: (A) yearling chinook and (B) steelhead 49 ...... 19. Travel time, in days, vs flow in John Day Pool ...... 50 20. Smolt-to-adult (SAR) return of spring chinook and mean flow at Lower Granite Dam (April 20-May 30) for (A) Rapid River Hatchery and (B) Marsh Creek releases 1977-1987 52 .... 21. Snake River wild/natural "B' steelhead passage as a function of average September flow and temperature...... 54

iv LIST OF TABLES

Pa2e

1. Maximum consumption rates (smolts/predator/dav ) of northern squawfish upon @ salmonid prey at various temperatures ...... ')o 2. Fish flow in recommendations, kcfs: minimum instantaneous/dailv average 44 ......

v SUMMARY

The anadromous salmon the Columbia of River Basin are: chinook ( tshawYcha), coho (0. kisutch), sockeye (0. nerka), chum (0. keta) salmon, and stcelhead (0. mykiss). They have evolved over millennia to migrate from fresh water to salt water habitats in the higher water velocities that occurred during spring freshets. Part physiological of their transition to prepare them for downstream migration and saltwater entry includes a decrease in swimming ability, which aids rapid migration to salt water during the high velocities of the spring freshets. The timing of changes in physiological condition and higher flows combine to form a "biological window". This biological window is a limited time period during which a fish has the optimum physiological capability to survive the transition to salt water. The development the hydroelectric of system on the Columbia and Snake rivers dramatically increased the cross-sectional the river, area of decreasing the water velocities which occurred Nvith the natural runoff. In addition, the operation the hydroelectric of system shifted naturally high spring flows to the fall and winter months. The construction of mainstem hydroelectric projects also resulted in site specific project passage mortalities and delays. The development and operation of the hydrosystem completely changed the hydrologic conditions under which the salmon life history evolved. The Columbia Basin Fish and Wildlife Authority (Authority flow proposal will improve migratio n conditions within the it will in present system, but not result pre-impoundment migration conditions. The Authority proposal recommends flows that are realistic and achievable within the current system. These flow recommendations optimum levels. are not due to the inability to recreate pre-impouridinent conditions. However, they are given the adequate best available information at this time. Travel time is a key migrational characteristic reflecting the dynamics of the downstream migration of juvenile anadrornous salmonids. The physiological condition of sincilts chanaes over the time thev are migrating. Travel time determines whether the smolts arrive at the estuary during the 'biological window".

so they can successfully survive the is transition to salt water. Travel time inversely related to flow. With the present hydrosystern, even extremely high flows achieve cannot pre-dam water velocities. Therefore, pre- impoundment smolt travel times and migration conditions cannot be achieved. The objective of the flow is proposal to provide flows throughout the Columbia River Basin that will survival during critical life maximize stages of anadromous salmonids and other fish stocks. The recommended flows should move juveniles from freshwater to the estuary within the appropriate "biological window"; stimulate upstream migration of adult salmon; provide optimal spawning habitat and water conditions for survival emerging fry of and rearing juvenile fish; and should also allow for facility operations that provide safe project passage for migrating juvenile and adult fish.

vi LIACKGROUND A. Introduction

Pacific salmon (Oncorhynchus spp.) are anadromous fish that hatch from eggs laid in the gravel in freshwater rivers, lakes. streams, and After rearing in fresh water anywhere from a few days up to three

years, juvenile salmon (smolts) migrate downstream to the ocean where they grow to maturity before returning to their natal stream or lake to spawn and complete their life cycle.

When smolts migrate they rely primarily on passive transport by water currents, rather than actively swimming downstream. Therefore, their timely and successful seaward migration is closely linked to river flows, which determines how fast the smolts complete theirjourney. Migratory behavior and the physiological

ability to make the transition from freshwater to saltwater are expressed in smolts for only a limited time.

Hence, the fish must reach the saltwater environment of the estuary within this "biological window". The trait is saln!ionids. migration not unique to anadromous Manv animals, ranging from butterflies to caribou, travel vast distances within relatively short time periods or 'windows" in order to survive. Their migratory behavior is genetically determined as an adaptation to maximize survival, and delays in migration can reduce their survival as individuals. and as populations.

The Columbia River Basin is internationally known for its stocks of anadromous salmonids. The large geographic range and abundance of steelhead (0. mykiss) and chinock (0. tshawytcha), coho (0. kisutch), sockeye (0. nerka) and chum (0. keta) salmon were critical to the past econo mic growth of the region, and are important to the cultural needs and economic productivity in the future. Columbia When the and Snake rivers were in their natural state, the rivers' water velocitv was relatively high, particularly during the high in runoff periods the spring and ear1v summer. The timing of the smolts' seaward migration coincided with this increased runoff period (Mains and Smith 1964). It took migracoR, smolts only 22 days to travel from the Salmon River in Idaho to the lower Columbia River below Bonneville (Ebel 1977). Today, river is Dam the a series of reservoirs with large cross-sectional area with lower water velocity for moving smelts downstream. In addition, much of the peak spring and summer runoff is now captured in upper river storage reservoirs and released in the fall and winter months, which further reduces water velocity in the mainstem reservoirs. Thesmolts'journeyfromtheSalmonRivertothelowerColumbiaRivercannowtakeover5Odays(EbeI .

1977), roughly twice as long as in the past. During the record low runoff of 1977, victual travel time for from River smelts the Salmon to The Dalles Dam was 57 days. These fish arrived at The Dalles Dam in mid- June, than early May (Sims aL 1978). rather et Increases in migration time result in increases in exposure predators, which thrive in the to reservoirs. These delays also subject migratory fish to the higher water temperatures later in the year. It is speculated that these additional exposures adversely affect the health condition sarefts. Smolts and physiological of that are delayed and finaily do reach the estuary may no longer be physiologically fit to make the transition to saltwater, they may saltwater after the time when or enter . ocean conditions are most favorable for survival. 0 The state and federal fishery and wildlife agencies and Indian tribes (agencies and tribes) have lo ng recognized that adequate flows for migration critical are for survival and rebuilding of anadromous fish runs in the Columbia River Basin. The effects flow of river on smolt travel time and survival rate were established during the 1968 through 1983 period when last major the hydroelectric development of the Columbia and - Snake rivers was taking place. These investigations provided evidence that the development and operation ,r--of the hydroelectric projects significantly increased the travel time of smolts migrating to the ocean. These ' same studies also enabled biologists to conclude that flow was positively correlated with juvenile migrant survival. Higher rates of survival occurred in higher flow and lowest survival years occurred in low flow years. Information collected since 1983 through the Smelt Monitoring Program of the Columbia Basin Fish and Wildlife Authority (Authority) has confirmed these earlier data. No information has ever been collected that indicates the successful migration and survival higher of juvenile salmon can be achieved without adequate and timely flows.

Changes in river operations will continue to occur as a result of the Non-Treatv Storage Agreement. revision of the Canadian entitlement, renegotiation of power sales contracts, potential revisions to the Pacific Northwest Coordination Agreement, and firming up non-firm power. Additionally, changes in irrigati1010 withdraw will further shape flow patterns the patterns of the Columbia River. As a result, it will becom more and more difficult to achieve fish flow needs for spawning, rearing, and migration in the Columbia River Basin, unless the needs are incorporated into power system planning to guarantee they are met under any conditions. In recognition these of needs, and in response to a request from the Bonneville Power Administration, the agencies and tribes of the Authority identified bioicigicattv based Fish flow needs for salmon in a entitled, report "Proposed Mainstem Flows for Columbia Basin Anadromous Fish" (CBFWA 1990b). This document describes the biological and technical rationale for the Cishery tlow needs identified in the Authority's March 1990 proposal. This justification has been joirlily prepared by the member agencies and tribes of the Authoritv. B. Historical Prospective

The Columbia River Basin is the dominant drainage system in the Northwest region of the United States. Beginning in southwestern Canada, the Columbia River journeys almost 1.200, miles before entering the Pacific Ocean. The basin encompasses 219,000 square miles in the United States and 40,000 square miles in Canada. Major tributaries to the Columbia River include the Snake. Willamette, Yakima. Kootenai. Pend Oreille, Deschutes, and John Dav rivers.

Precipitation in the basin is concentrated as snow in the mountainous rezions durin(T the winter months. Consequently, winter streamflow is low generally (except in the tributaries West of the Cascade Mountain Range) followed by high runoff in the spring and early summer. The historical runoff pattern contributed approximately 60 per of the annual May, cent runoff during June and July. The Columbia River has an average annual runoff of 198 million acre-feet (MAF) at the mouth of the river (U.S. Army Corps of

Engineers 1985), and is second only in to the Missouri-Mississippi River system the United States in terms of average annual runoff. Prior to the development of the region the Columbia River supported the largest chinook and steethead populations in the world (Evans 1977). These fish anadromous stocks of the Columbia River Basin have been relied upon by the human, avian. and mammalian inhabitants of the region for thousands of years. The estimated annual catch by Native Americans in the basin prior to European settlement was 2.4 million fish (Hewes 1947). In the last half of the 1800's, salmon stocks major were natural resources for the growing economy of the area. In-river commercial, recreational, ceremonial. and subsistence fishery harvests have risen or fallen in concert with the salmon runs of the Columbia River. Ocean commercial and recreational fisheries from southern Oregon to Alaska also have benefitted from salmon (primarily chinook and coho) produced in the Columbia River Basin.

Since the settlement of the Columbia Basin in the mid 1800's, the water in the Columbia River has been used for transportation; mining; municipal and industrial use; and recreation. The water has been for by appropriated use agriculture (7.6 million acres irrigated in 1980) (CRWMG 1983), and more recently appropriated for in use power generation (34,000 megawatts annually (NPPC 1986). To enhance the use the river as a of transportation corridor, and develop the fertile and accessible flood plains of the basin, flood extensive control and navigation projects have been developed by the U.S. Army Corps of Engineers. The use and control the Columbia of River's water has meant the development of an, elaborate system of , locks. canals, storage reservoirs, water diversions, and power generation facilities. dams in Construction of began the 1880's on the Spokane and Willamette rivers, and by the early 1900's several dams small and hydropower projects were constructed throughout the basin (NPPC 1986). Many of these "run-of-the-river" with little were projects storage capability. Storage projects, or projects designed to retain and store runoff for later use were proposed in the 1930's by the U.S. Army Corps of Engineers in "308" their report, which detailed plans for developing the hydropower potential of the Columbia River. During the 1940's and 1950's, additional plans were developed to further harness the water resources of the Columbia. The driving forces behind these plans continued to be flood control and the ever increasing economic incentive to provide power in the postwar development era. The key to much of the development was an international agreement between the United States and Canada. the Columbia River Treaty (Treaty) which was signed in 1961 and ratified in 1964. The benefits of this Treary are power generation opportunities in Canada and the United States (2,800 megawatts) plus a vast amount of flood control storage (15.5 million acre-feet in Canada alone). Under terms of the Treaty Canada was paid approximately S65 million for the flood control benefits (8.5 MAF, with 7 an additional MAF available for additional cost). Canada was also entitled half the to power generation capability realized under the Treaty (approximately 1,400 megawatts). When the dams completed were and the Canadian reservoirs were filled in 1973 the storage capability of the

3 system was doubled, all the storage was convertible to power generation, and the high spring and surrinio flows were controlled.

Another significant milestone in the history of water development in the Columbia Basin was the concurrent formation of the Pacific Northwest Coordination Agreement. Created in 1964 to coordinate operation of the U.S. portion of the hydrosystem as a,"single utility" it provided guidelines and criteria describing how downstream benefits derived- from the 1964 Treary would be allocated. It also prescribed optimal use of the system's hydropower generation capability. The control of river runoff patterns has modified the physical characteristics of the river channel and the hydrograph of the river in comparison to historic patterns. Storage reservoirs and mainstem run-of-the-river reservoirs have drastically changed the water vtlocity, water depth, and water quality (in terms of temperature, turbidity and dissolved gases such as oxygen and nitrogen). These changes to the aquatic environment have affected the fishery resources in many wavs, which are not vet completely understood. An increase in the cross sectional profile the river of channel has increased the travel time for water through the system. By increasing riffles water depth, and cascades have become inundated. which decreases the river's ability to regulate dissolved gases in the This water. change in composition of pools and riffles also has a large impact on the fish species in found river reaches; in essence the reservoirs have created settling basins. As stated earlier and shown in Figure 1, the historic flow in pattern the Columbia Basin was high spring run-off (April through mid-June), decreasing summer and fall flows (mid-June through December), and low winter flows (December through March). Anadromous salmon have adapted to this flow pattern through the millennia by successfully migrating down the river as juveniles during high flows. The regulated flow transfers much the pattern of spring freshet (May through June) to the winter (November through March) with low flows in the summer and earlv fall (July through late Auaust). This regulated Llow pattern satisfies the power and flood control needs of the region, but alters the flow pattern for the fish from that in which anadromous fish adapted a to pattern that no longer matches the needs of the anadromous fish stocks native to the basin.

Pre-impoundment spring runoff afforded downstream hiLyh migrants water velocities, turbid water and large water volumes which diminished their vulnerability to predators and minimized their energy requirements to reach the Post-impoundment ocean. conditions have increased the smelt's exposure to predators by increasing travel time, iricreasin 9 water claritv and smelt visibility. These conditions have also increased the energy requirements of smolts to move through the reservoirs. Raymond (1968 and 1979) estimated the of migration for juvenile rate salmonids in a free flowing stream was 24 to 54 krn/day. on flow. depending For an impounded river the migration rates were 8 to 24 km/dav. Passage problems at the dams have also impacted the survival of juvenile salmonids. Mortality rates of juvenile salmonids at the various dams on the-Columbia and Snake rivers vary by species and flow conditions. Mortality can also be influenced by configuration of turbines, flow, amount of spill, condition of the fish, and

4 NaEurai Flow

ReLyumrd Flow ......

JAN FEB MAR ;kPR MAY J`UNE DE'C TULY AUG SE, T OCT NOV

Figure 1. Generalized effect of hydroelectric operations on mainstern Columbia flows ; River at The Dalles, Oregon. Taken from A Question of Balance', Pacific Northwest Regional Commission, 1978. physical structure of the facility.

Adult salmonids migrating upstream may be delayed by factors such as temperature, facility design, altered flow patterns, and nitrogen supersaturation. Salmon do not feed during their upstream migration and energy be reserves may depleted prior to the spawning period, thus reducing spawning success. Higher flows at the dams increase the search time and effort to locate small attraction flow at fishwav entrances. Work conducted by French and Wahle (1966) showed a 2-4 day delay for chinook and sockeve salmon at Rock

Island Dam in 1954-1956. Schoning (1956) 2.5 3.0 in and Johnson showed a - day delay migration of fall

Chinook salmon at Bonneville Darn in 1948. Gibson et aL (1979) estimated that Bonneville Dam delayed spring chinook an average of 2 days during 1978. These increases in energy demand can increase migration time, add more passage stress, and modify the productive capacity of the returning adults through higher rnay prespawning mortality which be age- and sex-specific.

While changes to the historic riverine environment have impaired environmental conditions for anadromous fish, they have also enhanced habitat for competitors and predators. Fish communities in the

Columbia River evolved over thousands of years to cope with the natural flow regimes of the basin. flow Changing these patterns affected the dynamics of the river ecosystem, In response to declines in population levels of anadromous salinonids during the mid-1970's the agencies

tribes in and greatly restricter, allowable harvest on depressed stocks, some cases, allowing only token fisheries orclosingfisheriesentirely, Evenwith-fairlyseverefisherymanagementactionsvariousstocksofsalmonstio exhibited declines in population levels and displayed signs of stock collapse. The reduced survival of these stocks as juveniles in the altered freshwater environment had simply been too great to overcome by curtailing harvest impacts alone.

By the late 1970's, upper Snake River salmon runs were being considered for potential Federal listing as - threatened and endangered species "A (see Question of Balance", PNRC, 1978). The agencies and tribes, who do not formulate the policies affecting the in management of water and power the basin, were - unsuccessful in incorporating fishery resource recommendations in power plans. In 1980, in part due to the concern over potential endangered species action, the possibility of the agencies and tribes making recommendations about the management of the hydrosystem was enhanced through the inclusion of the fish and wildlife provisions in the Pacific Northwest Electric Power Planning and Conservation Act of 1980 (Act) (16 U.S.C. 839 et seq.) Provisions in the Act supported the protection, mitigation. and enhancement of fish and wildlife influenced by hydroelectric operations and development in the basin. These provisions also allowed the agencies and tribes to bring forward issues for improving fish migration through the hydropower system.

In the early 1980's, the agencies and tribes developed instrearn flow recommendations to enhance fishin* survival (see "Initial Recommendations for Protection, Mitigation and Enhancement of Anadromous Fish the Columbia River Basin", Anonymous 1981). 'Me agencies and tribes flow recommendations were never implemented incorporated as or hard constraints in hydropower systern operations. 'Me first efforts to give fish greater consideration to needs in hydropower systern generation came in 1982 in the form of a compromise called the "Water Budget". The Water Budeet was designed by the Northwest Power Planning Council (Council) for the migration of juvenile salmonids as an alternative Lo the flow by regimes proposed the agencies and tribes. is The Water Budget a volume of water designated for use in providing enhanced river flows during the spring run-off period (April 15 through June 15). The Water Budget volume is 4.64 MAF 1.19 in of water. MAF the Snake River measured at Lower Granite Dam and 31.45 MAF in the Mid-Columbia River measured at Priest Rapids Dam. In-season recommendations are provided by the agencies and tribes to the power and water managers (U.S. Army Corps of Engineers and Bonneville Power Administration) to approximate the recommended flow regimes Cor spring migrating juvenile salmonids.

The Water Budget has not provided specified flow in levels as hard constraints hydropower system operations. Tle Council did define flow not what quantities or levelswere sufficient to improve production, migration and survival of anadromous fish. Rather than meeting specified flow levels at critical points in the river, the provision of the Water Budget volume has become the biological standard for determining whether an action is harmful fish to (see Bonneville Power Administration's "Proposed Non-Treaty Storage Environmental Agreement Assessment" March 1990). The fact is that the Water Budget inadequate to meet

6 even minimum fish flow needs during much of the time period when it can be used (April 15 through June 15) has been ignored.

A serious consequence of the Council failing to specify flow levels at critical points in the Columbia River is the inability of the agencies and tribes to use the Water Budget to improve flows below the confluence of the Columbia and Snake rivers. After smolts successfully migrate through the upper Columbia and Snake river reaches they must then migrate through the two largest reservoirs encountered during their passage to the sea. McNary and John Day reservoirs are 4 to 5 times as large as the upper river reservoirs, have much greater distances bank to bank, and lower water velocities through them. When velocities through the Snake

River reservoirs are decreased because of low flow conditions, the ability of the Water Budget to improve migration conditions through the lower Columbia River is reduced because the Council's plan limits flows to 140 thousand cubic feet per second (kcfs) at Priest Rapids Dam for fish passage purposes. eliminating the possibility of using the upper Columbia River flow to improve conditions in the lower Columbia River.

Ironically flows of 140 kcfs or higher are often provided at Priest Rapids Dam later in the year to meet power marketing needs.

Since the provision of the Water Budget "volume" is the only hard constraint on hydropower system operations, river operations to meet power needs have continued to expand and evolve without regard to the impact on fish needs. For example, when the Water Budget was established, the agencies and tribes understood that power flows would be sufficient to allow for safe migration after June 15 during the summer migration period. However, non-firra power created by the Water Budget is exchanged with California and other Northwest utilities for power returned during June and July. This exchange power allows the Columbia

River system to store water and reduce flows during the summer. 'Merefore, when the Water Budeet is used to increase spring flows, a corresponding decrease in summer flows occurs. Low summer flows in recent years have substantially increased the migration time for subvearling chinook and increased their exposure to predators and higher water temperatures.

The largest juvenile outmigration. in terms of absolute numbers. occurs during this summer period. and includes the depressed mid-Columbia summer chinook stock and Snake River fall chinook stocks, the latter currently being reviewed for listing under the Endangered Species Act. Columbia River upriver fall chinook, which support international ocean fisheries. in-river tribal subsistence and commercial fisheries. and non-tribal recreational and commercial fisheries. are also present during this period.

7 IL IDENTIFICATION OF THE NEED FOR FLOW A. Juvenile Salmonids

Hydroelectric development affects survival of juvenile salmonids in two ways: the first relates to the mortality incurred as a consequence of passage past the dams, and the second results from the physical alteration of the environment. Specifically, hydroelectric development changed a once free flowing river into a series of reservoirs and impoundments with greatly reduced water velocities. To counteract the effects of the physical dams structure, have been altered to improve the passage of both juveniles and adults. Other mitigation actions to date have included a program to transport fish around the hydroelectric system, via barge truck, and installation or the of structural modifications in spill bays to decrease the levels of nitrogen supersaturation during periods of spill. Despite these measures, spring and summer Chinook runs have not responded, and steelhead have responded only minimally. When the Columbia and Snake rivers were in their natural state. thev were free flowing rivers which followed the natural cycle the of melting snowpack. The spring freshet and the onset of salmon migratory behavior coincided. Free-flowing waters were cooler, and the higher turbi dity encountered benefitted migrants by providing concealment from predators (McRimmon 1954). The rate of migration of Chinook salmon sinoits has been found to increase with increased turbidity (Thomas 1975).

Hydroetectric'development changed the free-flowing rivers and created a series of reservoirs where water velocity decreased, resulting in an increase in the amount of time required for a smolt to travel from its point of origin to entry into salt water ("travel time"). Smolt passage through reservoirs slowed to one-half to one- third as fast as it through was free-floxing stretches of river (Raymond 1988). Bentley and Raymond (1976) showed that the travel time from the Salmon River to the Ice Harbor Dam site for vearlini, Chinook ranged from 9 to 15 days, before the installation of Lower Monumental and Ice Harbor dams. whereas the average travel time ranged from 20 31 days to after installation. In addition to duration. the migration timing of some stocks has been altered. However. in the two-month delay migration timing at Bonneville Dam for subyearling Chinook (Park 1969) may be in part.attributed to the late release of fish from hatcheries above the dam.

Sims and Ossiander (1981) demonstrated the relationship between river flow and travel time. This relationship has been verified and expanded upon with current monitoring data from the Smolt Monitoring Program (Smolt Monitoring Program 1987@ 1986; 1988; 1989). Sims and Ossiander postulated that if travel time affects the survival juveniles of there would be a positive relationship between river flow and survival. They found a positive correlation between estimated average smolt survival per project and flows at Ice Dam Harbor during the period of peak migration from 1973 to 1979. Raymond (1988) evaluated in trends the survival to adulthood of spring and summer Chinook and steelhead that had originated from Columbia the and Snake rivers. He concluded that the survival rates of adult fish, which return of returned from juveniles that had out-mierated between 1962-1984. had declined

8 as a result of the hydroelectric development of the river. Earlier studies had demonstrated a positive relationship between smoit survival and rate of return of adults (Raymond 1979). Together these studies confirmed that formation of the hydroelectric system negatively impacted the survival of juvenile salmon,

which resulted in reduced numbers of adults returning to the mid-Columbia and Snake rivers.

From the previously described research studies the role of river flow in determining juvenile survival, and

subsequently adult returns, was described. It was recognized that juvenile survival was a function of travel is time, and it travel time that is inversely related to river flow. Increased travel time and migration delav low due to flows reduce juvenile survival as a result of: 1) residualism in reservoirs; 2) a reduced ability of

juvenile migrants to tolerate saltwater; 3) a delayed entry into the estuary and ocean beyond the time period when conditions are most favorable for survival; 4) an increased exposure time of juvenile migrants to predators; and, 5) exposure to higher water temperatures. The following sections describe in more detail how the factors that determine juvenile survival are functions of time. 1. Migratory Behavior

a. Biology

onset of migratory behavior is closely associated with the smoltification process in juvenile salmonids.

Smoltification includes changes in both morphology and physiology, resulting in migratory behavior and the

ability to live in seawater (Bern 1978; Folmar and Dickhoff 1980).' Numerous morphological changes such

as the weight to length ratio, coloration, change in caudal peduncle shape, fin shape and coloration, and

development of recurve teeth in the mouth result in a smoit profoundly changed from the freshwater parr (Vanstone and Market 1968; Gorbman et aL 1980@ Winans and Nishicka 1987). Many physic)logical changes

are related to each of these general changes and collectively typify smoldrication (Folmar and Dickhoff 1980: Wedentever et aL 1980; Hoar 1988). Behavioral changes associated with smoitification include restlessness, elimination of territoriality, onset of schooling behavior, and becoming senti-petagic (Hoar 1965. McKeown 1984). the cumulative effect of the above changes is that smelts are no longer adapted to remain in freshwater habitats, but are well adapted for saltwater entry.

The migration of juvenile salmonids from their freshwater habitats to the ocean must be bv actively swimming, passively by transported the current, or both. In considering these modes Thorpe et aL (1981) "It stated would be energetically inefficient and ecologically imprudent for smolts to swim acLivetv downstream river when a could transport them passively over the same route. Pressure Lo evolve such active behavior would only arise if the passive transport system was too slow, or resulted in the delivery of smotts into the sea at an inappropriate season". Smith (1982) postulated that smelts actively swim upstream. but because of their reduced swimming performance are swept downstream. In fact, the only active migration of smoits that occurs routinely appears to be associated with sockeye migration through lakes (Johnson and Grocite t963; Groote 1965). Therefore, we will focus our consideration to the passive mode of migration and how it is related to flow.

9 A mostly passive mode migration, of taking advantage of downstream displacement by water currents,6 made possible by several mechanisms: development of negative rheotaxis (the movement of an organism in response a a to current); decrease in swimming proficiency; and, a decline in swimming stamina in smolts when compared to part (Folmar and Dickhoff 1980; McCormick and Saunders 1987). Annual rhythms in rheota.xis has observed in been Atlantic salmon (Salmo salar) with strong negative rheotaxis in smolting juveniles (Lundquist and Eriksson 1985). A reduction in swimming stamina among smolts compared to part has also been observed (Folmar and Dickhoff 1980). The swimming ability for coho salmon parr is 3.5-7.3 body lengths (BI-s-1) per second and for coho salmon smolts about 2-5.5 BLs-1 (Glova and McInerney 1977; Smith 1982). A similar decline for BU-1 Atlantic salmon from up to 7 for part to about 2.0-2.5 BLs-1 for smolts indicates that this is not unique to coho salmon, but may be common among all salmonid smolts (McCleave and Stred 1975; 1978). 'norpe and Morgan These chances in swimming behavior provide a mechanism for downstream displacement by flow and a resulting migration that is largely passive. depending mostly on water flow to determine migration speed and direction. Early observations on chinook salmon support the hypothesis of a mostly passive migration. In a studv conducted on Sacramento the River from 1896 to 1901, Rutter (1904) stated "there is no doubt that in migrating the fry drift downstream tail first, keeping the head upstream for ease in breathing as well as fC* convenience in catching food floating in the water" (his reference to fry is somewhat misleading in that the Fish were about 5 cm in length). The hypothesis of passive migration is also supported by numerous observations on Atlantic salmon. Studies on Atlantic salmon by Thorpe and Morgan (1978), Tyder et al. (1978), and Thorpe aL (1981) in et Scottish rivers, lochs and estuaries and by Fried et aL (1978) in the

Penobscot River is estuary suggest the Migratory behavior mostly passive. In each study juveniles drifted with the night 6 9 current at for to hours. Although random movements occurred for various lengths of time during the night, the overall displacement was downstream at a speed consistent with the current velocity. A passive mode of migration is by further supported the close relation between observed migration rates chinook of and steelhead and the expected rates based on the water particle travel time (Figure 2). In this particle in case the water travel time days was estimated by dividing the total reservoir volume in a reach by daily flow an average at the downstream end of the reach (known as the "storage replacement method" developed by U.S. the Army Corps of Engineers). Of particular interest in this relation between travel time of smolts between Lower Granite Dam and John Day Dam and Snake River flows is that travel times of W to 15 days appear possible over a wide range of relatively high flows. b. Control of Migratory Behavior

Migratory behavior is by controlled genetic and environmental factors (Randall et aL 1987). Genetic* selection favors genoty pes controlling behavior patterns that improve the survival of their carriers (Smith 1985). As early as the 1920's the migration juvenile I patterns of chinook salmon were considered to be inherited by subsequent generations in the Columbia River (Rich and Holmes 1929). In a review, Randall

10 Z'

SDring Chinook 35 Steelhead Nviter Particle

30 V) >1

25 E

20

15 0

10

5

40 60 so 100 120 140 160 180 Flow (kcfs) -

Figure 2. Travel time of juvenile Spring chinook and steelhead from Lower Granite Dam to John Day Dam and the calculated water particle travel time as a function of.flows at Ice Harbor Dam. et aL (1987) pointed out that the genetic influences on the age of smolting within species have been underestimated in the past. Recent findings indicate chinook in the Nanaimo River, . which are characterized by a specific age and size at seaward migration, can be associated with significantly different frequency of allozymes and are seemingly a genetically distinct sub-population (Carl and Healey 1984). At the turn of the century apparently a wide variety of migratory traits e)dsted, as Rich (1922) observed juvenile chinook salmon in the Columbia River estuary throughout the year. Current knowledge suggests that wide migration variety of patterns among hatchery and wild stocks has a genetic basis, and efforts to protect these are desirable.

Environmental cues serve to synchronize the initiation of miaratcry behavior and the more general endogenous rhythmicity associated with smoltification. The mechanisms controlling smoltification are mediated through endocrine control at two levels: the first interfaces environmental inputs with the pituitary

involves in and the second the function of hormones the endocrine system (Groote 198L Schreck 198L 1986). Barron Important environmental factors involved with the development of a disposition to migrate photoperiod, are water temperature and stream discharge. When fish are in a proper state of migratory

11 readiness, a proximal stimulus, such as lunar phase or stream flooding initiates migration (Hoar 1988). Photoperiod is a key environmental cue influencing the timing of downstream migration in juvenile steelhead (Wagner 1974). role Ile of photoperiod cues apparently result from the direction and rate of change of day length (Wedemeyer aL et 1980). Baggerman (1960) and Wagner (1974) emphasiz e that, while photoperiod-controlled changes may bring the animal into a state of preparedness, priming it for migration, other released stimuli initiate and maintain migration. Consequently McKeown (1984) concluded there is relatively little evidence in support of photoperiod being an important cue in the actual initiation of migration.

Temperature influences smoltification by contro Iling the rate of the physiological response to photoperiod, such that effects are apparent sooner at elevated temperatures (Wedemever et aL 1980; Hoar 1988). The migratory movements of Atlantic salmon smolts are closely correlated with water temperature with only small numbers moving below a threshold temperature (Solomon 1978). Similariv, water

explained 89 95 , temperature to percent of the yearly variation in the date of cumulative smolt migration. by Atlantic salmon through a combination of temperature increase and ambient river temperature during spring (Jonsson and Ruud-Hansen 1985). Average stream temperature explained 60 percent of the variation in the median emigration date of of coho salmon smolts from Carnation Creek, British Columbia (Holtby all streameet 1989). In contrast to these findings, Bjornn (1971) could not establish a causal relationship between

temperature and the seaward migration of salmon smolts. Although the smolt migrations coincided with increasing in stream temperatures the spring, the increasing temperatures seemed coincidental since steelhead reared in a spring-fed pond migrated from the relatively constant temperature pond at the usual time. 'Mains and Smith (1964) found that seaward migration of chinook salmon in the Snake River during 1954 and 1955 was predominantly in the spring. This migration coincided with the spring runoff. They stated that, "While temperatures may play an important role in initiating the downstream migration of chinook salmon. the First occurrence of the spring freshet was the primary factor responsible for stimulating this phenomenon. In both during years which this study was made, the discharge required in the Snake River before migration commenced was approximately 70,000 cfs". 2. Physiological Development

Physiological changes in juvenile salmon encourage migration and prepare them for residence in seawater. The behavioral motivation for migration has long been recognized as having an endocrinological basis (Hoar 1958). The thyroid hormones have been implicated in behavioral changes associated with migration. but the relations have not been completely elucidated (Leatheriand 1982: Eales 1985: Dickhoff and Sullivan 1987@ Grau 1988). Godin et aL (1974) injected juvenile Atlantic salmon with thyroid hormones and observed that swimming activity, aggressive behavior, and upstream orientation were significantly reduced. They concluded that the hormones initiated the migratory tendencies. Similarly, others have concluded that increased plasma thyroxine permits smolting Atlantic salmon to resist displacement in high flows and orientate head-

12 downstream in flows, moderate thereby elevating ground speed at no extra metabolic cost (Youngson et aL 1985; Thorpe 1989). The thyroid hormones do have an endocrine role in controlling migration behavior, but

s (1988) a Hoar concluded, they do not regulate behavior per se.

The migratory behavior of smolts has also been related to the physiological changes associated with the

development of osmoregulatory capacity, particularly the level of gill sodium potassium adenosine tri- (gill activity. phosphotase AT?ase) (Zau gg and Wagner 1973; Wagner 1974; Zaugg et aL 1985; Rodgers ei

aL 1987). The coincidence of an increased percentage of juvenil e steelhead migrating from experimental

releases and the seasonal rise in gill ATPase has been demonstrated for winter steelhead from the Alsea

River, and for summer steelhead at Dworshak National Fish Hatchery Idaho (Wagner 1974; Zaugg 1981a;

Zaugg 1981b). The same general relationship has been observed in yearling spring chinook salmon from the Deschutes River, Oregon that were allowed to migrate in an artificial stream (Hart 1981). Many of the observed relations between migratory behavior and aill ATPase activitv in smolts are derived from juvenile salmon held in the captive environments of the laboratory or hatchery. We know that smoits released to migrate freely usually exhibit remarkable smolt development indicated by rapid increase in activity aL ATPase (Ewing et 19K Zaugg 1981; Zaugg 1981b: Zaugg et aL 1985). In contrast, the smolt gill development, including ATPase and plasma thyroxine responses, of fish held in the captive environment is aL often suppressed (Zaugg et 1985; Nishioka et aL 1985; Patino et aL 1986; Rodgers et aL 1987; Maule

et aL 1988).

The duration of the elevated gill ATPase levels among migrants is of interest because a decline may indicate a reversion to a part status accompanied by a loss of migratory behavior. Zaugg (1981b) found that

yearling coho held at hatcheries beyond normal May releases showed a decline in ATPase levels and a reversion to the parr appearance. Despite this reversion, fish released in June and July rapidly migrated 2@17Pase seaward and experienced renewed high levels. Although it is apparent that at least coho can regenerate high ATPase levels, it is not known how long high levels are normally sustained in migrants. Although the migration experience is stirmulatory ATPase activity collected from miQrating smolts of hatchery

and wild origin suggest an early June decline similar to the seasonal rhythmicity observed in captive environments (Figure 3).

a. Delay and Residualism

Migratorv behavior of smolts is seasonal and fish that do not complete the migration become holdovers or residuals. A holdover juvenile salmonid over-winters in the reservoirs. sometimes completing the migration during the next spring. A residual undergoes a reversion to the freshwater form and becomes a resident in the reservoirs, not completing the migration to the ocean. The problem of steelhead holdover and residuals in has been most apparent the Snake River during exceptionally low flow years such as 1973 and 1977. During 1977 only about 50 percent of the steelhead smolts starting their migration in tributaries reached

Lower Granite Dam during the spring migration. That most of the fish arriving late were lost to mortaliLv

13 40

30

20

>1

>

U 10 12 27 12 Apr May Jun Ca

a

30

20

10

1'2 27 12 Apr May Jun

Date

Figure 3. L) MeangillATPaseactivitv(pmolesPiomgProt-loh of (A) hatcherv and (B) wild steelbead during the spring migration. 1989.

14 is evident, because only 11.000 fish were collected at Lower Granite Dam. These fish were collected in

October and November and transported from the Snake River (Park et aL 1978). Steelhead holdover or

residualism also occurred in 1987, a low flow year. During a creel census of the adult steelhead harvest in

Lower Granite and Little Goose reservoirs between November, 1987 and January, 1988, a total of 213 juvenile steelhead bearing marks associated with coded wire tags were incidentally collected. Among the juvenile steelhead identified from tags, 51 percent originated in Oregon and 43 percent in Idaho hatcheries (Mark Schuck, Washington Department of Wildlife, personal communication). Purse seine collections in the summer of 1988 in Lower Granite reservoir revealed the presence of residual steelhead. Steelhead

contribution to the collection was about 25 percent of the level observed during the spring quarter, based on catch per effort. Subsequent low catches of steelhead in the fall purse seine collections indicated that

either over-summer survival was low or they emigrated out of Lower Granite reservoir (Bennett and

Chandler 1990). From these recent observations. there is continuing evidence that steelhead migrating as far as Snake River reservoirs are remaining in freshwater well beyond the accepted norm. The likely consequence is low survival due to disease, predation, and fishing. b. Salinity Preference and Tolerance

The development of osmoregutatory capabilities is concurrent with a change in behavior that results in a strong salinity preference (Baggerman 1960; Otto and McInerney 1970). Salinity preference has been

proposed as an orientation mechanism for migration, particularly in the estuary (McInerney 1964). The salinity preference is a behavioral attribute of smoits that is restricted to a limited time (Baggerman 1960;

McInerney 1964). Experimental results show a preference for salinity at the time of migration and a reversion to freshwater preference if the migrants continue their freshwater residency.

The migratory disposition in juvenile steelhead and coho salmon has been found to be preceded by the development of salinity tolerance from as much as several weeks to six months (Conte and Wagner 1965@

Conte et aL 1966). The development of some salinity tolerance among juvenile sainionids in a wide range of sizes and physiological conditions independent of migratory behavior is not surprising, but the high saliniEv tolerance and subsequent rapid seawater growth without stunting is an attribute of smolts (Kepshire and McNeil 1972; Woo et aL 1978: McCormick and Saunders 1987). Continued freshwater residency is associated with reversion to a parr-like fish with a lower salinity tolerance in steelhead. coho. and chinook salmon

(Conte and Wagner 1965; Wagner 1974; Woo et aL 1978). Chrisp and Biorn (1978) concluded that hatcherv and wild steelhead could not tolerate saltwater at a concentration equal to 10 parts per thousand in a 10 day by challenge, the time the migration from the upriver areas terminated in early June. Similarly Adams et aL (1975) concluded that saltwater survival of steelhead transferred directly to saltwater at 10 to I L')'C was low in eariv March, near 100 percent in mid-April. and declining by early May. Fall chinook differ from other salmonids since their seawater adaptability increases in early May and remains high well into July (Clark and Blackburn 1978; Clarke and Shelbourn 1.982). In early August, the latter part of the subvearling chinook

15 migration at McNary Dam exhibited a reduced osmoregulatory a bility (Maule et aL 1988; Schreck et aL 1984) Similarly, fall chinook from Spring Creek National Fish Hatcherv. on the lower Columbia river, exhibited sharp decline in ability to withstand direct exposure to sea water in the laboratory (Gould et aL 1985). "Windows' c. of Biological Timeliness and Their Management Implications

Numerous investigators have concluded that " salmonid smelts have biological windows" to successfully enter the estuary and ocean (Walters aL 1978; Bilton aL 1982; et et Boeuf and Harache 1982: Holtby et aL. 1989). The concept of temporal-spatial windows has been developed primarily to explain the timing of smolts relative to coastal predators, marine productivity and oceanographic conditions that are likely determinants of early marine survival. However, the smolt migration can also be considered to have windows limited by photoperiod, temperature, and factors other controlling the behavior and the physiology of smolts. The duration of such windows is delineated by the onset and decline of migratory behavior, seawater preference, seawater tolerance, and selected physiological attributes such as gill ATPase. Flow not only determines the time required by smolts to reach but it the ocean, ensures that smolts are at the appropriate location and time and in an acceptable condition.

is Since smolt migration mostly passive their migration is rate controlled primarily by river water velocity. The effect of mainstem Columbia River is reservoirs to increase the cross section area of a river, and therefore reduce its water velocity compared to the free-flowing reaches. Raymond (1968; 1969) made

several comparisons of yearling chinook h salmon travel throu 9 McNarv reservoir before and after its impoundment and concluded that fish moved only about one-third to one-half as fast. Raymond (1968) supported his conclusion based on the relation between passive migration and river flow and the observations of Nelson et aL (1966), who found that aver@ge water velocities through McNarv reservoir were between 30 and 42 percent of the average in water velocity the unimpounded Columbia River below McNarv Dam. There is also evidence that juvenile sockeve salmon experience a similar delay. The median dates of passage of juvenile sockeye salmon at Bonneville Dam during 1987-89, were May 20 to 24 (Fish Passage Managers 1989). During 1946-53 the median passage dates between I were April 2J and May 1 (Davidson 1965), about four one to weeks earlier than under conditions. Slow is recent migration through reservoirs not unique to Pacific salmonids. Atlantic salmon in smelts released lakes were considerably delaved in their downstream migration compared to river released smolts (Hansen et aL 1984). Therefore, even if seasonal flow regimes remained unchanged from historical flow patterns, the reduced water veiocitv in reservoirs would result in slower migration rates for salmonid smotts.

in Delay migration can expose smolts to seasonally rising water temperatures. Since gill ATPase activitv and migratory disposition are sensitive elevated to temperatures. exposure Lo such temperatures during migration may have deleterious effects. The temperature effects on steelhead are of particular concern because steelhead migrate later than yearling chinook and are more temperature sensitive than coho salmon. Based on laboratory experiments, water temperatures of 15'C (59F) caused a steep decline (Figure 4) in the

16 Na+, K+- stimulated adenosine triposphatase activity (micromoles of inorganic phosphorus per milligram of protein per hour)

eb

OZ

C3,

0 rj + tz q 7-0. U1 + > rt cm Cr 0

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Z gill ATPase activity of yearling steelhead, the authors suggested an upper limit of 12'C (54'F) (ZaUgg t 1972; Adams et aL i975; Zaugg 1981a). The effect of temperature on the elevated gill ATPase activity o

migrants that have migrated for weeks rising is under seasonally water temperatures of concern. This condition is fundamentally different from increases in gill ATPase activity observed among juvenile steelhead a few days after hatchery release, when they may experience rising water temperatures in the natural environment. Furthermore, temperature limits are further supported by the reduced number of smelts that were observed from seawater rearing of steelhead in net-pens when temperatures rose above 12'C (Prentice et aL 1981). As a consequence. temperature limits were incorporated into the "Initial Recommendations for the Protection, Mitigation, and Enhancement of Anadromous Fish in the Columbia River Basin" by the agencies and tribes (Anonymous 1981). They reasoned that the smolt migration usually begins in the Snake River in mid-April and there is only about 30 days smoits to move through the mainstern rivers before water rise above 12'C in the temperatures lower Columbia River. Temperatures of 15'C were reached at Bonneville Dam in late May to early June during most recent years. Therefore. median arrival dates of June 9, 1973 and June 21, 1977 at The Dalles Dam (Sims and Ossiander 1981) are unacceptable with respect to knowledge gill current of seasonal ATPase patterns, water temperatures. and expected migratory behavior. 3. Predation Losses and Flow Fish predators are important sources of mortality to outmigratin@Z juvenile salmonids in the Columbia River (Rieman aL 1988; et Uremovich 1980). Northern squawfish (Ptychocheilits oregonensis), walleye (Stizostedion vitreum), smallmouth bass (Micropterus dolomieui), and channel catfish (1cralitruspunctatus) are the major fish predators in John Day reservoir, with northern squawt7ish accounting for 78 percent of the estimated smolts lost piscivores to (Rieman et aL 1988). Northern squawfish predation upon juvenile salmonids is influenced by many factors, including prey density, prey species. predator size. arid time of the year (Poe et aL 1988; Vigg 1988). et A Changes in temperature and smolt residence times caused by hydroelectric development and altered now regimes are especially important factors with respect to predation. insufficient water.flows, causing extended residence times of smelts during high temperature periods. may severely exacerbate predation the problem. Also, water velocity will affect the distribution of both prey and predators. The following sections describe how temperature and residence times influence predation losses. especially from squawfish. Smolt northern residence time within a rese[voir, associated temperature effects of extended or shortened residence times, and water velocity are discussed below as potential mechanisms of how flow manipulation could alter predation losses.

a. Residence Time Effects on Predation Losses Travel time is function a of Row and, therefore, residence time is also a function of flow. Smolt losses caused by northern squawfish predation varies with smolt residence time. and therefore with flow. Modeling exercises of northern squawfish predation in John Day reservoir have estimated the effects of residence time, flow and other variables. Beamesderfer aL (1991) et used a system of differential equations solved at dai[v

18 intervals to predict smolt mortality during passage. In their sensitivity studies, flows alone had a relatively

small effect upon smolt mortality; however, mortality increased rapidly as increased. Bledsoe I I temperature P (1989) used the Columbia River Ecosystem Model (CREM), composed of ordinary differential equations,

O to simulate predation in John Day reservoir. CREM simulations of subyearling chinook salmon showed t at increasing residence times from 7 to 18.5 days caused mortality to increase from 38 percent, to about 65 percent. Residence times as long as 3 months caused mortality to be about 85 percent. Neither of these els have been used to examine passage through a series of reservoirs.

The impact of low flows upon predation losses are likely the result of both long residence time in a

reservoir (long exposure to predators) and higher predation rates caused by the increased temperature during the summer months. Temperature increases about VC every 9 days between May and August. and northern squawfish predation has been shown to increase rapidly with temperature (Beyer et aL 1988: Vigg 1988). Beamesderfer et aL (1991) estimated that'150.000 smalts were lost in John Day reservoir for each I'C rise in temperature. Increases in mortality with longer residence times observed with the CREM model are also linked to temperature changes,. since temperature continues to rise throughout the simulation. b. Temperature Effects on Predation Losses

is Temperature probably the most important physical variable affecting the consumption rate and growth of predatory fishes (Brett 1979; Kitchell 1983). Consumption rate increases from low values at low temperatures to a maximum rate at an optimum temperature. Above the optimum temperature, consumption rate usually declines rapidly, eventually failing to zero near the maximum lethal temperature for the species. Consumption rate of northern squawfish, as a function of temperature, has been examined in the field and in the laboratory. Average consumption rate was significantly affected by temperature, prev density and predator weight in analyses of John Dav reservoir data (unpublished data.'S.Vigg, U.S. Fish and Wildlife

Service). Analyses showed that consumption increased rapidly with increasing temperature. Laboratory studies on digestion rates of northern squawfish showed faster digestion and prey evacuation at high temperatures (Falter 1969: Steigenberger and Larkin 1974; Bever et al. 1988). Laboratory experiments (Vigg and Burley, MS in review) demonstrated that maximum consumption of salmonid prey increased from 0.5 smolts/dav at 47'F to 7 smolts/dav at 71'F (Table 1; Vigg and Burley MS in review).

The most rapid increase in maximum consumption occurred between about 60 and 70'F (Table 1).

Summer temperatures in Columbia River reservoirs commoniv reach 70'F. or hiaher. and mav remain this high for 2 or 3 months (U.S. Army Corps of Engineers. Annual Fish Passage Reports).

c. Water Velocity

in Increased water velocity the tailrace of a dam, caused by increasing the water flow through the turbines spillway, or over the changes the behavior of predators. This in turn may alter the predation rate in the area. Faler et al. (1988) monitored the movements of northern squawfish by radiotelemetry ire the McNary Dam tailrace, and observed that northern squawfish avoided areas with high water velocity. When dam discharge

19 rates were high during spring or carly summer, northern squawfish remained in protected near shore arcas.1p Northern squawfish moved closer to the dam and the juvenile bypass outflow area when discharge rate was low in mid to late Northern in summer. squawfish did not usualtv reside areas with water velocities of 100 cm/sec greater. Therefore, if flow or rates were increased during the summer, predation on juvenile salmonids in dam tailraces might be reduced by forcing predators away from bypass outlets, where smotts - appear to be especially susceptible aL to predators (Vigg et 1988). In addition, higher water velocities would likely reduce the exposure time of smelts to northern squawfish in the if tailrace zone, even velocities were - not high enough to displace the predators. 4. Conclusions

All of the aspects of smolt survival discussed thus far share a common foundation - the magnitude of their impact is function time. As a of the time spent migrating increases, the "biological window" of opportunity for successful migration is lost. Longer residence times increase mortality due to predation and disease. Later migrations occur under more adverse temperature conditions, when increased predation rates and reduced salinity tolerance 'Me solution these "time is occur. to related" mortalities to reduce the amount of time that juveniles spend migrating through the system. Past studies have shown that adult production will

be increased by increasing smolt survival. and smolt survival be by can increased decreasing the time spent migrating.

Predation alone, whether it is management predator removal, protecting prey or a combination, will not restore smolt survival to pre-impoundment levels. Physical injury and the stress of dam passage, the onset and decline migratory behavior, of seawater preference and seawater tolerance would still be some of the important determinants of how many smolts migrate, residualize, or survive entry to the estuary. Therefore. increases in flow essential improving are to smolt survival and rebuilding the runs. B. Adult Salmonids 1. Table Maximum consumption rates f@lowingwatercanproducevelocitiesandmodifv (smolts/predator/dav) of northern squawfish upon salmonid prev at varions the physical and chemical characteristics of river temperatures. Data are from Vigg and Burley (N-IS in reaches in ways that influence the upstream preparation). migration and spawning success of adult Ma:.. Consumption Rate ariadromous salmonids. Favorable migration 47 0.5 54 1.2 conditions minimize delavs and overexertion that 1 63 4.2 may prematurely exhaust limited energy reserves. 71 7.0 Adult salmoruds are attracted to fast-flowing

water, and moderate velocities may enhance

migratory performance relative low velocities. Velocities in to Columbia and Snake river mainstem reservoirs appear inadequate to orient and stimulate adult migrants, although other mechanisms apparently permit successful transit impoundments. of Numerous examples of spawning migrations that correspond with

20 seasonal peak flows suggest that high flows favor upstream migration. However, extremely high flows can

impede passage at dams, since adult fishways were not designed to adequately pass both high flows and adult salmonids.

1. Migratory Behavior

a. Migration Timing

The timing of many runs of adult anadrornous salmonids corresponds with peak flow (Collins 1892;

Pritchard 1936; Cramer and Hammack 1952; Andrew and Geen 1960; reviews in Major and Mighell 1966,

and Banks 1969; CDE and IPSFC 1971; Baker 1978). There is evidence that prior to decimation of the summer run the mode of the Columbia River chinook salmon migration corresponded with the time of highest river discharge (Figure 5). The former runs of sockeye salmon in the Columbia River were initiated with the rising waters of spring (Collins 1892), and the present run of that species still corresponds closely with the time of historical high flows (Figure 5). Season, climatic conditions. and characteristics of the

discharged water are usually correlated with flow, so it is not clear whether flow itself was or is the primary

determinant of run timing (Banks 1969). The association between migration timing and flows, as well as the

behavior of migrating adults with respect to velocity (discussed in a later section), suggests that high flows may provide a favorable migrating environment for adult salmonids. b. Bioenergetics of the Spawning Migration

Delays caused by an unfavorable migrating environment may contribute to reproductive failure. Adults, which fast during what may be a months-long upstream journey, exhaust virtually all energy reserves prior

to spawning and death (Idler and Clemens 1959; Gilhousen 1980). Delays of as little as 3 to 4 days at

migration barriers have been associated with premature mortalities (Godfrey et aL 1954-. Andrew and Geen

1960: CDE and IPSFC 1971), and unusual exertion (DeLacy et aL 1956: Paulik 1960).

Extreme flows, both high and low, and high water temperatures are understood to cause delays in the spawning migrations of some salmonid stocks in the Columbia, Snake, and other rivers (Thompson 1945: Fish

and Hanavan 1948; Cramer and Hammack 1952; Major and Mighell 1966: ODFW 1977; Johnson et aL 1982;

Liscom et aL 1985; Shew et aL 1985). High temperatures, in addition to blocking migration, can increase the rate at which limited energy is consumed for standard metabolism (Fry 1971). Females suffer more from some delays (Godfrey et aL 1954), perhaps because they have less surplus energy than males (Gilhousen

1980). There are also differences among runs, and between early and late components of runs, with respect to energy reserves and swimming ability (Gaulev 1960; Gauley and Thompson 1963@ Gilhousen 1980).

c. Spawning and Incubation

Some important anadromous stocks are mainstern spawners with special flow needs for spawning. The Hanford Reach of the mid-Coltimbia River hosts spawning populations of summer steelhead and upriver fall chinook (Becker 1985). Typically, about 70 percent of the adult spawning for Columbia upper river fall chinook occurs within the Hanford Reach (Carlson and Dell 1990). Spawning also occurs in mainstem areas

21 400 - 1918-25. 1927-31 350 -

300 1983-1986 E - ...... 250 -

200 -

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50 -

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Chinook Harvest 1876

Sockeye 4 60 Counts, 10-year Average ...... >1 Chinook Counts, 3 40 10-year Average ......

5 2 I - Z 20

0 0 Feb Jan Mar Apr fkfav Jun Jul Aug Sep Oct Nov Dec Month

Figure 5. Streamflow and timing of adult chinook and sockeve salmon runs: (A) Annual bNYdrograph of the Columbia River below the present site of Priest Rapids Dam (RM 396) in vears prior to project development (1918-1932. except 1925 and 1926), and in recent vears (1983-1986). Data from U.S. Geological Survey Portland. Oregon; (B) Chinook salmon harvest timing at Asto Oregon (RM 13) in 1876 day-' (fish - boat" for Astoria gillnetters. adapted from Thomp 1951, Figure 6), 10-year average chinook daily passage at Bonneville Dam, 1981-1990 (unpublished data, Fish Passage Center, Portland, Oregon), and 10-year average sockeve daily passage Bonneville Dam at for 1980-1989 (unpublished data. P.Lumley, Columbia River Inter- Tribal Fish Commission, Portland, Oregon).

22 above the Hanford Reach (Watson 1970, Rogers et aL 1988) and is suspected to occur in lower river reservoirs (Liscorn and Stuehrenberg 1983). Strong cohorts of upriver fall chinook have made large

contributions to ocean and in-river fisheries in recent years (PSC 1988: ODFW and WDF 1989). The Hells

Canyon reach is the principal natural spawning area of Snake River fall chinook (Howell et aL 1985). This stock is presently being considered for listing as threatened or endangered under the Endangered Species Act.

Water velocities are an important factor in redd site selection and construction (Chambers 1956, 1960;

Meekin 1967; McCart 1969), and seasonally high flows can play an important role in flushing harmful fine

material from spawning gravel (Reiser et aL 1985). Without high seasonal peak flow, which historically

occurred during spring and summer months when fry had already emerged from the substrate (Figure 5),

there has been little mass bedload movement in the Hanford Reach (Chapman et aL 1986). Gravel can become sedimented except where spawning is concentrated each year (Roy Beatv, Columbia River Inter-

Tribal Fish Commission, personal communication). The tendency of spawners to concentrate in high-use

spawning areas in the Hanford Reach (Dauble and Watson 1990) may reflect the high relative suitability of

gravel that has been cleansed of fines by redd construction in prior years. High flows during spawning can provide a greater wetted area for spawning when space is limiting, but

of equal or greater importance is the maintenance, until fry have emerged, of flow levels close to those that

prevailed during spawning (Thompson 1974; Graham et aL 1980; Chapman et aL 1986). At Vernita Bar, a heavily-used spawning area at the upstream end of the Hanford Reach and just below Priest Rapids Dam,

daily vertical fluctuations of 2 to 3 meters commonly resulted from power peaking operations (Chapman et

aL 1986) (Figure 6). Dewatering of redds during special low flow operations has been known to cause large

eg,g/fry mortalities (Bauersfeld 1978), although an agreement has been reached to regulate flow levels during some periods of spawning and incubationiernergence (FERC 1988). Redd dewatering and losses of fall

Chinook are also believed to be occurring in the Snake River, even when Idaho Power Company operates the Hells Canyon complex within Federal Energy Regulatory Commission (formerly Federal Power 4 Commission) license stipulations. Such abiotic. density independent mortalities during incubation may determine vear class strength in some salmonid stocks (McNeil 1969). Greater regulation of flow, particularly

is in the Snake River, required to safeguard important stocks of salmonids during a very vulnerable period of their life cvc1e.

Spawning and incubation temperatures have changed with development of these rivers. and it is very likely that high water temperatures during spawning and early incubation are affecting the productivity and population composition of at least the Columbia River upriver fall chinook (13caty, in prep.) and probably

Snake River fall chinook, as well.

2. Flow Effects on Passage

a. Reservoir Passage

23 0 180 -

1987 160 - 140 - 1926

120

U 100

80

60

40 ...... 20

0

Nov Dec . i" A pr Month

mid-Columbia* Figure 6. Hydrographs daily of average flow at Vernita. Bar in the Hanford Reach of the River (RM 392) during spawning, incubation and emergence of upriver fall Chinook salmon for a typical pre-development water year (1926) and a post-development water year (19S7), (FERC 1988). 1

Reservoir velocities may be inadequate evoke a to strong upstream orientation and movement of adult salmonids. Radio tracking studies have shown fish that movement in the relatively slow waters of reservoirs mav be disoriented, at least lacking or a strong "upstream" orientation (Trefethen and Sutherland 1968: Liscorn et aL 1978). Numerous radio tracking studies have documented well a phenomenon at Bonneville Dam, where radio-tagged adults exiting the Bradford Island fishway orient along the shoreline (or an isobath), follow it around Bradford Island, and pass before the spillway bays. where they frequently fall back when the spillway is operating (Monan and Liscorn 1973@ Liscom aL et 1977; Ross 1983: Turner et aL 1984b). That the tendencv to follow the Bradford Island shoreline downstream was less pronounced w hen higher velocities prevailed at the tip of Bradford Island (Liscom aL 1978), et suggests that forebay and reservoir velocities may be adequate not to evoke a rheotactic response under low to moderate flows, In the absence of sufficient velocities, alternative means of navigating through reservoirs may be available. Shoreline or isobath orientation, which may occur even in flowing dams (%I I waters below onari and Liscom 1973; Liscom 1976@ 1983)* and Monan Monan and Liscorn 1976; aL 1982@ Johnson et Ross 1981: Turner et aL is one alternative, as are celestial and in geomagnetic orientation (reviews Hasler 1971, and McKeown 1984). Such mechanisms may answer the riddle of successful spawning migrations through still-or-slow-water lakes

24 and reservoirs (Andrew and Geen 1960; review in Trefethen and Sutherland 1968). Adults bearing radio tags have migrated through Columbia and Snake River reservoirs at average rates of 0.8-2.0 mi h-' (1.3-3.2 km h-1) 1976@ Liscom (Monan and Liscom 1975, et aL 1978; Turner et aL 1983, 1984a). Reducing nighttime and weekend flows to project minimums significantly reduced the probability that steelhead passing Lower

Monumental Dam would reach Little Goose Dam within 150 hours (Liscom et aL 1985), which indicated that flow pattern, apart from daily average flow, can influence migration time.

In summary, velocities through reservoirs appear to be generally inadequate to direct and stimulate adult upstream migration, although migrants may employ methods other than rheotaxis to complete their journey.

is There evidence that higher velocities (Liscom et aL 1978), and more uniform flow patterns (Liscom et A 1985) can facilitate adult passage. b. Dam Passage

The structure and operation of dams determine in large part how various flow levels affect adult fish passage. Passage at the darns themselves is probably expedited at low flow levels: turbulence and extreme velocities in the tailrace are reduced; fishwav attraction flows are a higher percentage of total flow and are easier to locate; and minimal spill reduces fallback and nitrogen supersaturation. However, as will be discussed in subsequent sections; extreme low flows, independent of water quality conditions, adversely affect adult passage at dams.-

High flows, and how the projects handle them, can also have major negative impacts on adult passage. Project operations can affect approach to, and use of the fish passage facilities (Monan and Liscom 1971;

Liscom and Monan 1976; Moran and Liscom 1976; Turner et aL 1983, 1984a). Adult passage facilities may not be designed to perform adequately during periods of extremely high'flows or under certain operations

(Turner et aL 1983, 1984a).

Spillway operation contributes to dam fallback and nitrogen supersaturation. two problems frequently associated with unusually high flows. Many researchers have estimated fallback rates ranging up to 58 percent, as was measured for radio-tav 9 ed summer chinook salmon using the Bradford Island fishway at

Bonneville Dam in 1974 (Monan and Liscom 1975). Although fallback occurs via routes other than the spillways, spill level is often a major factor in fallback (Monan and Liscom 1975, 1976; Uscom and Monan

1976). However, the physical siting of fishway exits with respect to shore lines, forebay currents, and spillways, particularly for the Bradford Island Cishway at Bonneville Dam. may be the single most important factor contributing to fallback (Monan and Liscom 1973, 1975; Liscom et aL 1977; ODFW 1977@ Gibson er aL 1979; Ross i983). Most adults that fall back over spillways are able to continue their migration (Monan and Liscom 1975; Liscom et aL 1977), but it is not known how the delays and additional exertion to reascend the dam (sometimes repeatedly) ultimately reduce their ability to reproduce. In addition, Gibson et aL (1979) measured significant mortality in passage at Bonneville, The Dalles and John Day dams. It is reasonable to assume that fish which fall back have at least the same mortality rate on each attempt at passage. Therefore,

25 fallback increases mortality at dams. SPRI Nitrogen supersaturation, which can result from water spilling over dams and plunging deeply into basins, is a greater potential problem with high flows (Ebel 1969). The lethal and sublethal effects of nitrogen gas supersaturation on adult salmonids are well-known (Beiningen and Ebel 1970; review in Ebel aL 1975; aL et see also Colt et 1986), and may be exacerbated by high temperatures and prolonged exposure by delays caused (Beiningen and Ebel 1970). Spillway deflectors can reduce the hydraulic conditions that contribute nitrogen to supersaturation (Johnsen and Dawley 1974), but deflectors have not yet been installed on all dams. Dam passage problems associated with high extremely flows may not be the result of the flows themselves,

but rather the failure of the dam designers and operators to pass those flows safely. Regardless, it is dam passage problems that contribute to the relationship sometimes observed between extremely high flows and poor survival of adult salmonids through mainstem reaches containing multiple dams and reservoirs. c. Reach Passage Mortality 20 rates of percent for adult spring and summer chinook at John Day Dam have been reported as independent of flow (Gibson et aL 1979). Examining John Day and Little Goose dams. Mauseth & Associates, Inc. (1980) found significant no correlation between lagged daily conversion rates (from the next wiO down stream project) and flows. They did find significant but inconsistent (between species) correlations flow proportion of being spilled. Failure to find a relationship between flow and an index of survival does that survival is not prove independent of flow. The relationship found for spill may be influenced by some of the dam operations problems discussed earlier.

Filardo (1990) found that a multiple regression model (using a limited data set (1985-1989) for conversion

rates that had been provided by the U.S. v. Oregon Technical Advisorv Committee) based on September flows and in water temperatures the Snake River explained a significant amount of the variation in conversion rates of wild/natural Snake River "B" steeihead from the Zone 6 fishery (between Bonneville and McNary dams) to Lower Granite Dam.

The effect of flow on travel time and survival of adult migrants in dammed and impounded reaches is

difficult to establish because of the overlap between dam-related and reservoir related passage mortalities. Inter-dam conversion travel rates and times for groups of unmarked or batch-marked fish do not appear to be highly Also@ reliable. much of the early research was conducted when dams lacked some modifications (e.g., spillway deflectors) that reduced the adverse effects (e.g., nitrogen supersaturation) of high flow. 3. Water Quality Associated with Flows quality There are many water factors correlated with flow level. however, only temperature will b addressed here.

a. Temperature

is Much known about the biological significance of temperature for poikilothermic animals such as

26 salmonids, and numerous observations suggest that. under some conditions, water temperature exerts a great influence on the success of adult passage. Columbia River water temperatures commonly exceed 21'C (70'F)

during August (Collins 1963: Shew et aL 1985: Meyer 1989), while those in the Snake River can exceed 240C

(75'F) (Sylvester 1958; Collins 1963: Thompson 1974; Liscom et al. 1985). These levels equal or exceed lethal temperatures that have been measured for Columbia River steethead and chinook stocks migrating during the hottest seasons, and "ecological death" due to equilibrium loss occurs at even lower temperatures (Coutant 1970). High temperatures can also aggravate the adverse effects of nitrogen supersaturation

(Coutant and Genoway 1968; Ebel 1969; Beiningen and Ebel 1970), stress (Trefethen and Sutherland 1968), low dissolved oxygen (Fry 1971), and other factors (Andrew and Geen 1960).

There is abundant evidence that existing Columbia and Snake river mainstem temperatures are indeed adult detrimental to migrants. Temperatures appear to explain some of the variation in conversion rates of some steelhead stocks migrating through the lower reaches of the Snake River (Filardo 1990), and high

temperatures were probably important in failures of radio-tagged fish to pass upstream (Trefethen and Sutherland 1968; Shew et aL 1985). Migration delays caused by high temperatures have often been observed, particularly where tributary temperatures exceed those of the mainstem Columbia River (Major and Mighell

1966; ODFW 1977; Johnson et aL 1982: Liscom et aL 1985). Most mortalities or reductions in spawning success caused by high temperatures probably go undetected. Impoundments have probably contributed to the problem of high temperatures. which may have already existed to a lesser extent. Crawford et aL (1976) concluded that the water temperatures of the Columbia River and its tributaries have been altered bv man's activities. While the nature and extent of these changes is uncertain. the thermal cycle appears to have been shifted from pre-impoundment conditions, so that lower water temperatures occur in the spring and early summer and hiizher water temperatures occur in the fall. The result of this widening ranQe of water temperatures may not change the average water temperature in the basin. This increased range does. however, present new risks to the anadromous fish species that depend on acceptable seasonal water temperatures to survive critical migration. spawning, emergent and rearing stages. Sylvester (1958) describes harmful temperatures in the Snake prior to construction of the lar ge mainstem projects and notes factors that determine how impoundments affect downstream temperatures.

Large impoundments with deep outlets (e.g., Lake Roosevelt behind ) can actually moderate seasonal temperature extremes and delay the annual temperature cycle (Jaske and Goebel 1967: Raleigh and Ebel 1968; Gregoire and Charapeau 1984; Zimmerman 1984). Thermal stratification, which permits this buffering effect, can also occur in at least one Columbia River "run-of-the-river" project (e.g., see Beiningen and Ebel 1970; Raphael 1961, cited Liscom et aL 1985).

The lowest flows of the vear, which formerlv occurred in winter, now occur durina some of the hottest months, July-October. When flows are. low, climatological conditions have a greater effect on water temperature (Sylvester 1958; see discussion in Andrew and Geen 1960: Ralston 1974). Also, when water

27 is velocity relatively independent of flow through a river reach (eg., Hells Canyon on the Snake River), flow

(quantity of water) becomes more important for heat accumulation (Koski 1974). Tanovan-(U.S. Army Corps

of Engineers, personal communication) modeled temperature response to within-day flow shaping on the Snake River (ie. "zero" nighttime flows). He concluded that water temperature. which often exceeded 20'C, was relatively insensitive to flow changes over such a short period. During filling of Lake Roosevelt in 1941,

many sockeye perished below Celilo Falls during unseasonably low flows and high water temperatures (Fish and Hanavan 1948). The correspondence between the lowest annual flows and the highest annual

temperatures, combined with the increased sensitivity of in-river water temperatures to atmospheric conditions, leads to the observation that water temperature is strongly and negatively correlated with flow. 4. Conclusions

Widespread occurrence of salmonid spawning migrations that are correlated with peak annual flows suggest the importance of relatively high flows for reaching the spawning grounds and reproducing successfully. There are mainstem-spawning stocks which require greater regulation of news to protect incubating eggs and fry from highly variable power peaking flows. This is particularly true for the Columbia River fall chinook which spawn in the Vernita Bar area of the Hanford reach below Priest Rapids Dam.

Migration delays and undue use of limited energy reserves can result in prespawning mortality. High

velocity water appears to be very attractive to upstream migrants. and moderate velocities may improve migratory performance, between dams. However, high project flows can interfere with passage at dams because fishway attraction flows become a small percentage of total flow. While high flows do exacerbate the problem, inadequate facilities and operation of the dams are also part of the problem. Mortality of adult salmonids at mainstem dams increases under high flows and is probably caused by delay associated with the inability of adults to find fishway entrances and fallback associated with high levels of spill.

Numerous mortalities and delays have been associated with excessive high water temperatures. These high temperatures are common in the mainstern Columbia and Snake rivers during August and September.

Temperature may be as important as flow for adult passage during this time. High temperatures are probably impacting the spawners and their progeny. Mainstem reservoirs lack may velocities that are adequate for orienting and stimulating upstream

migration. Salmonids LI can apparently use behavioral mechanisms other than rheotaXiS LO Mi rate through slack-water reservoirs. C. Other Species

White Sturgeon (Acipenser transmontanus) is another species of significance that has been impacted by

the development and operation of the Columbia River hydropower system. Originally this species was so

it fi abundant that posed a nuisance to salmon ishers. Smith (1895) quoted an early cannery operator, M.J. "In 1879 Kinney as stating the sturgeon were so thick in Baker Bay that we did not consider it safe, early in gill fish the season. to put our nets out. The were so numerous and laree that thev were able to destroy a

28 great amount of netting."

However, in the late 1880's freezing plants for processing the commercial catch were built. An intense Sturgeon fishery rapidly developed and the harvest peaked at nearly 5.5 million pounds in 1892 before collapsing to less than 75,000 pounds in 1895; the population did not recover until the 1950's when restrictive regulations were adopted (Craig and Hacker 1940). Although over-harvest originally decimated the population, the development and operation of the hydropower system is also considered to have impacted white sturgeon populations by blocking their migrations, and altering the free-flowing habitat to more stillwater conditions. Presently, white sturgeon populations below Bonneville Dam -are considered healthy and since 1979 have supported the principal sport fishery.in the river (ODFW/WDF 1987). However, the status of their populations upriver from Bonneville Dam ranges from uncertain in most reservoirs, toseverely depressed in the Kootenai and Snake rivers (Graham 1981; Cochnauer 1983).

1. Biology

a. Spawning

Whitesturgeonspawnduringthespringatpreferredwatertemperaturesofabout 10-15'C. Temperatures exceeding 18'C may cause substantial mortality (Wang et aL 1985).

Eggs have been collected in the Columbia River from 19 April to 19 July, at water temperatures of 10 to 20'C, or temperatures exceeding levels known to cause mortality. Spawning activity has been determined to be interrupted, or be extended past the preferred temperature range, when the discharge from the dams has dropped or been below average (La Voy et aL 1989; Parsley et aL 1989).

'ne eggs are randomly broadcast over rocky substrate in turbulent areas caused by high current velocities (Scott and Crossman 1973). Spawning in these areas ensures the huge numbers of adhesive eggs are widely scattered, thereby reducing the probability of predation and spread of fungi, which would occur on dense concentrations of eggs. In the Columbia River, white sturgeon spawning has only been documented immediately downstream Bonneville, of The Dalles, John Day and McNarv dams - the areas of highest water velocity (Kreitman 1983; Palmer et aL 19M Parsley et aL 1989). b. Larval Stage and Flow

When the eggs hatch, the larvae swim up into the water column and are dispersed downstream by the current. The larvae remain in the water column for longer time periods at lower current velocities than when velocities are higher (Brannon et aL 1985). After this period of dispersal. the larvae seek cover from predators by burrowing among rocks or detritus while their yolk sac is absorbed. Upon emergence after 10-

14 days the larvae begin actively feeding. If food is unavailable, they swim-up into the water column and are again displaced downstream to new foraging areas by the current.

In the below average flow years of 1987 and 1988 very weak year-classcs of white sturgeon were established in Bonneville, The Dalles and John Day reservoirs (Parsley et aL 1989). In those years, it is not known whether current velocities were insufficient to stimulate reproduction, disperse eggs, or disperse larvae.

29 Water temperatures, bottom substrates, food and supply did appear adequate in these reservoirs for success reproduction and rearing of white Therefore, sturgeon. water velocities appear to be the most limiting factor. But, the magnitude velocities, of as well as the magnitude of the flows necessary to provide these velocities in different river reaches, is unknown.

30 III. FLOW AND WATER VELOCYfY

Earlier discussion in this report covered the importance of moving smolts through the Columbia and

Snake river reservoir system in a biologically timely manner if an acceptable smelt survival, and subsequent adult production level is to be achieved. Because of the relation between water movement and smolt travel time survival, is and a critical concern the rate of water movement through the present reservoir system compared to the pre-project, free-flowing conditions., A. Water Particle Movement 1. Storage Replacement Method

Figures 7A, 713, and 7C illustrate water particle travel time through each project in the lower Snake, mid-

Columbia, and lower Columbia river reaches, respectively, at different streamflow rates. These curves, and related curves presented later, are based upon the, "storage replacement method" using reservoir data provided by the Reservoir Control Center. AJso shown on each figure are the combined effect of all the projects in each reach on water particle travel time through the reach. and the May flow level needed for smolt migration, as listed in the agencies and tribes flow proposal.

Snake River reservoirs all have similar characteristics in terms of water particle travel time at different flow through rates. Water takes the longest to move Little Goose reservoir, which has the largest storage volume of the four reservoirs, and takes the least time to move through Lower Monumental reservoir, which has the smallest storage volume. Travel time curves are shown on Figure 7A for these two projects. Since travel times in Lower Granite and Ice Harbor reservoirs fall between the curves plotted, they are left out for clarity.

Water particle travel time through the entire I3)8 mile reach from the head of Lower Granite reservoir

is I to Ice Harbor Dam neariv 3 davs at a flow of 85 kcfs. Travel time rapidly decreases as flows are increased, to about six davs at 140 kcfs.

7B Figure presents the same information for the five mid-Columbia River projects. There is a greater difference among these projects than among the lower Snake projects. with Wanapum reservoir producing the longest water particle travel time and Rock Island reservoir the shortest. Since nearly the same total is water volume stored, all five mid-Columbia projects combined produce an effect similar to that of the lower

Snake projects. Therefore, nearly six days are required for water particle movement through the mid- flow 140 kcfs. in Columbia reach at a of The same rapid increase travel time with decreasing [lows occurs. reaching about 14 days at 60 kcfs.

Figure 7C shows a much greater difference among. the four lower Columbia projects. As would be expected. water particles take much longer to move through John Day reservoir, because it has a verv large volume of water to move. Unlike the other run-of-river projects, John Day is actually classified as a storage project. The Dalles reservoir has the shortest travel time of the four. Water particles take nearly 8 days to move through the lower Columbia reach at a flow level of 300 kcfs. with about one-half of that time movincy

31 ((A)

20

is 14 Pmj@ LOS

1140 kcf3 10

0

0 100 200 300 4GO Flow (kcfs)

20

15

140 kch

10

5

0

0 100 2 0 ;00 400 Flow (kcfs)

@o

15

300 kfs lo

5

n

0 loo 200 300 4 5 0 600 Flow (kcfs)

Figure 7. Water particle travel time, in days, through: (A) lower Snake River; (B) mid-Columbia Riven and (C) lower Columbia River reservoirs.

32 through John Day reservoir. Travel time is substantially increased with decreasing flows is 13 and approximately days at 200 kcfs.

The significance of the present water particle travel times becomes more evident when pre-project, free-flowing compared with water movement. This is comparison made in Figures 8A, 813, and 8C for the lower Snake, mid-Columbia, and lower Columbia. respectively. In a free-flowing state before projects water particles were built, traveled the entire lower Snake River reach in less than one day at all flows above 40 (Figure 8A). Under kcfs present conditions, it would take extremely high flows begin historical to to approach those travel times. The same statements can be for repeated the mid-Columbia River reach because the (Figure 8B) of great similarity to the lower Snake reach. For the lower Columbia under free flowing conditions (Figure 8C), water particles moved through the reach in less than two days all flows kcfs. Obviously, at above 80 the extreme slowing of water movement bv caused the I 1 projects under discussion created a has severe obstacle to smelts adapted to a mostly passive mode of transportation in higher flows. 2. Average Velocity Method.

To provide a common basis for direct comparison , of the relative effect. of each reservoir on movement, water particle water travel times in days were converted to average velocity in through water miles per hour each reservoir. This takes into account the effect of different cross-sectional The areas among reservoirs. results are illustrated in Figures 9A, 9B, and 9C for the lower Snake, mid-Columbia, projects, and lower Columbia respectively. Also shown are the average velocities through each reach at different flows, both with the projects and under free-flowing, historical conditions, and the Mav flow proposal levels. Water velocity versus flow plots rate as a straight line for the reservoirs because, with the volume of moving being the water constant, velocity increases proportionately as the flow increases. Free-flowing plot conditions as, a curve because the volume of water, or cross-sectional ;ncreases area. as flow increases. It should be noted, however, that the water velocitv in a reservoir will be near Ehat of free-flowing conditions head of the reservoir near the where the water is the shallowest and the cross-sectional area the will decrease smallest. The velocitv as the reservoir becomes deeper and the cross-sectional larger, area becoming much slower than the averages shown in Figure 9 at the location of maximum cross-sectional This also flowing area. applies to the free- streams, which have very high velocities through shallow steep, sections and low in pools. velocities deer)

The lower Snake projects do not differ appreciably in their reservoir volume length in similar to ratios, resulting average water velocity a given at of flow. Figure sho.@s rate 9A that Lower Mon has the urnental reservoir greatest slowing effect and Little Goose reservoir the least. The water velocitv curve for Granite reservoir Lower nearly coincides with the curve for the entire reach. and is shown level not separately. At a tlo%v of 140 kefs, average velocity water through the four is I reservoirs about one mile per hour, to eight miles com p ared per hour with free-flovring conditions. The water velocity/flow relations vary much more among the mid-Columbia projects (Figure 9B). Of the

33 (A)

45

40

35

All 4 Proj.M 30 -

V 25 E 140 kcfs - 1 20 - I 15

10

5 PM.Pmj=

0

0 100 200 300 400 Flow (kcfs)

45 - 40 - 35 - 30 - AJI 5 Pmj@ 25 E 1140 kcf, 20 M > 0 415- i 10

5

0

0 100 200 300 400 How (kcfs)

45 - 40 - 35 - Al I 4 P,.,@ 30 - 2.5 300 kas 'E - 20 - 15

10

5 P.-P@jwt

0 I

0 100 200 300 400 500 600 Flow (kcfs)

Figure 8. Water particle travel time. in days, through: (A) lower Snake River: (B) mid-Columbia River; and (C) lower Columbia River reservoirs, showing pre-project (free-flowing) and post-project water movement.

34 (A)

9

8

6

140 kf. .2

4

E! 3

LGR 4 Ws

0

I

9 100 200 300 400 Flow (kcfs) -

9

a

7 RJS

E 6

140 kcf.

4

2 3 WEL RRH ?RD & MI 5 2 Wm

0 100 200 300 400 Flow (kcfs)

(C)

7

_E 6

5 1300 kcf.

4

WH 3 MA

NJ 4 ,MCN JDA

0

0 100 200 300 400 Soo 600 Flow (kcfs)

Figure 9. Water velocitv particle I in miles per hour (mph), for: (A) lower Snake (B) @ River; mid-Columbia River, and (C) lower Columbia River reservoirs, compared to pre-project water particle velocities.

35 live projects, water moves the slowest through Wanapum reservoir and the fastest through Rock IsIt'.0 reservoir. The relatively small volume, for its length, in Rock Island reservoir results in the water movin-a

nearly three times as fast compared Wanapum reservoir. Priest is to The Rapids reservoir curve not shown separately, because it coincides with the velocity average curve for the entire reach. Average water velocity is for the river reach about one mile per hour 140 kcfs, 8.5 at compared to about miles per hour under free- - flowing conditions.

Of the four lower Columbia projects, water moves the slowest through John Day reservoir, but not much - faster through McNary reservoir because it also has a fairly large storage volume relative to its length (see Figure 9C). Water moves the fastest through Bonneville reservoir at a given rate of flow. Average water velocity through the entire reach a flow 300 kds is at rate of about one mile per hour, compared to about 8.5 miles per hour under free-flowing conditions. . To tie all of the foregoing discussion of water movement and fish flow needs together, Figure 10 provides a rough theoretical example of total particle travel water time from the head of Lower Granite reservoir, to Bonneville Dam several flow levels, at compared to travel time under free-flowing conditions. The very short time increment for existing the free-flowing reach from Priest Rapids Dam to the head of McNary reservoir is included in calculations in not the or the figure. Travel times would apply for the river reach from the yed* head of Wells reservoir to Bonneville Dam. Under low flow in conditions such as have occurred recent (50 kcfs in the lower Snake River and 110 kcfs in the lower 3 Columbia River), water took about days to move through the river reach. from the head of Lower Granite reservoir to Bonneville Dam, under pre- project, free-flowing conditions. Today, under existing conditions. water takes about 39 days to travel the river reaches, in same a fifteen-fold decrease velocity under which anadromous fish evolved. At flow levels of 140 and 300 kcfs, pre-project travel time was about 1.7 days compared to today's travel time of about 14 davs a nine-fold increase. --- Even at hiah flows (200/430 kcfs) or at t@c lower end of flood flows (100/600 kcfs), @yater now takes about seven times as toniz to move through the -'!,,er reaches. compared to pre-project conditions. In short, pre-project travel times are unattainable with the present reservoir system, but travel can be substantially improved, times and sinolt migration conditions improved, by increasing flows over the low flow conditions characteristic of recent vears. B. Smolt Movement

1. Smolt Versus Water Particle Travel Time

Based upon smolt migration data for the 1971 nine years of through 1981, Sims et aL (1982), showed a close correlation between smolt travel time and water particle travei time through the reservoir system. Marked groups of hatchery yearling chinook (1982-1989) and steellie-ad (1982-1989) from the Snake Riv drainage, and in-river summer migrants from the lower river (1981-81; in 1986-88) John Day reservoir corroborate those earlier findings. Plots of water particle travel time, as calculated by the storage replacement method predicted versus sinolt travel time from these later years are presented in Figures 11

36 40 39.3

WATER PARTICLE TRAVEL TIME from Head Of Lower Granite Reservoir to Bonneville Dam N

Pre-Project

E 20 Existing Conditions

lo

0

Low Flow High Flood Travel Time Flow Proposal Flow Flow Ratio 1:15 1:9 1:7 1:6

Figure 10. Water particle travel time, in davs, from the head of Lower Granite reservoir to Bonneville Dam at various streamilows, compared to water particle travel time tinder pr e-project river conditions.

37 A.) 20 -

18 Observed Fish - I travel time 1982-1990 16 - Predicted Fish Travel Time 14 >1 - Water Particle Travel Time . . . . 12 -

10 -

8 > Ca

6

4

2

0

B.) 20

Observed Rish Travel Time 19824989 16 Predicted Fish Travel Time 14 Witer Particle Travel Time . . . . 12

F- lo

6

4

2

0

40 60 80 100 120 140 160 180 200 Flow (kcfs)

it Ice Harbor Dam

Figure 11. Water particle travel time, predicted fish travel time. and, observed rish travel time vs flow for: (A) Snake River vearling chinook salmon; and (B) Snake River steelhead.

38 and 12. The predicted smolt travel times are from the bivariate relationship between smolt travel time and flow.

Smolt travel time closely parallels particle travel time water --- decreases with increasing flow levels and increases with decreasing flow levels, reflecting the earlier discussion of biological 'factors affecting migration.

The important aspect of these plots is the striking similarity in the general form of the, relation between average flow and either water particle travel time, or fish travel time, Water particle travel time is simply a function flow of and the cross@sectional area of the waterway. The similarity of fish travel time to water particle travel time indicate a causative, rather than simply a correlative, relation between flow and travel time of juvenile salmonids.

C. Wavs to Meet Increased Water Velocitv Needs 1. Reservoir Drawdown

One option for decreasing travel time that has been suggested. especially through the lower Snake River

is reservoirs, to operate at lower reservoir elevations, thereby decreasing storage volumes to be moved, which would in turn increase water velocity. The following section discusses the physical potential of that option,

applied to the lower Snake River, without examining the substantial potential adverse impacts on the various

uses of the reservoir system. A family of curves showing water particle travel time at different rates of flow for Lower Granite reservoir operated at different elevations is shown in Figure 13. The curves are for 10

foot increments of drawdown from full pool elevation of 740 feet above mean sea level to 90 feet of drawdown at elevation 650 feet. Normal operating range at Lower Granite Dam is between elevations 733 and 738 feet.

At the desired flow level of 140 kcfs, and with the reservoir at full pool, water travel time through the is reservoir nearly two days (about 45 hours). To maintain that same travel time at a lower flow rate, say 85

kcfs, would require operating about elevation 710 a 30 t is at feet --- fee drawdown. If the purpose to meet fish flow needs with an even lower flow rate, sav 50 kcfs. a 50 feet drawdown to an operating elevation of 690 feet would be required. If the intent is to reduce both the flow rate and travel time, even larger drawdowns would be required, the magnitude depending upon the flow rate and travel time objectives selected. In anv case, it is evident that applying this method to meet fish passage needs without providing higher flows would require major reservoir drawdown,

A variation of the above option was examined, based upon the fact that the very process of drawing down

a reservoir results in increased outflow during the time that the reservoir is being drawn down. Figure 14A shows the magnitude and length of time of flow augmentation that could be sustained at different controlled drawdown rates to different elevations using Lower Granite reservoir. For example. a sustained flow increase

10 of kcfs could be maintained for about three days by drawing down within the normal operating range of 738 733 feet. elevation to That same flow increase could be sustained for four days by drawing down to elevation 730 feet, and for six days by going to elevation 725 feet.

39 40 - Observed Fish 35 Travel Time - 1981-1983, 1986-1988 30 - Predicted Fish Travel Time >1 25 - %ter Particle Travel Time 20 00 E- -

> 15 - 400 0

10 -

5

......

0 ------50 100 ISO 200 250 300 350 400 Flow (kcfs) at John Dav Dam

Figure 12. Water particle travel time, predicted fish travel time. and observedfish travel time vs flow for subyearling chinook salmon in John Dav reservoir.

Carrying this approach one step further. Figure 14B shows the flow augmentation possible if all four lower Snake reservoirs were simultaneously drawn down within their normal operating ranzes. The Lower Granite reservoir curve is shown Separately for comparative purposes. For example, usine Lower Granite reservoir alone (see Figure I 1) a flow increase 5 kcfs of could be sustained for about five davs. When the simultaneous increase from each reservoir is added, five the dav sustained flow increase at Ice Harbor reservoir would be 15 kcfs.

In view of the very low spring (April - June) flows that often in occur the tower Snake River, it does riot appear that the above approach is a useful for fish option meeting flow needs over the necessary time period. even without considering the mariv adverse effects, such as the need to later reduce flows in order to refill the reservoirs.

D. Conclusion subsequently* In summarv the relationship between water particle travel time and Smelt travel time. and smolt Survival and adult production, further supports the smolts' need for flow. The relation between river now and water particle velocity changed dramatically between pre- and post-hydroelectric development

40 Geller

9 - 740

7 - 730

6 - 7:00

140 kcEs

E 7

4 > 700 3 6:

6 0 2 670 660 650

0 50 100 150 200 250 300 Flow (kcfs)

Figure 13. Water particle travel time, in days, for various strearnflows through Lower Granite reservoir. periods. Increases in cross-sectional area now require much larger changes in flow, even above historic flows, simulate to pre-impoundment water velocities. Pre-development water particle velocities. which were the velocities upon which salmon evolved. can never be met in the present river configuration. Reservoir drawdown as decrease to the cross-sectional area and increase velocities mav not be a viable option. Compared in to increasing flows order to substantially decrease water particle travel time, and thereby increase smolt migration rate.

41 No. of Sustained How I)ays No. of Sustained Flow Days

0 4@

CM

rj IV. JUSTIFICATION OF FLOWS PRESENTED IN THE FLOW PROPOSAL A. Fish Flow Objectives

In order to avoid the implementation problems the Water Budget faces today and to address concerns for it is the future, absolutely essential that fish flow requirements be incorporated into power planning as

hard constraints and as part of system rule curves. Otherwise, there is no assurance that fish flows will

actually be met, and a strong probability that they will not. The most effective way to implement hard constraints and rule curves for fish flows is to establish a fishery flow objective, and the flow levels necessary

to achieve the objective. The objective of the flow proposal is to provide flows throughout the Columbia

River that will yield adequate survival of critical life stages of anadromous salmonid stocks. Flows should move juveniles from fresh water to the estuary Arithin the appropriate "biological window" discussed earlier; stimulate upstream migration of adult salmon; provide optimal spawning habitat and water conditions for survival of emerging fry and rearing of fry and juvenile salmonids; and also allow Cor project passage facility operations (Table 2) (CBFWA 1990b).

While there are undoubtedly costs associated with this change from current practice. there are significant advantages as well. First, hard constraints and rule curves are intended by the agencies and tribes to accomplish putting fish flow planning and operation at the same level as power and other uses of the hvdrosvstem. Second, they will provide improved predictability and certainty for power planning, which should allow for more efficient generation. Third, they will put an end to costly and time consuming disputes @ '6 A -t over fish flows, including disputes over future power planning and operations. 1-t@w -@, B. Justification For Flows

The recommended criteria for fish flows are comprehensive. They include provisions of flow for: (1) successful juvenile migration in a timely manner: (2) adequate operation of adult passage facilities: (3) successful reservoir migration by adults: (4) adequate spawning and emergence of wild fish: and. (5) reservoir levels that are sufficient for resident fish (resident fish needs are considered in a separate Authority flow in document). The levels identiliccl this proposal are based on the best available technical data as to the biological needs of the fish. Although biological data concerning anadromous fish may be subject to differing interpretations, the recommended flow levels reflect a careful analysis and evaluation of available ckita performed over a period of several years, and the best collective judgement of the agencies and tribes on biolodcally adequate flows for fish. The objective of these flows is not to recreate pre-impoundmenL migration conditions or travel Limes. The travel time to the ocean is substantially greater under the flow proposal than the travel time under the natural conditions. It was recognized earIv on that historic velocities travel fish and times that the runs adapted to over thousands of years could not be achieved in the present hydropower system. because of the reservoirs. However, the flow levels chosen would achieve substantial improvement over existing migration conditions in terms of reduced travel time, reduced exposure to

43 2. Table Fish flow recommendations, kefs: minimum instantaneous/dailv average

MONTH THE DALLES PRIEST RAPIDS ICE HARBOR

January 80/80a 70/70" 10/20

February 80/80a 70/,70b 10/20

March 80/80a 70/70" 10/30c

April 1-15 8(/250d 70/14(d 30/1 00@ April 16-30 80/300d 70/140d 30/140d

May 80/3(od 70/140d 30/140d

June 1-15 80/300" 36/140d 30/140d

June 16-30 80aooe 36/12W 30/8V

July 1-15 80/200c 36112V 30/80e

July 16-31 80/160e 36/110e 30/50 C

August 80/160e 36/11W 10150c

September 80/803 36/40 30/35c

October 1-15 80/80a 36/40 30/35c

October 16-31 80/80a 50/125' 30/35c

November 1-15 SO/Soa 501125' 30/35c

November 16-30 80/80' 50/125' 30/35c

December 80/803 70/70b 30/35c

Adult passage, Bonneville Dam b Incubation, emergence ' Adult passage d Juvenile migration: steelhead, sockeye, chinook. coho Juvenile migration: subyearling chinook Spawning/maximum

predators high and water temperatures. and increased survival. In the view of the agencies and tribes. these flows "adequate" biological represent flows for fish rather than the "optimum" flows based an historic conditions that fish stocks adapted to.

AR juvenile salmon travel,time versus flow relationships are characteristically curvilinear. At low flows anv increase in flow is bv matched a substantial decrease in travel time. Above certain flows the corresponding decrease in travel time is not as large. The flow levels chosen represent the area of the curvilinear relations where increases in flow do not yield corresponding large decreases in travel time.

1. Spring (April I - June 15)

The agencies and tribes have recommended flows of 140 kcfs for the lower Snake. and mid-Columbia and 300 kcfs for the lower rivers Columbia River. These flow levels were selected to substantially reduce

44 smolt travel time and increase smolt survival to provide significant progress towards the Authority and Council goal of doubling adult runs in the Columbia Basin (CBFWA 1990a).

The recommendation of 140 kcfs for the lower Snake River is based on travel time versus flow and survival versus flow relationship from 1973-79 migration studies of yearling chinook and steelhead in the Snake River by Sims and Ossiander (1991). Significant relationships were found between travel time and survival of yearling chinook and steelhead migrating between Little Goose or Lower Granite dams and The Dalles, Dam and flow at Ice Harbor Dam (Figures 15 and 16). These travel time versus flow relationships are similar to earlier findings (Bentley and Raymond 1968; Raymond and Sims 1979; and Raymond 1979) and to relationships found for 1981-89 hatchery chinook and steelhead (see Figure 11).

The agencies and tribes flow recommendation submitted to the Council in 1981 called for flows of 300.

kcts in the lower Columbia River during the spring migration. This recommendation is based on travel time

versus flow and survival versus How relationship of Snake River yearling chinook and steelhead by Raymond

1968, Collins et al. 1975, and Sims and Ossiander 1981 (Figures 17 and 18) which is consistent with travel time versus flow relationships found for yearling chinook in the John Day reservoir during 1973-83 and 1986- 90 (Figure 19). The significant correlations of travel time and survival of Snake River yearling chinook and

steelhead to flows at Ice Harbor Dam (Figures 15 and 16) and the Dalles Dam (Figures 17 and 18)

demonstrate that the travel time and survival of chinook and steelhead from the Snake River to the lower Columbia River are dependent on flow conditions in both the lower Snake and Columbia rivers. Based on simulations with the Svstern Planning Model for the Middle Fork Salmon River (CBFWA

19190a), inoreasingflows at Lower-Granite Darafrom average (93 kc1s) to therecommended(140kcfs)spring flows would increase escapement of wild spring chinook by 2.8 fold (729 to 21843 adults) and steelhead by

.1.1 fold (5152 to 16177 adults). -'Additional adult recruitment modelinQ by P c trosky (1991) (Figure 20) and

the Authoritv during development of the Integrated SyIs tem Plan (CBFWA 1990a), also suggest that recommended spring flows may increase adult returns to the Snake River. Although a lot of the variation is unaccounted for in these relationships, there is a trend toward increased adult production of wild and hatcherv chinook as flow increases from 40-146 kcfs.

16 2. Summer (June - August 31)

During the negotiations of the flow mitigation concept that led to the present Fish and Wildlife Program

(Program), the needs of summer migrants were not well understood. Studies conducted to that time inferred

that the migrational characteristics and needs of summer downstream migrants were not the same as those that had been documented for spring migrants. The simulations of the mitigation flows for fish indicated that

summer power base flows to meet loads would be adequate to protect summer migrants in all but critical periods. The Program was subsequently adopted without specific flow provisions for protection of summer migrants. However, the 1981 Program included language regarding the further investigation of the migration of summer migrants.

45 (A)

40 1977 Ln Y 7.79 0.87Ln X R2 = 0.90 (P<0.01) Predicted Smolt Travel Time ..... %ter Particle Travel Time 30

1973

20

1976 1979

197 ...... :975 >1 ...... 1974 10

40 60 80 100 120 140 160 ISO

> 40 1977 Ln 6.14 Y - 0.73Ln X R2 = 0.58 (P<0.05) Predicted Smoft Travel Time ..... %ter Particle Travel Time 30

20 1973 1976

1 75 10

40 60 so 100 120 7110 160 130

Flow (kcfs) it Ice Harbor Dim

Figure 15. Travel Of time. in davs, Snake River. (A) yearling chinook and (B) steelhead from LOWer Granite or Little Goose dams to The Dulles Dam vs. flow at Ice Harbor Dam (t7 days of migration peak) (from Sims and Ossiander 1981).

46 (A)

60

Ln Y -7.42 + 2.29Ln X R2 (P<0.01) 50 =.83 Predicted Smolt Survival

40 74 1978

30

1979 1975 20

10

> 1973 0

40 60 80 100 120 140 160 180 C4 (1) D I CZ / Ca (-D) C 60

La Y = -9.53 + 2.65Ln X R2 .92 (P<0.0 1) so Predicted Smoit Sumval 1975

40

30 1976

1978

20

10

0 j 40 60 80 100 120 140 160 180 Flow (kcfs) at Ice Harbor Dam

Survival Figure 16. of Snake River. (A) yearling chinook and (B) steelhead from Lower Granite or Little Goose dams to The Dalles Dam vs. flow at Ice Harbor Dam (±7 davs of migration peak)(Sints & Ossiander 1981).

47 (A)

40

1977 X La Y = 7.77 - 0.90La R2 0.79 (P<0.01) Predicted Smolt Travel Time

30

1 3 20

1976

>1 10 - 100 125 150 175 E 200 225 250 275 300 325 350

> CZ 40 1@977

La Y = 7.58 - 0.87Ln X R2 = 0.67 (P<0.05) -Predicted Smelt Travel Time

30

20 1.(73 1976

1974

1978 10 1975

100 125 ICO 200 225 250 275 300 325 350

Flow (kcfs) a[ The Diiles Dam

Figure 17. Travel time, in days, of Snake River. (A) yearling Chinook and (B) steelhead from Lower Granite or Little 6oose The dams to Dalles Dam vs. flow at The Dalles Dam (±7 davs of migration peak) (from Sims and Ossiander 1981).

48 (A)

50 Ln Y -11.03 + 2.51Ln X R2 0.87 (P<0.0 1) Predicted Smolt 1974 40 Survival 1978

30

1975 1979

1976 20

10

> 1977 7 0

C40 100 125 150 175 200 225 250 275 300 325 350

CZ

rz Q so U La Y = -13.60 + 2.95Ln X R2 0.93 (P<0.01) Predicted Smolt 19;5 40 Survival

30

1976 1978

1974 20

10

197

0

100 125 150 175 200 225 250 175 300 325 350

Flow (kcfs) it The Dalles Dam

18. Figure Survival of Snake River: (A) yearling Chinook and (B) steelhead from Lower Granite or Little Goose dams to The Dalles Dam vs. flow at The Dalles Dam (±7 days of migration peak)(Sims & Ossiander 1981).

49 14 -

Travel Time 12 - 1986-90 SMP %ter Particle lo Travel Time ...... ;>1 - Travel Time 1973-83 MNFS 8 E Observed Fsh Travel Time 6

>

4 %

2

0

100 150 200 250 300 350 400 Mow (kcfs) at John Day Dam

19. an* Figure Travel time. in days, vs flow in John Day Pool for (1) Yearling chinook freeze branded released at McNary Dam between 1986-1990 under the Smolt Monitoring Program; (2) for yearling chinook freeze branded and released at McNary Dam between 1973-1993 by the National Marine Fisheries Service; and (3) water particle travel time.

The changes in hydrosystern operations, the decline of summer flows, the use of energy exchanae agreements, and the changes in extreme marketing and load shape over the recent years have created serious for concerns summer downstream mi-rants. The changing dynamics of the hvdrosvstem manipulates flows so that flows higher in fall are and winter months when power markets are strong, and lower in spring and

is summer months when demand less. For example, summer flows in 1989. with a January-July runoff volume of 90.1 MA-F, as low were as flows that occurred in 1987 and 1988. when runoff volumes were 77 and 76 MAF, respectively. Flows in 1977, which had a lower Januarv-Julv runoff volume, were higher than the flows which occurred in 1987, 1988 and 1989. The flow summer requirements contained in this proposal were developed from a[[ the existing passage and migration data on summer miarants collected through the Smolt Monitoring Program and earlier monitoring efforts. The flow requirements of summer migrating juvenile salmon were monitored in 1981- 1983. The results -that of investigation were largely inconclusive. The studv was not desianed to address all of the variables that affect the movement of summer migrants. The study did indicate that the migration

needs of summer migrants were different from those of spring militants. However, the "low flow" travel time estimates generated for the 1981-1983 studies were for groups that were migrating later in the season. In

50 general, summer flow levels decrease throughout the season. The travel time estimates associated with the late migrants may be influenced by other variables, such as size and increased physiological development.

Subsequent to the 1983 study, summer migrants were monitored through the Smolt Monitoring Program.

Mark groups from hatcheries and other studies, such as transportation controls, were utilized to monitor summer migrant movement. The re-analysis of the 1981-1983 summer migrant data combined with migration data from 1986-88 indicated that a statistically significant relationship exists between flow and travel time for summer migrants (see Figure 12).

1 3. Fall (September - November 30)

The Columbia River upriver fall Chinook, which primarily spawn at Vernita Bar and throughout the

Hanford Reach of the mid-Columbia River, are an extre mely important stock of Chinook salmon due to

Columbia River and North Pacific fishery management considerations. This stock of Chinook has been very

productive in recent years (1986-1988), with adult populations in excess of 500,000 and recent spawning escapements in excess of 100,000. '17his currently productive stock, although showing signs of decline in 1989 and 1990, contributes substantially to ceremonial, commercial, and recreational fisheries from the Columbia

River to Southeast Alaska.

To maintain the viability and productivity of this stock, it is essential to protect the natural spawning and

rearing habitat at Vernita Bar and within the Hanford Reach. There are no tributaries to the Columbia River between Priest Rapids Dam and the head of McNary reservoir. 'ne flow and flow regulation (fluctuations at Priest Rapids Dam completely controls the available spawning and rearing habitat in the

Hanford Reach. Because Vernita Bar is near the upstream end of the Hanford Reach, four miles below

Priest Rapids Dam, the habitat at Vernita Bar is more dramatically influenced by flow regulation at Priest Rapids than the remainder of the Hanford Reach.

Adult Columbia River upriver fall Chinook enter the Hanford Reach beginning about mid-August and continue through November. The peak period of adult entry to this area is the month of September. The mature adults seek out suitable spawning habitat and begin spawning about October 15. '17he female Chinook select the redd (spawning nest) location, and construct their redd bv excavating a pit in the river bottom with

their tail. The precise environmental parameters that individual female Chinook utilize in site selection for redds are not totally known, but primary parameters include water depth. water velocity, and substrate. The combinations of depth, velocity. and substrate selected result in redds constructed (1) where the females can dig effectively in the substrate: and (2) where the eggs deposited have adequate inter-gravel flow to maintain oxy 9 en supply during incubation. An additional parameter which can strongly influence site selection for redds is the occurrence of springs or other groundwater return flow. Ground water return flow is substantial at Vernita Bar, and is one of the reasons that this area is heavily utilized as spawning habitat.

'ne physical condition at Vernita Bar presents substrate that is suitable for spawning to an elevation equivalent to a flow level at Priest Rapids Dam of 85 kcfs. Under the terms of the current Vernita Bar

51 (A)

1983 SAR K/f 1 + [(K-yo)/y(] [j-bQ) O=FIOW, 0.4 where b=0.06908, K=.292, YO=O.(0183 RI 0.37 (P<0.05) Predicted SAR 1986 1985 1984

0.3

1979

0.2

78 1982 9 0.1

1981 19

0.0 ------20 40 60 80 100 120 140 160

1 8 1@6

10982 SAR K/(I+[(K-YO)[YO][e(-bO)p where Q=Flow, b=0.04086, K=1.36, YO=0.010201 1.4, R 2 0.26 (P

1.0

0.8

984 0.6

0.4 83

0.2 6

197 7 1979 0.0 1980

20 40 60 80 100 120 140 160

Flow (kcfs) at LoNver Granite Dam

Figure 20. Smolt-to-adult (SAR) return of spring Chinook and mean flow Lower 20 at Granite Dam (April - May 30) for (A) Rapid River Hatchery and (B) Marsh Creek releases. 1977-1987 migration years (Petrosky 1991).

52 Settlement Agreement (FERC-Docket No. E-9569-0,00), flows during spawning, October 15 to November 25, are regulated to discourage the utilization of spawning habitat above the 70 kcfs elevation. This is done to ease the constraints to the hydropower system during the incubation and emergence period, November

through May 15, when flow levels must be maintained to protect incubating eggs and emerged fry. Once is spawning completed, a redd location survey is conducted at Vernita Bar to set incubation and emergence flow levels.

flow The regulation during the spawning periods at Vernita Bar consists of attempting to keep flows during daylight hours (0700-1700) below the 70 kcfs level at Priest Rapids Dam. To accomplish this

objective, the daily average flow at Priest Rapids Dam must be less than or equal to 125 kcfs, unless Grant County Public Utility District is willing to spill water at night in excess of powerhouse capacity. The reduction of daylight flows has not been completely successful in discouraging spawning above the 70 kcfs

level at Vernita Bar. In the 1988 spawning season, 51 redds were located above this elevation in the index

area surveyed at Vernita Bar. This was apparently due to female chinook selecting sites and spawning at

night, when flows were substantially above the 70 kcfs level. Nighttime flows at Priest Rapids Dam were typically 150 kcfs at the peak of spawning, November 10-20, 1988.

'Me redd survey mentioned above sets a protection level which must be maintained by minimum flow releases from Priest Rapids Dam during the period November through completion of emergence, approximately May 15. This upper limit or cap on incubation/emergence protection is not providing 100 percent protection, as dramatically demonstrated in 1988.

The fall spawning period flow regulation appears to be fairly onerous to the hydropower operators, and not completely successful in terms of controlling the location of redds via reverse load factoring, i.e. relatively

low daytime and high nighttime flows. Our flow recommendation assumes 50 kcfs minimum flow will be in

place for November 1-30, a 70 kc[s minimum 1 15 and flow from December - May for incubation and

emergence protection, maximum flow restraint in the fall spawning period, agreement to continue conducting redd survevs after end of the spawning season, and the setting of annual incubation/emergence flow levels based upon actual placement of the highest redds on Vernita Bar.

Additionally, there are flow requirements in the fall for adult steeihead migrants entering the Snake River

is (Figure 21). Temperature an,important factor determining entry of these fish to the Snake River, but a analysis multiple regression performed on data for wild/natural "B' run steelhead shows that flows greater than 35 kcfs were the primary factor contributing to good conversion rates for these fish. These findings corroborate a previous report (Wagner 1971) which states "...there appears to be a correlation between Snake

River discharge and fish passage at Ice Harbor. The of flow be 20 amount necessary appears to ... between and 30 kcfs.'

4. 1 Winter (December - March 30)

Minimum instantaneous flows required for operation of adult fishways at the various Snake and Columbia

53 09 LI) C/)

It I, r

It rz a CL

uo ......

CL 0

CID River dams were developed under the auspices of the Columbia Basin Fishery Technical Committee, and were presented in the Wagner (1971) report, "Recommendations for Flow Regulation at Columbia and Snake

River Projects". For Bonneville Dam, the report states that the fish passage facilities will not operate when is the tailwater below elevation 8. Hourly flow data for Bonneville Dam indicate that flows of 80 kcfs can result in tailwater elevations of 7.7-8.2. Therefore, a minimum instantaneous flow of 80 kcfs is required for fish adult passage facility operation at Bonneville Dam. For minimum tailwater elevations at The Dalles, John Day, and McNary dams, the report recommends flows of 70, 65, and 63 kcfs. respectively. Since requirements at Bonneville override these requirements, our recommendation is 80 kcfs. For the Snake River projects, minimum design criteria for the adult fishways are the basis for minimum tailwater elevations. These tailwater elevation requirements result in minimum flow recommendations of 28 and 27 kcfs at the two lower and two upper projects, respectively. All mid-Columbia project adult fishways are expected to operate at the FERC minimum flow criteria of 36 kcfs. It should be pointed out that adult lish passage investigations have not been conducted at the mid-Columbia projects. C. Conclusions

Fish need flows that are guaranteed in the spring, summer, fall, and winter. As presented earlier in this report, the anadromous Fish of the Columbia River Basin adapted over thousands of years to the water flows and velocities present during the fishes spawning, rearing and migration periods. These important criteria for stages in the life history of anadromous fish do not allow the fish to cope with the altered flow and in velocitV patterns now present the system. Appropriate changes must be made to allow the needs of fish to be met. This requires specific flows established as hard constraints within the management of the

Columbia River hydrosystem. Establishment of a flow regime can address future power operations, can litigation, prevent controversy and can provide certainty for both fish and power, and can assure equitable treatment for fish and wildlife. This report is the technical recommendation of the Columbia Basin Fish and Wildlife Authority. V. CITED tEFERENCES

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