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1 PATRICK R. BERGIN (SBN 269672) FREDERICKS PEEBLES & MORGAN LLP 2 2020 L Street, Suite 250 3 Sacramento, CA 95811 Telephone: (916) 441-2700 4 Facsimile: (916) 441-2067 [email protected] 5 [Proposed] Amicus Curiae Modoc Tribe of 6 7 8 9 DISTRICT COURT 10 NORTHERN DISTRICT OF CALIFORNIA - SAN FRANCISCO DIVISION 11 THE , a federally Case No: 3:18-cv-03078-WHO 12 recognized Indian Tribe, [Related Case Nos. 3:16-cv-06863-WHO and 3:18-cv-03078-WHO] 13 Plaintiff, 14 vs. DECLARATION OF BLAKE FOLLIS IN SUPPORT OF AMICUS CURIAE 15 UNITED STATED BUREAU OF COUNTY OF SISKIYOU, COUNTY OF RECLAMATION; UNITED STATES FISH & MODOC, KLAMATH COUNTY, AND 16 WILDLIFE SERVICE; NATIONAL MARINE THE MODOC TRIBE IN SUPPORT OF FISHERIES SERVICE, DEFENDANTS UNITED STATES 17 BUREAU OF RECLAMATION ET AL. Defendants, AND DEFENDANT-INTERVENORS 18 KLAMATH WATER USERS and ASSOCIATION ET AL. 19 KLAMATH WATER USERS ASSOCIATION, 20 SUNNYSIDE IRRIGATION DISTRICT, and BEN DuVAL, 21 Defendant-Intervenors. 22 23 24 25 26 27 28

Case No. 3:18-cv-03078-WHO DECLARATION OF BLAKE FOLLIS 56509710 Case 3:18-cv-03078-WHO Document 50 Filed 07/03/18 Page 2 of 5

1 DECLARATION OF BLAKE FOLLIS 2 I, Blake Follis, declare as follows: 3 1. I am the Attorney General of the Modoc Tribe of Oklahoma (“Modoc Tribe”), a 4 federally recognized Indian Tribe headquartered in Miami, Oklahoma.1 I am duly licensed to 5 practice law in all courts of the Modoc Tribe and the State of Missouri. I am also an enrolled 6 member of the Modoc Tribe. Unless otherwise stated on information and belief, I have personal 7 knowledge of the facts stated herein and, if called as a witness, could testify competently thereto. 8 2. As Attorney General, I am familiar with the Modoc Tribe’s history, laws, 9 governmental and business operations, and treaties with the United States. I have reviewed the 10 Klamath Tribes’ Motion for Injunction, ECF No. 13, filed in the above-captioned lawsuit. I 11 submit this declaration in support of the joint amicus curiae brief of Siskiyou County, Modoc 12 County, Klamath County, and the Modoc Tribe. 13 3. The Modoc Tribe’s ancestral home is an area along the California-Oregon border, 14 extending from the summit of the Cascades near Happy Bend, CA to the summit of the Warner 15 Mountains in the Sierra Nevada mountain range and the lands between. As Euro-Americans 16 began settling the area in large numbers, the Modoc Tribe, along with the Klamath Tribe and the 17 Yahooskin Band of Snake Indians, entered a joint treaty with the United States in 1864, ceding 18 their lands and agreeing to live together on a reservation in present-day Oregon. See Treaty with 19 the Klamath Indians, 1864., Art. I, 16 Stat. 707. 20 4. Unable to live peaceably alongside the Klamath, the majority of the members of 21 the Modoc Tribe led by Captain Jack left the reservation in 1865 and later returned to the 22 reservation in December of 1869; departing again in April, 1870. The Modoc again returned to 23 their ancestral homelands, and requested that a separate Modoc reservation be established there. 24 The United States refused. The United States Army repeatedly tried to force the Modoc onto the 25 Klamath reservation, leading to an armed conflict initiated by orders from the Bureau of Indian 26

27 1 Also known as the Modoc Nation, Modoc Tribal Nation, Captain Jack’s Band of Modocs, 28 Modoc Indians in Oklahoma, and the Modoc Tribe of Oklahoma; to be distinguished from other groups and individuals promoting themselves as Modoc. - 1 - Case No. 3:18-cv-03078-WHO DECLARATION OF BLAKE FOLLIS 56509710 Case 3:18-cv-03078-WHO Document 50 Filed 07/03/18 Page 3 of 5

1 Affairs that was known as the Battle of and the beginning of the , which 2 was fought in 1872-73, in and around the lava beds south of Tule Lake, California. 3 5. At the end of the Modoc War, the United States executed Captain Jack and three 4 other Modocs, and sentenced two others to life imprisonment at Alcatraz. The United States 5 removed the belligerent Modocs to , taking 153 individuals east by railroad under 6 armed guard, originally to , Wyoming, then to Fort McPherson, Nebraska, down 7 through Kansas City, Missouri and unloading the Modocs in Baxter Springs, Kansas and 8 ultimately settling them at a location in the Agency, in the northeastern corner of 9 present-day Oklahoma. 10 6. The Act of March 3, 1909, placed the removed Modoc to be enrolled with the 11 Klamath Agency in Oregon. In 1954, the United States terminated federal supervision of “the 12 Klamath Tribe of Indians consisting of the Klamath and Modoc Tribes and Yahooskin Band of 13 Snake Indians.” 14 7. In 1978, Congress enacted a law recognizing the “Modoc Indian Tribe of 15 Oklahoma.” The law stated that the recognized tribe “shall consist of those Modoc Indians who 16 are direct lineal descendants of those Modocs removed to Indian territory (now Oklahoma) in 17 November 1873, and who did not return to Klamath, Oregon, pursuant to the Act of March 9, 18 1909 (35 Stat. 751), as determined by the Secretary of the Interior, and the descendants of such 19 Indians who otherwise meet the membership requirements adopted by the Tribe.” The law 20 repealed the application of the 1954 Klamath Termination Act to the Modocs in Oklahoma. The 21 Modoc Tribe remains an unaffiliated independent government from the Klamath Tribes. 22 8. The Modoc Tribe owns 800 acres in the County of Siskiyou and is in the process 23 of reacquiring additional land within its aboriginal territory as part of a governmental effort to 24 return home. The Modoc Tribe also possesses “the exclusive right of taking fish in the streams 25 and lakes” and “gathering edible roots, seeds, and berries” from areas now irrigated by the 26 Project. Treaty with the Klamath Indians, 1864., Art. I, 16 Stat. 707. See also, United States v. 27 Adair, 478 F. Supp. 336 (D. Or. 1979), aff'd as modified, 723 F.2d 1394 (9th Cir. 1983) (Treaty

28 signers reserved water rights associated with fishing and gathering have not been abrogated.).

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1 Even in exile, the Modoc Tribe members continue to possess these reserved rights. See Kimball 2 v. Callahan, 493 F.2d 564, 568 (9th Cir. 1974) (Treaty rights to hunt and fish are, however, 3 rights of the individual Indians.). Thus, the Modoc Tribe has governmental, jurisdictional, and 4 economic interests in the California Counties of Modoc and Siskiyou, as well as in the Oregon 5 Counties of Lake, Jackson and Klamath. 6 9. The land ceded by the Modoc Tribe under the 1864 treaty includes the area now 7 irrigated by the Klamath Irrigation Project. Water from the Lost River enters the Klamath River 8 basin from natural flows from Clear Lake and man-made flows from the Upper Klamath Lake, 9 which is the water source at issue in this lawsuit. The headwaters of the Klamath River are 10 sourced from the Upper Klamath Lake. Today, Tule Lake and Lost River depend on the 11 irrigation flows and diversions of the Klamath Irrigation Project to sustain water levels in the 12 Lost River and Tule Lake. The Lost River, along with its terminus, Tule Lake, serves as a main 13 water source for the Modoc Tribe’s aboriginal territory. It is among these water sources that the 14 Modoc Tribe historically relied upon on for subsistence. Accordingly, the Modoc Tribe has an 15 interest in preserving water flows to its aboriginal territory in the area irrigated by the Klamath 16 Irrigation Project. 17 10. The Modoc Tribe is acquiring land in California for the benefit of the Modoc 18 people and the natural resources under its jurisdiction. Among the needs of the Modoc Tribe is a 19 sustainable supply of water within the Klamath Irrigation Project area. In addition, the Modoc 20 Tribe intends to bring much needed economic development to this area, such as agricultural 21 products and services, which rely on ample water supplies. The inability to receive irrigable 22 water, as would occur if the Tribes’ injunction is granted, will irreparably harm the governmental 23 interests of the Modoc Tribe. Moreover, increased retention of water in the Upper Klamath 24 Lake, as would be needed to satisfy the Tribes requested lake levels, will ensure that essentially 25 no water is made available downstream to the Modoc Tribe and its neighbors, including Klamath 26 Project irrigators. 27 11. It is among my duties as Attorney General to monitor litigation, as well as local,

28 state and federal administrative actions concerning the Modoc Tribe’s ancestral lands. In doing

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1 so, I have become aware of relevant informational material on the Lost River sucker fish 2 published by the California Department of Fish & Wildlife and the United States Fish & Wildlife 3 Service. Attached hereto as Exhibit A is a true and correct copy of the informational page on 4 the Lost River Sucker published by the Endangered Species Project of the California Department 5 of Fish & Wildlife. Attached hereto as Exhibit B is a true and correct copy of the informational 6 page on the Lost River Sucker by the United States Fish & Wildlife Service and published by the 7 Oregon Fish and Wildlife Office. 8 I declare under penalty of perjury under the laws of the United States of America that the 9 foregoing is true and correct. 10 Executed July 3, 2018, at Miami, Oklahoma. 11 12 13

14 Blake Follis 15 16 17 18 19 20 21 22 23 24 25 26 27 28

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Lost River Sucker (Deltistes luxatus) Lost River Sucker and Shortnose Sucker Status -- Federal: Endangered; California: Endangered reclamation projects devel- Shortnose Sucker (Chasmistes brevirostris) oped in the early 1900s, Status -- Federal: Endangered; California: Endangered resulted in the loss of over The Lost River sucker is a large fish, measuring up to 3 feet (1m) in total length, 250,000 acres of wetlands in and weighing up to 10 lbs (4.5 kg). Its small mouth is positioned below its elon- the Upper Klamath Basin. gated snout. Its coloration varies with location, Upper Klamath Lake (Oregon) The loss of these wetlands fish have dark backs and sides has had large scale impacts to fading to yellow or white on the the quality and quantity of belly, while Clear Lake Reservoir suitable sucker habitat. (California) fish are light brown Currently, less than 75,000 above and white or tan below. acres of wetlands remain in Capable of living for more than the Basin. 40 years, Lost River suckers usu- Range: Today, substantial ally inhabit deep lakes or river populations of Lost River Photos: John Brode, CDFG Lost River Sucker Current Range suckers are only found in pools. Adult suckers are thought Current and Historic Range to feed near the bottom on invertebrates found in the sediments and zooplankton. Tule Lake, part of the Lost The Shortnose sucker is smaller than the Lost River sucker, measuring less than 20 River, and Lower Klamath Lake in Siskiyou County, and in Clear Lake Reservoir inches (50 cm) in length, and weighs 3.5 lbs (1.5 kg) when fully grown. It has a big in Modoc County. In California, the present distribution of Shortnose suckers is head and its body is nearly cylindrical. It is recognized by its large, flexible mouth. similar to that of the Lost River sucker. Gerber Reservoir in Oregon is the only The fish is also characterized by its blunt, turned-up snouth. Body coloring ranges habitat with a Shortnose sucker population that does not also have a Lost River from dark-to-light brown above and sucker population. from tan to white below. They are Breeding: Lost River suckers reach sexual maturity between the ages 6 to 14 known to live at least 33 years. For years. From early February through May, they begin their runs up tributary most of the year, they live in large, streams in order to spawn. Females release their eggs in stretches of stream that shallow lakes and sluggish rivers, flow swiftly over rubble bottoms, depositing 44,000 - 231,000 eggs each. After where they feed on zoo-plankton, hatching, larvae move downstream to the lake under cover of darkness. Shortnose suckers reach sexual maturity at age 6 or 7. They begin their runs in algae, and benthic (sediments) Shortnose Sucker invertebrates. March, migrating up tributary rivers to spawn. In stretches of riffles and smooth Both sucker species shared a similar historical distribution, in Upper Klamath Lake runs of water, over gravel -or rubble-covered stream bottoms, females broadcast and its tributaries, and the Lost River system. Lost River suckers also occupied the tens of thousands of eggs. Some suckers in both species spawn along the shores of waters of Tule Lake, Lower Klamath Lake, and Sheepy Lake. Large scale water lakes and springs. Endangerment: The combined effects of damming of rivers, instream flow California Department of Pesticide Regulation diversions, draining of marshes, dredging of Upper Klamath Lake, and other water manipulations have threatened both species with extinction. Additionally, water Endangered Species Project quality degradation in the Klamath Basin watershed has led to large-scale fish kills California Department of Fish & Game related to algal bloom cycles. Introduced exotic fishes may reduce recruitment www.cdpr.ca.gov through competition with, or predation upon, suckers and sucker larvae. EXHIBIT A Case 3:18-cv-03078-WHO Document 50-2 Filed 07/03/18 Page 1 of 2

6/29/2018 OFWO - Lost River sucker FOLLOW THE FWS ONLINE

Oregon Fish and Wildlife Office Working with you to conserve the natural resources of Oregon

Lost River sucker

Scientific name: Deltistes luxatus

Status: Endangered

Critical Habitat: Proposed

Listing: The Lost River sucker was federally listed as endangered in 1988. A recovery plan was published in 1993. Critical habitat was proposed in 1994, but not designated. A status review was conducted in 2004, and a five-year review was done in 2007.

Historical Status and Current Trends Early records indicate that Lost River suckers were once widespread and abundant in the upper Klamath Basin of Oregon and California. This area historically contained over 350,000 acres of wetlands and floodplains. These wetlands protected sucker habitat by controlling erosion, recycling organic and inorganic nutrients, and maintaining water quality. Because suckers were historically very abundant, they were a major food source for Native Americans and local settlers in the late 1800s. Canneries were established along the Lost River to process suckers into oil, dried fish, and other products. However, agricultural development and associated water and land use changes in the basin have contributed to the significant loss of wetland habitat and a significant decline in sucker populations. Although overharvesting and pollution may have played a role in the species decline, it is believed that the combined effects of the construction of dams, the draining or dredging of lakes, and other alterations of natural stream flow have reduced the reproductive success of Lost River suckers by as much as 95 percent through the degradation of suitable breeding habitat. At the time the Lost River sucker was listed as endangered, it was noted that there had been no significant addition of young into the population in 18 years.

Currently, the Lost River sucker occupies only a fraction of its former range and is restricted to a few areas in the Upper Klamath Basin, such as the drainages of Upper Klamath Lake, Tule Lake, and Clear Lake. Poor water quality, reduced suitable habitat for all sizes and ages, and the impacts of non-native fishes continue to threaten remaining Lost River sucker populations.

Description and Life History Locally known as mullet, the Lost River sucker is a large, long-lived sucker that can reach 43 years of age. It has unique triangular-shaped gill structures which are used to strain a diet of detritus (decomposing organic matter), zooplankton (tiny floating aquatic animals), algae, and aquatic insects from the water. Lost River suckers typically begin to reproduce at nine years, when they first participate in spawning migration. Adult suckers migrate from the quiet waters of lakes into fast moving streams from March through May in order to spawn. They may also spawn in lakeshore springs from February to mid-April when the water temperature is a constant 15 C (60 F). Thousands of eggs (from 44,000 for smaller fish to 218,000 for larger suckers) are typically laid near the stream bottom in areas where gravel or cobble is available. Once the eggs hatch, the

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larval6/29/2018 fish begin their migration back to calmer waters. They generally OFWOmigrate - Lostat night River and sucker stay in shallow, shoreline areas and in aquatic vegetation during the day. Upon their return to the lake, larvae may be preyed upon by largemouth bass, yellow perch, or other non-native predatory fish, and larger juveniles may compete for food with non-native fishes such as fathead minnows, yellow perch, and others.

Habitat The Lost River sucker dwells in the deeper water of lakes and spawns in springs or tributary streams upstream of the home lake. Areas with gravel or close-set stone ("cobble") bottoms at springs or in moderate to fast-flowing springs are preferred for spawning. In addition, the spawning streams have a fairly shallow shoreline with abundant aquatic vegetation; these areas provide a safe haven for the young larvae during their journey back downstream to their home lakes or the deep, quiet waters of rivers.

Reasons for Decline Although a number of factors have contributed to the decline of the Lost River sucker, habitat degradation is considered the primary cause. Streams, rivers, and lakes have been modified by channelization and dams. Grazing in the riparian zone has eliminated streambank vegetation, and has added nutrients and sediment to river systems. Eggs and larvae, for example, suffocate when the water is cloudy, or dry out or get eaten by other fish when they are not protected by aquatic vegetation. Loss of streambank vegetation due to overgrazing, logging activities, agricultural practices, and road construction has also led to increases in stream temperatures, high levels of nutrients (which encourages the buildup of excess algae and bacteria), and serious erosion and sedimentation problems in streams. Such water quality problems have reduced the availability of suitable Lost River sucker habitat and have resulted in major fish mortality. Entire age classes of young suckers are routinely lost due to poor water quality conditions. As a result, few young suckers survive to sexual maturity, and therefore, do not increase the population size. Other factors affecting the decline of the Lost River sucker include previous overharvesting, chemical pollution from pesticides, herbicides, and forestry practices, and predation and competition from native and non-native fishes such as largemouth bass, blue chub, yellow perch, fathead minnows, and rainbow trout.

Conservation Measures Conservation efforts for the Lost River sucker focus on the re-establishment of a more naturally functioning ecosystem in the Klamath Basin. Fencing portions of streams to reduce cattle-caused erosion, replanting streambanks with native vegetation, improving forestry and agricultural practices, and assuring adequate water levels in reservoirs will contribute to the recovery of this species. Through coordination of the actions of land use agencies and private landowners, further degradation of sucker habitat can be avoided and steps can be taken to improve current conditions. By minimizing the impacts of future modifications to spawning habitat and restoring waters to a more natural state, recovery of Lost River sucker populations is possible in the Klamath Basin.

References and Links U.S. Fish and Wildlife Service. 1988. Determination of Endangered Status for the Shortnose Sucker and Lost River Sucker. FR 53:27130-27134. (https://ecos.fws.gov/docs/frdocs/1988/88-16062.pdf)

U.S. Fish and Wildlife Service. 1993. Shortnose Sucker (Chasinistes brevirostris) and Lost River (Deltistes luxatus) Sucker Recovery Plan (http://ecos.fws.gov/docs/recovery_plans/1993/930317.pdf). Portland, Oregon 108pp.

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1 NOSSAMAN LLP PAUL S. WEILAND (SBN 237058) 2 [email protected] 3 ASHLEY J. REMILLARD (SBN 252374) [email protected] 4 18101 Von Karman Avenue, Suite 1800 Irvine, CA 92612 5 Telephone: 949.833.7800 Facsimile: 949.833.7878 6 7 [Proposed] Amicus Curiae Siskiyou County 8 9 10 UNITED STATES DISTRICT COURT 11 12 NORTHERN DISTRICT OF CALIFORNIA - SAN FRANCISCO DIVISION 13 THE KLAMATH TRIBES, a federally Case No: 3:18-cv-03078-WHO recognized Indian Tribe, [Related Case Nos. 3:16-cv-06863-WHO and 14 3:18-cv-03078-WHO] Plaintiff, 15 vs. DECLARATION OF RICHARD A. VALDEZ IN SUPPORT OF AMICUS 16 UNITED STATED BUREAU OF CURIAE COUNTY OF SISKIYOU, COUNTY OF MODOC, KLAMATH 17 RECLAMATION; UNITED STATES FISH & WILDLIFE SERVICE; NATIONAL MARINE COUNTY, AND THE MODOC TRIBE IN FISHERIES SERVICE, SUPPORT OF DEFENDANTS UNITED 18 STATES BUREAU OF RECLAMATION Defendants, ET AL. AND DEFENDANT- 19 INTERVENORS KLAMATH WATER USERS ASSOCIATION ET AL. 20 and 21 KLAMATH WATER USERS ASSOCIATION, SUNNYSIDE IRRIGATION DISTRICT, and 22 BEN DuVAL, 23 Defendant-Intervenors. 24 25 26 27 28

Case No. 3:18-cv-03078-WHO DECLARATION OF RICHARD A. VALDEZ 56499896 Case 3:18-cv-03078-WHO Document 50-3 Filed 07/03/18 Page 2 of 5

1 DECLARATION OF RICHARD A. VALDEZ 2 I, Richard A. Valdez, declare as follows: 3 1. I am a certified fisheries biologist with an undergraduate degree in wildlife 4 management from New Mexico State University, Las Cruces, and a master’s degree in fisheries, 5 a doctoral degree in fisheries ecology, and a post-doctoral degree in stream ecology from Utah 6 State University, Logan. I am currently employed as a Senior Scientist with SWCA 7 Environmental Consultants, a position that I have held since 1996. I have 40 years of experience 8 in streams and lakes of western North America. I have expertise in fish population dynamics, 9 abundance estimates, stock assessment and recruitment models, population viability analysis, 10 and design and implementation of monitoring. A copy of my curriculum vitae is attached hereto 11 as Exhibit 1. I have personal knowledge of the facts stated herein and, if called as a witness, 12 could testify competently thereto. 13 2. I am familiar with the scientific literature published regarding Lost River sucker 14 (Deltistes luxatus) and shortnose sucker (Chasmistes brevirostris) and their habitat, including the 15 Upper Klamath Lake. This literature includes: (1) National Academy of Sciences, Scientific 16 Evaluation of Biological Opinions on Endangered and Threatened Fishes in the Klamath River 17 Basin: Interim Report (2002) (“2002 Report”); (2) National Academy of Sciences, Endangered 18 and Threatened Fishes in the Klamath River Basin: Causes of Decline and Strategies for 19 Recovery (2004) (“2004 Report”);1 (3) National Academy of Sciences, Hydrology, Ecology, and 20 Fishes of the Klamath River Basin (2008) (“2008 Report”),2 (4) Martin & Saiki, Effects of 21 Ambient Water Quality on the Endangered Lost River Sucker in Upper Klamath Lake, Oregon, 22 Transactions of the American Fisheries Society 128:953–961 (1999) (“Martin & Saiki”); and (5) 23 Saiki et al., Lethal levels of selected water quality variables to larval and juvenile Lost River and 24 shortnose suckers, Environmental Pollution 105 (1999) 37-44 (“Saiki et al.”). True and correct 25 26

27 1 Available at: http://www.westcoast.fisheries.noaa.gov/publications/Klamath/nrc_2004.pdf. 2 28 Available at: http://www.westcoast.fisheries.noaa.gov/publications/Klamath/nrc_2008_hydrol ogy_ecology_and_fishes_of_the_klamath.pdf. - 1 - Case No. 3:18-cv-03078-WHO DECLARATION OF RICHARD A. VALDEZ 56499896 Case 3:18-cv-03078-WHO Document 50-3 Filed 07/03/18 Page 3 of 5

1 copies of the 2002 Report, Martin & Saiki, and Saiki et al. are attached hereto as Exhibit 2, 2 Exhibit 3, and Exhibit 4, respectively. 3 3. I have reviewed the Declaration of Mark Buettner in Support of the Klamath 4 Tribes’ Motion for Injunction, ECF No. 13-3, filed in the above-captioned lawsuit. I submit this 5 declaration in support of Siskiyou County’s amicus brief. 6 4. Populations of Lost River sucker and shortnose sucker in Upper Klamath Lake 7 have declined substantially over the past half century. This decline is likely a consequence of 8 diminishing recruitment over time. However, the cause(s) of such diminishing recruitment is not 9 well understood. 10 5. The hypothesis underlying the Klamath Tribes’ position—that Upper Klamath 11 Lake levels are the cause of poor sucker recruitment—originated in the early 2000s. This has led 12 the U.S. Bureau of Reclamation and the U.S. Fish and Wildlife Service to increase lake levels 13 since that time. Most recently, the 2013 Klamath Project Biological Opinion, which was 14 developed through consultation pursuant to the Endangered Species Act, sets forth minimum 15 threshold elevations in Upper Klamath Lake to protect the imperiled species. 16 6. Despite the fact that Upper Klamath Lake levels have been maintained above 17 prior levels for over a decade, including most recently pursuant to the 2013 Klamath Project 18 Biological Opinion, there is no evidence of improving recruitment. This suggests that simply 19 altering lake levels will not result in improved sucker population levels. 20 7. The Klamath Tribes assert that higher lake levels—namely those identified as 21 “2018 Transition Levels” and “C’waam and Koptu Conservation Levels” in their briefing—are 22 sufficient to cause the population levels of Lost River sucker and shortnose sucker to improve. 23 There is no empirical scientific evidence to support this assertion. Put another way, even if 24 Upper Klamath Lake levels are increased to the levels requested by the Klamath Tribes, it is my 25 expert opinion that this alone will not improve sucker recruitment such that it will increase their 26 population and insure their viability. Rather, other variables in Upper Klamath Lake (e.g., 27 predation of larvae by fathead minnows; algae blooms causing oxygen depletions, etc.) all 28

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1 impact sucker recruitment. As such, merely increasing lake elevations in Upper Klamath Lake 2 will not improve sucker populations. 3 8. This conclusion—that higher lake levels do not necessarily improve sucker 4 populations—was confirmed by two blue-ribbon science panels established by the National 5 Research Council, a branch of the National Academies. The panels produced peer-reviewed 6 reports entitled Endangered and Threatened Fishes in the Klamath River Basin: Causes of 7 Decline and Strategies for Recovery in 2004 (the 2004 Report) and Scientific Evaluation of 8 Biological Opinions on Endangered and Threatened Fishes in the Klamath River Basin in 2002 9 (the 2002 Report). In short, the National Academy science panels concluded that there is no 10 scientifically justifiable connection between lake levels and improved sucker population levels. 11 See, e.g., 2004 Report at 226-227; 2002 Report at 18-19. 12 9. In the 2004 Report, the expert panel addressed the same two aspects of sucker 13 survival now put forth by the Klamath Tribes: physical habitat and water quality. With respect 14 to physical habitat, the expert panel concluded, with respect to both sucker species, that: “For 15 the present there is no indication of a strong relationship between spawning success, as inferred 16 from abundance of larvae, and water level in Upper Klamath Lake." 2004 Report at 226. The 17 panel continued, stating: “Overall, maintaining full pool elevation for promotion of spawning, 18 although intuitively appealing, is difficult to defend scientifically.” Id. at 227. 19 10. Regarding water quality, the panel stated that “monitoring data show no 20 relationship between pH and water level,” concluding that “the weight of current evidence does 21 not support the argument that higher lake levels will mitigate problems associated with high pH.” 22 2004 Report at 115-116. 23 11. Other literature, including publications by Martin & Saiki and Saiki et al., confirm 24 that fish mortality in the Upper Klamath Lake is not directly attributable to water quality 25 concerns. See, e.g., Martin & Saiki, Effects of Ambient Water Quality on the Endangered Lost 26 River Sucker in Upper Klamath Lake, Oregon at 959 (“Collectively, these findings indicate that 27 water temperatures and un-ionized ammonia concentrations in Upper Klamath Lake were not

28 directly responsible for the occasional episodes of high fish mortality…”).

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1 12. The conclusions in the 2002 Report and 2004 Report were endorsed by another 2 National Academy panel in the 2008 Report, which stated: “The [2002 Report] concluded that 3 … available scientific data did not support the higher minimum lake levels or the higher 4 minimum river flows recommended in the biological opinions to benefit the species listed under 5 the Endangered Species Act. The [2004 Report] confirmed those conclusions…. This 6 committee endorses the recommendations of the earlier reports ….” 7 13. The specific higher lake levels requested by the Klamath Tribes, and advanced by 8 Mr. Buettner, are based on correlation and personal judgment, rather than empirical research and 9 causation. 10 14. There is an absence of empirical research suggesting that the lake levels requested 11 by the Klamath Tribes, as compared to those already maintained pursuant to the 2013 Klamath 12 Project Biological Opinion, will contribute to improved recruitment and therefore conservation 13 of the suckers. 14 I declare under penalty of perjury under the laws of the United States of America that the 15 foregoing is true and correct. 16

17 Executed June 27, 2018, at Logan, Utah. 18 19 20 21 22 23 24 25 26 27 28

- 4 - Case No. 3:18-cv-03078-WHO DECLARATION OF RICHARD A. VALDEZ 56499896 Case 3:18-cv-03078-WHO Document 50-4 Filed 07/03/18 Page 1 of 2 RICHARD A. VALDEZ, Ph.D., CFS Senior Scientist 6/1/2018

Education / Training Relevant Projects • Post-Doctorate, Stream Ecology, Utah • Member, Brown Trout Management Team, Glen State Univ., Logan, 1977 Canyon Dam and Grand Canyon Adaptive • Ph.D., Fisheries Ecology, Utah State Management Control Options, U.S. Geological Univ., Logan, 1975 Survey, U.S.G.S. Circular Publication (2017-present). • M.S., Fisheries, Utah State Univ., Logan, 1971 • Science Panel Chair, Feasibility of Reintroducing the • B.S., Wildlife Management, New Endangered Razorback Sucker into Lower Grand Mexico State Univ., Las Cruces, 1968 Canyon, U.S. Bureau of Reclamation (2010-2012, 2017).

Registration / Certification • Subject Matter Expert, Razorback Sucker Species • Certified Fisheries Scientist Status Assessment, U.S. Fish and Wildlife Service, (CFS#2088), American Fisheries Denver, CO (2016-present). Society, 1993 • Subject Matter Expert, Rio Grande Silvery Minnow Science Panel, U.S. Army Corps of Engineers, Professional Positions Albuquerque, NM (2016-present). • SWCA Environmental Consultants, • Logan, UT (1996-present), Senior Leader, Humpback Chub Recovery Team, U.S. Fish Scientist and Wildlife Service, Denver, CO (2015-present). • Bio/West, Inc., Logan, UT (1986-1996), • Senior Science Advisor, New Mexico Interstate Fisheries Section Manager Stream Commission (NMISC), Albuquerque, NM • Ecosystems Research Institute, Logan, (1995-present). UT (1984-1986), Senior Aquatic • Ecologist Science Panel Chair, Glen Canyon Dam Long-Term Experimental and Management Plan. Colorado River • U.S. Fish and Wildlife Service, Reno, NV Basin States and Upper Colorado River Commission (1983-1984), Endangered Species (2012-2016). Coordinator • U.S. Fish and Wildlife Service, • Technical Fisheries Advisor, In the Matter of the Yellowstone National Park, WY (1981- Determination of the Relative Rights of the Waters of 1983), Assistant Project Leader the Klamath River, a Tributary of the Pacific Ocean, • U.S. Fish and Wildlife Service, Grand Case 282 and 286 (2007-2008). Junction, CO (1979-1981), Field • Advisory Team Member, Temperature Control Device Station Coordinator for Glen Canyon Dam. Advise Bureau of Reclamation and Grand Canyon Program on Expertise construction of a TCD (1999-2002). • 40 years experience in streams and • lakes of western North America Workshop Facilitator: • Fish population dynamics, • Population Estimates Workshops to monitor abundance estimators, stock endangered fishes of the Upper Colorado River assessment and recruitment models, Basin (2002), and population viability analysis, design and implementation of monitoring • Nonnative Fish Control Workshops for the Upper Colorado River Basin (2006-2008). • Native and endangered fish issues, including Endangered Species Act, • Project Manager, Grand Canyon Humpback Chub Sections 4, 7, 9, 10 Life History Studies, U.S. Bureau of Reclamation, • Fish passage and stream restoration Flagstaff, AZ (1989-1998). • Water supply, management, • Project Manager, Cataract Canyon Fisheries Studies, watershed assessment, hydrology, U.S. Bureau of Reclamation, Salt Lake City, UT (1985- and instream flow assessment, 1990). including TMDL, and NPDES

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RICHARD A. VALDEZ, Ph.D., CFS Senior Scientist 6/1/2018 Key Reports / Publications • Runge, M.C., Yackulic, C.B., Bair, L.S., Kennedy, T.A., Valdez, R.A., Ellsworth, C., Kershner J.L., Rogers, R.S., Trammell, M.A., and Young, K.L., 2018, Brown trout in the Lees Ferry reach of the Colorado River—Evaluation of causal hypotheses and potential interventions: U.S. Geological Survey Open-File Report 2018–1069, 83 p., https://doi.org/10.3133/ofr20181069. • U.S. Fish and Wildlife Service. 2017. Humpback Chub Species Status Assessment. U.S. Fish and Wildlife Service, Mountain-Prairie Region 6, Denver, CO. (Recovery Team Leader) • U.S. Fish and Wildlife Service. 2016. Colorado Pikeminnow (Ptychocheilus lucius) population viability analysis. U.S. Fish and Wildlife Service, Mountain-Prairie Region 6, Denver, CO. (assist PVA modeler) • U.S. Fish and Wildlife Service. 2015. Colorado Pikeminnow (Ptychocheilus lucius) draft recovery plan. U.S. Fish and Wildlife Service, Mountain-Prairie Region 6, Denver, CO. (Head of Writing Team) • Valdez, R.A. (Science Panel Chair), et al. 2012. Proposed alternative for the Long-Term Experimental and Management Plan for flow and non-flow actions related to the operation of Glen Canyon Dam. Colorado River Basin States and Upper Colorado River Commission. • Valdez, R.A. (Science Panel Chair), et al. 2012. The potential of habitat for the razorback sucker in the lower Grand Canyon and Colorado River inflow to Lake Mead: A Science Panel Report, Report Number 2. U.S. Bureau of Reclamation, Upper Colorado Region, Salt Lake City, UT. • Kehmeier, J.W., R.A. Valdez, C.N. Medley, and O.B. Meyers. 2007. Relationship of fish mesohabitat to flow in a sand-bed southwestern river. North American Journal of Fisheries Management 27:750-764. • Medley, C.N., Kehmeier, J.W., O.B. Meyers, and R.A. Valdez. 2007. Simulated transport and retention of pelagic fish eggs during an irrigation release in the Pecos River, New Mexico. Journal of Freshwater Ecology 22:499-513. • Valdez, R.A. (Chairman), et al. 2006. Study Plan for implementation and evaluation of flow and temperature recommendations for endangered fishes in the Green River downstream of Flaming Gorge Dam. U.S. Fish and Wildlife Service, Denver, CO. • Valdez, R.A. and R.T. Muth. 2005. Ecology and conservation of native fishes in the Upper Colorado River Basin, in J.N. Rinne, R.M. Hughes, and B. Calamusso (eds). Historical changes in large river fish assemblages of the Americas. American Fisheries Society, Bethesda, MD. • Valdez, R.A. and P. Nelson. 2005. Upper Colorado River Floodplain Management Plan. Upper Colorado River Endangered Fish Recovery Program, Project No. C-6, Denver, CO. • Valdez, R.A., T.L. Hoffnagle, W.C. Leibfried, C.C. McIvor, and T. McKinney. 2001. Effects of a test flood on fishes of the Colorado River in Grand Canyon, Arizona. Ecological Applications 11(3):686-700. • Webb, R.H., J.C. Schmidt, G.R. Marzolf, and R.A. Valdez (eds.). 1999. The Controlled Flood in Grand Canyon. Geophysical Monograph 110, American Geophysical Union, Washington, D.C. • Marzolf, G.R., R.A. Valdez, J.C. Schmidt, and R.G. Webb. 1998. Perspective on river restoration in the Grand Canyon. Bulletin of the Ecological Society of America 79(4):250-254. • Schmidt, J.C., R.H. Webb, Valdez, R.A. Valdez, G.R. Marzolf, and L.E. Stevens. 1998. Science and values in river restoration in the Grand Canyon. BioScience 48(9):735-747. • Converse, Y.K., C.P. Hawkins, and R.A. Valdez. 1998. Habitat relationships of subadult humpback chub in the Colorado River through Grand Canyon: Spatial variability and implications of flow regulation. Regulated Rivers 14:267-284. • Valdez, R.A. and R.J. Ryel. 1997. Life history and ecology of the humpback chub in the Colorado River in Grand Canyon, Arizona, in Proceedings of the Third Biennial Conference of Research on the Colorado Plateau, edited by C. van Riper III and E. T. Deshler. National Park Service, Natural Resources Publication Office.

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Scientific Evaluation of Biological Opinions on Endangered and Threatened Fishes in the Klamath River Basin: Interim Report

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SCIENTIFIC EVALUATION OF BIOLOGICAL OPINIONS ON ENDANGERED AND THREATENED FISHES IN THE KLAMATH RIVER BASIN

INTERIM REPORT

Committee on Endangered and Threatened Fishes in the Klamath River Basin Board on Environmental Studies and Toxicology Division on Earth and Life Studies National Research Council

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NATIONAL ACADEMY PRESS 2101 Constitution Ave., N.W.Washington, D.C. 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This project was supported by Grant No. 98210–1–G092 between the National Academy of Sciences and the U.S. Department of the Interior and the U.S. Department of Commerce. Any opin- ions, findings, conclusions, or recommendations expressed in this publication are those of the author (s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. International Standard Book Number: 0–309–08324–9 Additional copies of this report are available from: National Academy Press 2101 Constitution Ave., NW Box 285 Washington, DC20055 800–624–6242 202–334–3313(in the Washington metropolitan area) http://www.nap.edu Copyright 2002 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce M. Alberts is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Kenneth I. Shine is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy’s purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce M. Alberts and Dr. Wm. A. Wulf are chairman and vice chairman, respectively, of the National Research Council. www.national-academies.org

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COMMITTEE ON ENDANGERED AND THREATENED FISHES IN THE KLAMATH RIVER BASIN Members

WILLIAM M.LEWIS, JR. (Chair), University of Colorado, Boulder RICHARD M.ADAMS, Oregon State University, Corvallis ELLIS B.COWLING, North Carolina State University, Raleigh EUGENE S.HELFMAN, University of Georgia, Athens CHARLES D.D.HOWARD, Consulting Engineer, Victoria, British Columbia, Canada ROBERT J.HUGGETT, Michigan State University, East Lansing NANCY E.LANGSTON, University of Wisconsin, Madison JEFFREY F.MOUNT, University of California, Davis PETER B.MOYLE, University of California, Davis TAMMY J.NEWCOMB, Virginia Polytechnic Institute and State University, Blacksburg MICHAEL L.PACE, Institute for Ecosystem Studies, Millbrook, New York J.B.RUHL, Florida State University, Tallahassee Staff

SUZANNE VAN DRUNICK, Project Director RUTH E.CROSSGROVE, Editor JENNIFER SAUNDERS, Research Assistant MIRSADA KARALIC-LONCAREVIC, Research Assistant HEATHER A.MCDONALD, Project Assistant KELLY CLARK, Editorial Assistant Sponsors

NATIONAL MARINE FISHERIES SERVICE U.S.BUREAU OF RECLAMATION U.S.FISH AND WILDLIFE SERVICE

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BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY1

Members

GORDON ORIANS (Chair), University of Washington, Seattle JOHN DOULL (Vice Chair), University of Kansas Medical Center, Kansas City DAVID ALLEN, University of Texas, Austin INGRID C.BURKE, Colorado State University, Fort Collins THOMAS BURKE, Johns Hopkins University, Baltimore, Maryland WILLIAM L.CHAMEIDES, Georgia Institute of Technology, Atlanta CHRISTOPHER B.FIELD, Carnegie Institute of Washington, Stanford, California J.PAUL GlLMAN, Celera Genomics, Rockville, Maryland DANIEL S.GREENBAUM, Health Effects Institute, Cambridge, Massachusetts BRUCE D.HAMMOCK, University of California, Davis ROGENE HENDERSON, Lovelace Respiratory Research Institute, Albuquerque, New Mexico CAROL HENRY, American Chemistry Council, Arlington, Virginia ROBERT HUGGETT, Michigan State University, East Lansing JAMES H.JOHNSON, Howard University, Washington, District of Columbia JAMES F.KITCHELL, University of Wisconsin, Madison DANIEL KREWSKI, University of Ottawa, Ottawa, Ontario JAMES A.MACMAHON, Utah State University, Logan WlLLEM F.PASSCHIER, Health Council of the Netherlands, The Hague, Netherlands ANN POWERS, Pace University School of Law, White Plains, New York LOUISE M.RYAN, Harvard University, Boston, Massachusetts KIRK SMITH, University of California, Berkeley LISA SPEER, Natural Resources Defense Council, New York, New York Senior Staff

JAMES J.REISA, Director DAVID J.POLICANSKY, Associate Director and Senior Program Director for Applied Ecology RAYMOND A.WASSEL, Senior Program Director for Environmental Sciences and Engineering KULBIR BAKSHI, Program Director for Committee on Toxicology ROBERTA M.WEDGE, Program Director for Risk Analysis K.JOHN HOLMES, Senior Staff Officer SUZANNE VAN DRUNICK, Senior Staff Officer RUTH E.CROSSGROVE, Managing Editor

1This study was planned, overseen, and supported by the Board on Environmental Studies and Toxicology. Copyright National Academy of Sciences. All rights reserved. EXHIBIT 2, Page 8 of 56 CaseScientific 3:18-cv-03078-WHO Evaluation of Biological Opinions Document on Endangered 50-5 and Threatened Filed Fishes07/03/18 in the Klamath Page River 9... of 56

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OTHER REPORTS OF THE BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY The Airliner Cabin Environment and Health of Passengers and Crew (2002) Arsenic in Drinking Water: 2001 Update (2001) Evaluating Vehicle Emissions Inspection and Maintenance Programs (2001) Compensating for Wetland Losses Under the Clean Water Act (2001) A Risk-Management Strategy for PCB-Contaminated Sediments (2001) Toxicological Effects of Methylmercury (2000) Strengthening Science at the U.S. Environmental Protection Agency: Research- Management and Peer-Review Practices (2000) Scientific Frontiers in Developmental Toxicology and Risk Assessment (2000) Modeling Mobile-Source Emissions (2000) Toxicological Risks of Selected Flame-Retardant Chemicals (2000) Copper in Drinking Water (2000) Ecological Indicators for the Nation (2000) Waste Incineration and Public Health (1999) Hormonally Active Agents in the Environment (1999) Research Priorities for Airborne Particulate Matter: I. Immediate Priorities and a Long-Range Research Portfolio (1998); II. Evaluating Research Progress and Updating the Portfolio (1999); III. Early Research Progress (2001) Ozone-Forming Potential of Reformulated Gasoline (1999) Risk-Based Waste Classification in California (1999) Arsenic in Drinking Water (1999) Brucellosis in the Greater Yellowstone Area (1998) The National Research Council's Committee on Toxicology: The First 50 Years (1997) Carcinogens and Anticarcinogens in the Human Diet (1996) Upstream: Salmon and Society in the Pacific Northwest (1996) Science and the Endangered Species Act (1995) Wetlands: Characteristics and Boundaries (1995) Biologic Markers (5 reports, 1989–1995) Review of EPA's Environmental Monitoring and Assessment Program (3 reports, 1994–1995) Science and Judgment in Risk Assessment (1994) Ranking Hazardous Waste Sites for Remedial Action (1994) Pesticides in the Diets of Infants and Children (1993) Setting Priorities for Land Conservation (1993) Protecting Visibility in National Parks and Wilderness Areas (1993) Dolphins and the Tuna Industry (1992) Science and the National Parks (1992)

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Assessment of the U.S. Outer Continental Shelf Environmental Studies Program, Volumes I-IV (1991–1993) Human Exposure Assessment for Airborne Pollutants (1991) Rethinking the Ozone Problem in Urban and Regional Air Pollution (1991) Decline of the Sea Turtles (1990) Copies of these reports may be ordered from the National Academy Press

(800) 624–6242 or (202) 334–3313

www.nap.edu

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ACKNOWLEDGMENTS xi

ACKNOWLEDGMENTS

This project was supported by the U.S. Bureau of Reclamation, U.S. Fish and Wildlife Service, and National Marine Fisheries Service. We are thankful for the assistance provided by the Bureau of Indian Affairs in planning the first meeting. We are grateful for the informative briefings provided by the following individuals: Michael Belchik, Water Management and Rights Protection Division James Carpenter, Carpenter Design Inc. Paul Cleary, Oregon Department of Water Resources John Crawford, Tule Lake Irrigation District Larry Dunsmoor, The Klamath Tribes Natural Resources Thomas Hardy, Utah Water Research Laboratory Jacob Kann, Aquatic Ecosystems Sciences, LLC Ron Larson, Klamath Falls Fish and Wildlife Office Steven Lewis, Klamath Falls Fish and Wildlife Office Todd Olson, PacifiCorp Felice Pace, Klamath Forest Alliance Donald Reck, National Marine Fisheries Service Michael Ryan, U.S. Bureau of Reclamation Kenneth Rykbost, Oregon State University Klamath Experiment Station Rip Shively, U.S. Geological Survey, Klamath Field Station

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ACKNOWLEDGMENTS xii

Glen Spain, Pacific Coast Federation of Fishermen's Associations Sue Ellen Wooldridge, U.S. Department of Interior The committee's work also benefited from the written and oral testimony submitted by the public, and we appreciate their participation.

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ACKNOWLEDGMENT OF REVIEW PARTICIPANTS xiii

ACKNOWLEDGMENT OF REVIEW PARTICIPANTS

This report has been reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the NRC's Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the institution in making its published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their review of this report: Michael T.Brett, University of Washington Alex J.Horne, University of California, Berkeley John J.Magnuson, University of Wisconsin, Madison Douglas F.Markle, Oregon State University John M.Melack, University of California, Santa Barbara Lisa Speer, National Resources Defense Council Edwin A.Theriot, U.S. Army Corps of Engineers David A.Vogel, Natural Resource Scientists, Inc. Eugene B.Welch, University of Washington Donald Siegel, Syracuse University Margaret Strand, Oppenheimer, Wolff & Donnelly, LLP

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ACKNOWLEDGMENT OF REVIEW PARTICIPANTS xiv

Although the reviewers listed above have provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations, nor did they see the final draft of the report before its release. The review of this report was overseen by John C.Bailar, III, University of Chicago, and Paul G.Risser, Oregon State University. Appointed by the National Research Council, they were responsible for making certain that an independent examination of this report was carried out in accordance with institutional procedures and that all review comments were carefully considered. Responsibility for the final content of this report rests entirely with the authoring committee and the institution.

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PREFACE xv

PREFACE

The federal Endangered Species Act of 1973 has been invoked extensively for the protection of aquatic species in the western United States. Aquatic fauna of the West show extensive endemism because of genetic isolation associated with aridity and with the drainage of many rivers directly to the Pacific. Human intervention in the water cycle of the West is especially pervasive because of the general scarcity of water and the extensive redistribution of water in support of economic growth. Also, the West is growing and developing very rapidly. Thus, an unusual combination of biogeographic, hydrologic, and socioeconomic circumstances conspire to raise the likelihood that the legal protection of aquatic species will come into conflict with development and use of water in the West. Fishes in the Klamath River Basin are the focus of perhaps the most prominent current conflict between traditional uses of water in the West and requirements established by law for the protection of threatened and endangered species. This case is especially interesting in that the federal government is playing two potentially conflicting roles. Through the U.S. Bureau of Reclamation, the Department of the Interior is attempting to serve the needs of irrigators for water that is derived from the federal Klamath Basin Project. Not only is the delivery of water a contractual obligation of the government, it also is traditional in the sense that water delivery has occurred through the project for almost a century. At the same time, the U.S. Fish and Wildlife Service of the Department of the Interior and the National Marine Fisheries Service of the Department of Commerce are attempting to protect three threatened or endangered fishes of the Klamath Basin drainage (the Lost River

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PREFACE xvi

sucker, the shortnose sucker, and the Klamath Basin coho salmon). Interested parties, some of whom have livelihoods or cultural traditions at stake, include farmers, commercial fishing interests, Native Americans, environmental interests, hunters, and hydropower production interests. Conflicts became openly angry during 2001 when irrigators were deprived during a severe drought of traditionally available water through the government's issuance of jeopardy opinions on the endangered and threatened fishes. Economic losses were substantial and the changes in water management were a source of great frustration to irrigators. The Endangered Species Act (ESA) sets a framework for determination of future water use and management in the Klamath River Basin. The ESA is tightly focused on the requirements for survival of the threatened and endangered fishes, the survival of which is not negotiable under the ESA. Therefore, if the fishes require more water, ESA directs that they shall have it, which would imply that water managers and users must augment their water supplies, reduce their demands, or reach other accommodations consistent with the requirements of the species. While the ESA gives priority to the needs of threatened and endangered species, it also requires that any allocation of resources to these species be justified on a scientific or technical basis. The burden for scientific and technical justification falls mainly on the federal agencies, and especially the U.S. Fish and Wildlife Service and National Marine Fisheries Service, which are the source of biological opinions on the species. Assessment of the requirements of any species in a manner that is scientifically or technically rigorous is difficult and often cannot be accomplished quickly. The agencies have assembled considerable data and have interpreted the data as showing need for higher flows in the Klamath main stem and higher lake levels in the upper part of the basin. External review increases confidence in scientific and technical judgments, and is especially important when such judgments underlie important policy decisions. Accordingly, the Department of the Interior and Department of Commerce have arranged through its agencies for the National Research Council to form the Committee on Endangered and Threatened Fishes in the Klamath River Basin, whose charge is to conduct an external review of the scientific basis for the biological opinions that resulted in changes of water management for year 2001. The committee is to conduct its work in two phases. The first phase, which is reported here, gives an interim assessment of the evidence behind the biological opinions. A second phase, which will

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PREFACE xvii

occur over approximately the next year, will take a broader approach to evaluation of evidence for long-term requirements of the threatened and endangered fishes. In formulating its interim assessment, the committee has been greatly assisted by individuals who have provided it with information orally and in written form. The committee is especially indebted to the invited speakers and members of the public who attended the first meeting of the committee and also to NRC staff members Heather McDonald, Jennifer Saunders, David Policansky, and Suzanne van Drunick and to Leslie Northcott of the University of Colorado. All NRC committee reports are subject to external peer review as well as internal quality control processes. The committee and the NRC are grateful to the reviewers who contributed their time and expertise to the review process. The NRC committee is pleased to provide scientific and technical assessments that it hopes will be helpful to federal agencies as they attempt the difficult process of guiding water management toward practices that are consistent with the welfare of threatened and endangered species while also accommodating to the fullest practical extent other uses of water in the Klamath River Basin. William M.Lewis, Jr., Chair Committee on Endangered and Threatened Fishes in the Klamath River Basin

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PREFACE xviii

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CONTENTS xix

CONTENTS

Summary 1

1 Introduction 6

2 Evaluation of the Biological Opinion on Shortnose 11 andLost River Suckers

3 Evaluation of the Biological Opinion on Klamath 21 BasinCoho Salmon

4 Conclusions 26

References 28

Appendix : Statement of Task 32 Figure 1 Map of the Upper Klamath River Basin showing surface 7 waters and landmarks mentioned in this report (modified from USFWS sources) Figure 2 Overview of monthly levels for Upper Klamath Lake proposed 14 by USBR through its biological assessment of 2001, USFWS through its biological opinion of 2001, and observed conditions for the years 1960–1998. Hydrologic categories used by USBR in its proposals (dry years, critical dry years) are explained in the text. Mean depths, excluding wetlands, corresponding to water levels are as follows (feet): 4,137=3.5; 4,139=4.8; 4,140 =5.7; 4,141=6.6; 4,142=7.6 (Welch and Burke 2001, Figure 2–5)

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CONTENTS xx

Figure 3 Historical record of level at the end of September for UpperK- 16 lamath Lake (from USBR sources) Figure 4 Relationship of chlorophyll a and median August lake level in 18 Upper Klamath Lake between 1991 and 1998. Chlorophyll data are averages as reported by Welch and Burke (2001). Recruitment and mortality information is as reported by USFWS (2001) Figure 5 Estimated age frequency distributions using opercles from Lost 19 River suckers and shortnose suckers collected from 1997 fish kill in Upper Klamath Lake, Oregon. Estimates did not include all suckers collected, but were calculated using only suckers from which a length measurement (fork length) was obtained. Data are truncated from 1987 to 1994, additional information exists on other year classes of suckers. Source: USGS, unpublished data, 2001 Figure 6 Three flow regimes for the Klamath River below Iron Gate 24 Dam: USBR (USBR 2001b, minima for dry and critical years) proposed historical mean minima for dry and critical dry years, and RPA minimum flows from NMFS (2001). Hydrologic categories used by USBR in its proposals (dry years, critical dry years) are explained in the text

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xxi

INTERIM REPORT: SCIENTIFIC EVALUATION of BIOLOGICAL OPINIONS on ENDANGERED AND THREATENED FISHES in the KLAMATH RIVER BASIN

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xxii

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SUMMARY 1

SUMMARY

The Klamath River Basin, which drains directly to the Pacific Ocean from parts of southern Oregon and northern California, contains endemic freshwater fishes and genetically distinctive stocks of anadromous fishes. Endemic freshwater fishes include the shortnose sucker (Chasmistes brevirostris) and the Lost River sucker (Deltistes luxatus). These long-lived and relatively large species, which live primarily in lakes but enter flowing waters or springs for spawning, were sufficiently abundant during the nineteenth and early twentieth centuries to support commercial fisheries. During the last half of the twentieth century, these species declined so much in abundance that they were listed in 1988 as endangered under the federal Endangered Species Act (ESA). In addition, the genetically distinctive Southern Oregon/Northern California Coast (SONCC) coho salmon (Oncorhynchus kisutch), an evolutionary significant unit (ESU) of the coho salmon, depends on the Klamath River main stem for migration and on tributary waters for spawning and growth before entering the Pacific for maturation. The Klamath Basin coho has declined substantially over the last several decades and was listed as threatened under the ESA in 1997. Factors contributing to the decline in abundance of the endangered suckers and threatened coho in the Klamath River Basin are diverse and, in some cases, incompletely documented. Factors thought to have contributed to the decline of the endangered suckers include degradation of spawning habitat, deterioration in the quality of water in Upper Klamath Lake, overexploitation by commercial and noncommercial fishing (now regulated), introduction of competitive or predaceous exotic species, blockage of migration routes, and

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SUMMARY 2

entrainment of fish of all ages in water-management structures. Factors contributing to the decline of coho salmon are thought to include earlier overexploitation by fishing as well as continuing degradation of tributary habitat and reduced access to spawning areas. The threatened coho salmon also may be affected by changes in hydrologic regime, substantial warming of the main stem and tributaries, and continuing introduction of large numbers of hatchery-reared coho, which are derived only partly from native stock. The U.S. Bureau of Reclamation's (USBR) Klamath Basin Project (Klamath Project) is a system of main-stem and tributary dams and diversion structures that store and deliver water for agricultural water users in the Upper Klamath Basin under contract with the USBR. After the listing of suckers in 1988 and coho in 1997, the USBR was required to assess the potential impairment of these fishes in the Klamath River Basin by operations of the Klamath Project. In the assessments, which were completed in 2001, the USBR concluded that operations of the project would be harmful to the welfare of the listed species without specific constraints on water levels in the lakes to protect the endangered suckers and on flows in the Klamath River main stem to protect the threatened coho salmon. After release of the USBR assessment on the endangered suckers (February 2001) and following procedures required by the ESA, the U.S. Fish and Wildlife Service (USFWS) in April 2001 issued a biological opinion based on an extensive analysis of the relevant literature and field data. The biological opinion states that the endangered suckers would be in jeopardy under USBR'S proposed Klamath Project operations. The USFWS proposed a reasonable and prudent alternative (RPA) for operation of the Klamath Project. The RPA requires screening of water-management structures to prevent entrainment of suckers, adequate dam passage facilities, habitat restoration, adaptive management of water quality, interagency coordination in the development plans for operating the Klamath Project during dry years, further studies of the sucker populations, and a schedule of lake levels higher than those recommended by the USBR in its assessment. The National Marine Fisheries Service (NMFS), which assumes responsibility for the coho because it is anadromous, issued a biological opinion in April 2001 indicating that the operation of the Klamath Project as proposed by the USBR assessment of January 2001 would leave the coho population in jeopardy. The NMFS formulated an RPA incorporating reduced rates of change in flow (ramping rates) below main-stem dams to prevent stranding of coho, interagency coordination intended to optimize use of water for multiple purposes, and minimum flows in the Klamath River main stem higher than those proposed by USBR.

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SUMMARY 3

During 2001, a severe drought occurred in the Klamath River Basin. The U.S. Department of the Interior (DOI) determined that the newly issued biological opinions and their RPAs must prevail; thus, water that would have gone to irrigators was directed almost entirely to attempts to maintain minimum lake levels and minimum flows as prescribed in the two RPAs. The severe economic consequences of this change in water management led DOI to request that the National Research Council (NRC) independently review the scientific and technical validity of the government's biological opinions and their RPAs. The NRC Committee on Endangered and Threatened Fishes in the Klamath River Basin was formed in response to this request. The committee was charged with filing an interim report after approximately less than 3 months of study and a final report after about 18 months of study (see statement of task, Appendix). The interim report, which is summarized here, focuses on the biological assessments of the USBR (2001) and the USFWS and NMFS biological opinions of 2001 regarding the effects of Klamath Project operations on the three listed fish species. The committee conducted a preliminary assessment of the scientific information used by the agencies and other relevant scientific information, and has considered the degree to which the biological opinions are supported by this information. During November and early December 2001, the committee studied written documentation, heard briefings from experts, and received oral and written testimony from the public, and used this information as the basis for its interim report.

THE COMMITTEE'S PRINCIPAL FINDINGS The NRC committee concludes that all components of the biological opinion issued by the USFWS on the endangered suckers have substantial scientific support except for the recommendations concerning minimum water levels for Upper Klamath Lake. A substantial data-collection and analytical effort by multiple agencies, tribes, and other parties has not shown a clear connection between water level in Upper Klamath Lake and conditions that are adverse to the welfare of the suckers. Incidents of adult mortality (fish kills), for example, have not been associated with years of low water level. Also, extremes of chemical conditions considered threatening to the welfare of the fish have not coincided with years of low water level, and the highest recorded recruitment of new individuals into the adult populations occurred through reproduction in a year of low water level. Thus, the committee concludes that there is presently no sound scientific basis for recommending an operating regime for the Klamath Project that seeks to ensure lake levels

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SUMMARY 4

higher on average than those occurring between 1990 and 2000. At the same time, the committee concludes that there is no scientific basis for operating the lake at mean minimum levels below the recent historical ones (1990– 2000), as would be allowed under the USBR proposal. Operations leading to lower lake levels would require acceptance of undocumented risk to the suckers. For the Klamath Basin coho, the NMFS RPA involves coordination of operations as well as reduction of ramping rates for flows below the mainstem dams and increased flows in the Klamath River main stem. Coordination and reduced ramping rates are well justified. However, the committee did not find clear scientific or technical support for increased minimum flows in the Klamath River main stem. Although the proposed higher flows are intended to increase the amount of habitat in the main stem, the increase in habitat space that can occur through adjustments in water management in dry years is small and possibly insignificant. Furthermore, tributary conditions appear to be the critical factor for this population; these conditions are not affected by operations of the Klamath Project and therefore are not addressed in the RPA. Finally, and most important, water added as necessary to sustain higher flows in the main stem during dry years would need to come from reservoirs, and this water could equal or exceed the lethal temperatures for coho salmon during the warmest months. The main stem already is excessively warm. At the same time, reduction in main-stem flows, as might occur if the USBR proposal were implemented, cannot be justified. Reduction of flows in the main stem would result in habitat conditions that are not documented, and thus present an unknown risk to the population.

CONCLUSION On the basis of its interim study, the committee concludes that there is no substantial scientific foundation at this time for changing the operation of the Klamath Project to maintain higher water levels in Upper Klamath Lake for the endangered sucker populations or higher minimum flows in the Klamath River main stem for the threatened coho population. The committee concludes that the USBR proposals also are unjustified, however, because they would leave open the possibility that water levels in Upper Klamath Lake and minimum flows in the Klamath River main stem could be lower than those occurring over the past 10 years for specific kinds of climatic conditions. Thus, the committee finds no substantial scientific evidence supporting changes in the operating practices that have produced the observed levels in

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SUMMARY 5

Upper Klamath Lake and the observed main-stem flows over the past 10 years. The committee's conclusions are subject to modification in the future if scientific evidence becomes available to show that alteration of flows or water levels would promote the welfare of the threatened and endangered species under consideration by the committee. The committee will make a more comprehensive and detailed assessment of the environmental requirements of the endangered suckers and threatened coho in the Klamath River Basin over the next year, during which time it will develop final conclusions.

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INTRODUCTION 6

1 INTRODUCTION

The Klamath River Basin is isolated from other fresh waters by its direct drainage into the Pacific Ocean (Figure 1). Its isolation and diverse freshwater habitats, including perennial tributary and main-stem flows, extensive marshlands, and large shallow lakes, have favored the genetic isolation of freshwater and anadromous fishes in the basin. Thus, the Klamath River Basin contains endemic freshwater fishes as well as genetically distinctive stocks of anadromous fishes that are shared with nearby basins on the Oregon and California coasts. Endemic freshwater fishes of the Klamath River Basin include the shortnose sucker (Chasmistes brevirostris) and the Lost River sucker (Deltistes luxatus). These two species, which are long-lived, reach relatively large sizes, and have high fecundity (Moyle 2002), occupy primarily lakes as adults but also use tributary streams and springs for spawning. The two sucker species were abundant in Upper Klamath Lake and elsewhere in the drainage prior to 1900; they were used extensively by Native Americans as well as settlers, and were the basis for commercial fisheries (USFWS 2001). During the twentieth century, particularly after the 1960s, the populations declined substantially. Reduction in abundance of the suckers has been generally attributed to changes in water quality, excessive harvesting, introduction of exotic fishes, alteration of flows, entrainment of fish into water-management structures, and physical degradation of spawning areas (USFWS 2001). Both the shortnose sucker and the Lost River sucker were classified as endangered under the federal Endangered Species Act (ESA) in 1988 (USFWS 1988).

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INTRODUCTION 7

FIGURE 1 Map of the Upper Klamath River Basin showing surface waters and landmarks mentioned in this report. Source: modified from USFWS.

The main stem and tributaries of the Klamath River support endemic populations of a genetically distinctive population of coho salmon (Oncorhynchus kisutch). This group of coho is part of the Southern Oregon/ Northern California Coasts (SONCC) evolutionarily significant unit

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INTRODUCTION 8

(ESU), which also occupies several other drainages near the Klamath River Basin. These fish mature in marine waters off the California and Oregon coasts, move up the Klamath main stem and into tributaries for spawning, descend back to the main stem for the smolt phase, and then exit to the Pacific. The present distribution of the species within the Klamath Basin extends to the Iron Gate Dam, although it probably extended farther upstream prior to the construction of main-stem dams (NMFS 2001). Stocks of native coho salmon have declined greatly in the Klamath River Basin over the past several decades. Potential causes of the decline include overexploitation (now largely curtailed), habitat degradation, manipulation of flows in the main stem, excessive warming of waters, degradation or blockage of tributaries, and introduction of large numbers of competitive hatchery-reared coho salmon only partially derived from the native stock (NMFS 2001). The SONCC coho ESU was classified as federally threatened under the ESA in 1997. In response to the listing of the two sucker species and the SONCC coho, the Bureau of Reclamation (USBR), which operates the Klamath River water distribution project (Klamath Project), prepared biological assessments of the effects of Klamath Project operations on the suckers and on the coho (USBR 2001a, b). Because the listing processes for these fish referenced water level in Upper Klamath Lake and other lakes in the Upper Klamath Basin and amounts of flow in the main stem of the Klamath River below Iron Gate Dam as potential points of concern for the welfare of the species, the USBR assessments were intended to make a case for specific flows and water levels in portions of the basin strongly affected by operations of the Klamath Project. In response to the USBR assessment of the endangered suckers, the U.S. Fish and Wildlife Service (USFWS) issued a biological opinion (USFWS 2001). A separate biological opinion was issued on the coho population by the National Marine Fisheries Service (NMFS 2001), which has the prime responsibility for ESA actions on these fish because they are anadromous. The two biological opinions differed sharply from the two corresponding USBR assessments by calling for maintenance of higher lake levels and higher main- stem flows. Year 2001 brought a severe drought to the Klamath River Basin. The U.S. Department of the Interior (DOI) determined that the actions to protect the endangered and threatened species called for in the biological opinions of the USFWS and NMFS must take priority over other uses of water. The amounts of water specified as reasonable and prudent alternatives (RPAs) in the biological opinions should be maintained to the degree possible before provision of water for consumptive use as specified by contracts between

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INTRODUCTION 9

irrigators and the USBR through its Klamath Project. Consequently, most of the water that would have been delivered to irrigators through the Klamath Project was not delivered. Substantial agricultural losses occurred, along with damage to the economic base of the Klamath River Basin (actual losses are still being estimated, but the work of Adams and Cho (1998) shows that direct losses to farmers alone would probably exceed $20 million). Given the strong economic consequences for implementation of the biological opinions through their effect on the Klamath Project, the DOI determined that the scientific basis for the two opinions should be reviewed. The National Research Council (NRC) was asked to form a committee to review the two opinions. Sponsors of the review are USBR and the USFWS of the DOI and the NMFS of the U.S. Department of Commerce. A portion of the work of the NRC committee and the committee's interim conclusions are summarized in this report. The two biological opinions and the two biological assessments contain valuable literature reviews. The committee cites these documents in lieu of the primary literature for much of the background subject matter of this report, but cites individual studies that are of particular importance to the committee's conclusions wherever appropriate.

TASKS OF THE NRC COMMITTEE The work of the NRC Committee on Endangered and Threatened Fishes in the Klamath River Basin is divided into two phases (see statement of task, Appendix). The first phase, reported here, involves a preliminary assessment of the scientific validity of the two biological opinions and their RPAs, particularly as they relate to the near-term operation of the Klamath Project. In a second phase, the committee will conduct a broad-based study of the evidence related to the welfare of the endangered and threatened species. Whereas the interim report focuses specifically on the biological opinions, the final report may extend beyond the biological opinions to deal more extensively with water pollution or other such subjects that are not directly under control of the Klamath Project. This effort will culminate in a second report that will give the committee's consensus view of the long-term requirements of the species. Although the interim report specifically deals with the two biological opinions, the committee also gives its conclusions about the two biological assessments upon which the biological opinions are based. If the biological opinions were rejected fully or in part, the presumed alternative for operation of the Klamath Project would be as prescribed in the USBR assessments.

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INTRODUCTION 10

Thus, the committee must not only evaluate the validity of the biological opinions, but also extend the same sort of evaluation to the assessments. The tasks of the committee encompass only the scientific and technical issues that are relevant to the endangered sucker and threatened coho species. The committee is not charged with investigating or reporting on economic dislocation or with forecasting the economic consequences of continued implementation of flows specified in the biological opinions. Given the background materials provided to the committee, however, all committee members are aware of the importance of any change in historical management of flows to water users in the Klamath Basin. Also, the committee is aware of the long-standing interest of Native American tribes of the Klamath River Basin in the maintenance and expansion of fish stocks, including Tribal Trust species not covered in this report, and of the interests of numerous other parties in water resources, wetlands, and the welfare of fishes and other aquatic life. Although the committee will not analyze economic or socioeconomic questions, it recognizes the interest of individuals and communities in the Klamath Basin in the conclusions of the committee. Not only from an economic and social point of view but also from a perspective of ecological and biological resources, the work of the NRC committee focuses on its statement of task and on the inherent requirements of the ESA, which prohibits federal actions that jeopardize continued existence of listed species through interference with their survival or recovery (50 CFR 402.02). The Klamath River Basin is home to hundreds of species of fish and wildlife and to distinctive native ecosystems, including wildlife refuges of national significance. Many of these natural resources have been greatly restricted or altered through human action. In fact, changes in the flow regime in the Klamath River may affect other fishes that have not yet been proposed for listing as threatened species but have not yet been listed (e.g., ESUs of steelhead and chinook salmon). The committee is charged, however, with studying the requirements of the shortnose and the Lost River suckers and the coho salmon and not those of other species in the Klamath River Basin.

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2 EVALUATION OF THE BIOLOGICAL OPINION ON SHORTNOSE AND LOST RIVER SUCKERS Populations of the shortnose and Lost River suckers currently are present within Upper Klamath Lake on the north side of the Klamath River drainage and within Clear Lake (which operates as a reservoir) and Gerber Reservoir on the Lost River to the southeast (Figure 1). Small groups of individuals, some or all of which may be nonreproducing, are found elsewhere in the Klamath River drainage, including Tule Lake sump (USFWS 2001). Conditions in the lakes are relevant to the USFWS biological opinion because of its proposals for minimum lake levels that are intended to reduce mortality and improve spawning success, recruitment (addition of new individuals to the population), growth, and condition of the suckers. The population sizes of endangered suckers in Upper Klamath Lake and elsewhere within the Klamath Basin are uncertain, but the abundances of these populations, which once were large enough to support commercial fisheries, are much lower than they were when agricultural development and water management began. Unfortunately, quantitative estimates of population sizes are not available. During the 1980s, qualitative evidence indicated that declines might have reduced the sucker populations in Upper Klamath Lake to just a few thousand old (greater than 10 years) fish (USFWS 1988). More recent estimates that were made possible incidentally by episodes of mass

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mortality suggest, however, that the populations are considerably larger than they appeared to be in the 1980s, and that some recruitment to the adult age classes has occurred in most or all years of the last decade (see below). Population sizes might range from a few tens of thousands to the low hundreds of thousands (USFWS 2001) but still are much lower than they were originally. Aside from the decline in abundance over the long term, other indications of problems within the sucker populations include absence of spawning at a number of sites historically used for spawning, apparent increase in mass mortality of adults (“fish kills”), and weak recruitment in most years (USFWS 2001). The water quality of Upper Klamath Lake has changed substantially over the past several decades. The lake appears to have been eutrophic (rich in nutrients and supporting high abundances of suspended algae) prior to any anthropogenic influence (Kann 1998). Mobilization of phosphorus from agriculture and other nonpoint sources (Walker 2001), appears, however, to have pushed the lake into an exaggerated state of eutrophication that involves algal blooms reaching or approaching the theoretical maximum abundances. In addition, algal populations now are strongly dominated by the single blue-green algal species Aphanizomenon flos-aquae (cyanobacteria) rather than the diatom taxa that apparently dominated blooms before nutrient enrichment (Kann 1998, Eilers et al. 2001). Evidence indicates that changes in the water quality of Upper Klamath Lake have increased mass mortality among adult suckers. Under certain conditions, the bottom portion of the water column in the lake develops oxygen depletion, either no oxygen (anoxia) or lower than normal oxygen levels (hypoxia), and accumulates high concentrations of ammonia. Mixture of those bottom waters with the surface waters under the influence of changes in the weather likely causes mass mortality (Vogel et al. 2001). Although mass mortality has been recorded over the observed history of the lake, its frequency appears to have increased (Perkins et al. 2000). Major incidents were recorded for 1995, 1996, and 1997; low dissolved oxygen appears to have been the direct cause of mortality in these years (Perkins et al. 2000). Impaired water quality also might stress fry through high pH in surface waters resulting from high rates of photosynthesis, although exposures to the highest pH probably are too brief to cause mortality (Saiki et al. 1999). In addition, the present trophic state of the lake potentially poses a threat of mortality in winter, when anoxia can occur under the ice if oxygen demand is high. Although not yet observed, winter mortality could occur in the future (Welch and Burke 2001). Factors of concern other than water quality include the presence of exotic species capable of inducing types of predation and competition that are

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evolutionarily foreign to these endemic species. Hybridization occurs but the degree of threat associated with it is unknown; the native suckers probably showed some interbreeding prior to human intervention (Markle et al. 2000). In addition, access of the suckers to historically significant spawning areas has in many cases been blocked or the spawning areas themselves have been physically degraded to such an extent that they cannot serve their former roles (USFWS 2001). Overfishing or habitat degradation might have eliminated portions of the population that were using specific spawning areas and although fishing no longer occurs, these subpopulations cannot be regenerated without manipulation of existing stocks in combination with habitat restoration. Suckers of all sizes are entrained by water-management structures (USFWS 2001). Although screening of these structures has long been recognized as an important means of reducing mortality of the endangered suckers, it has not yet been accomplished. Also, interaction of multiple stresses may increase vulnerability of the endangered suckers to disease, degrade their body condition, and cause them to show a high incidence of anatomical abnormalities. The USFWS biological opinion states that the Klamath Project contributes directly to mortality and adverse environmental conditions for the endangered suckers. On this basis, USFWS presents a reasonable and prudent alternative (RPA) consisting, in summary, of requirements for minimum lake levels, interagency coordination and adaptive management, screening to prevent entrainment of fish, creation of improved passage facilities, steps toward improvement of habitat and water quality, and additional studies. The RPA is intended to avoid jeopardizing listed species either directly or through adverse modification of critical habitat (50 CFR 402.02). With the exception of the recommendation on lake-level maintenance, there is good scientific or technical support for all the requirements listed in the RPA. Interagency coordination and adaptation of management are advisable, especially because the information base is evolving rapidly and because annual optimization of strategies for using water is an obvious need. Given the documented loss of suckers to entrainment and the blockage of their access to spawning waters at known locations (USFWS 2001), requirements of the RPA calling for mitigation of these problems also seems highly defensible. Potential for improvement of habitat and water quality must be viewed as incremental rather than comprehensive, but even incremental improvements offer the prospect of increasing the viability of the sucker populations and thus seem justified. Recommendations on water level are more difficult to evaluate, however.

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FIGURE 2 Overview of monthly levels for Upper Klamath Lake proposed by USBR through its biological assessment of 2001, USFWS through its biological opinion of 2001, and observed conditions for the years 1960–1998. Hydrologic categories used by USBR in its proposals (dry years or critical dry years) are explained in the text. Mean depths, excluding wetlands, corresponding to water levels are approximately as follows (feet): 4,137=3.5; 4,138=4.0; 4,139=4.8; 4,140=5.7; 4,141=6.6; 4,142 =7.6 (Welch and Burke 2001).

Figure 2 shows the water levels given by USFWS in its RPA (2001) as well as two other lake-level regimes (USBR recommended and historical). The USFWS requirements are given as absolute minimums (i.e., they do not vary from one type of water-level year to another). In contrast, assessment

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proposals of the USBR are framed for categories of water-level year. Categories shown in Figure 2 are characterized as critically dry (lowest 4%) and dry (approximately 12% of years just wetter than the critically dry ones). The span of lake level records that the USBR chose to use in its analysis (1960–1998) reflects the full interval of operations for the completed Klamath Project. Even earlier records are available, extending back to the creation of Link River Dam in 1919 (Figure 3), but the interval between 1919 and 1960 would not be typical from the viewpoint of current project operations. Records prior to 1919, extending back to 1905, also are available (Figure 3); they show higher maximum and minimum lake levels than have been typical of Upper Klamath Lake since closure of the dam. In addition, operation of the Klamath Project has created a higher amplitude of intraannual variation in lake level and a change in seasonality of intraannual change in lake level as compared with the original condition of the lake (USFWS 2001, III. 2., page 38). While the operating interval between 1960 and 1998 is very useful for judging the degree of variability that can be expected in lake levels over a long period of years with the Klamath Project in place, the possibility for use of lake- level data in environmental analysis is limited to a much shorter interval. Interaction between lake level and environmental variables or indicators of the welfare of the endangered fish is dependent on concurrent information for lake level, environmental conditions, and fish. While information of a sporadic or anecdotal nature is available over as much as 100 years, routinely collected data on environmental characteristics and fish are available only since 1990 or later. Thus, while the long-term lake level record seems to invite statistical analysis of the welfare of fish in relation to lake level, the information at hand is actually limited to a period often years or less. This limitation explains the focus of this report and of the USFWS biological opinion on data extending over approximately the last ten years. All three lake-level regimes (USFWS RPA, USBR recommended and historical) reflect seasonality that is partly inherent in the runoff reaching Upper Klamath Lake and partly a by-product of water withdrawals. The degree of seasonality in the USFWS RPA is considerably lower, however, than the seasonality of the other two regimes depicted in Figure 2, and minimum levels are highest overall for the USFWS RPA. The USBR proposed minimums are below the mean lake levels for the historical operating regime in each of the two dry-year categories, because the USBR used the lowest recorded monthly lake levels as its proposed minimums for each category. From the viewpoint of lake levels, water years are almost independent of each other because the lake has little capacity for interannual storage. The USBR proposal would allow more drawdown of lake level than has been characteristic in the past. Although the lake levels proposed by USBR

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have been observed over the past 40 years, the use of these 40-year minimums as year-to-year minimums indicates that drawdown to the 40-year minimums would be possible in any year of future operations if USBR'S proposals were accepted. If USBR chose to operate the project by using greater average drawdown than has been observed over the past 40 years, the result would be substantially lower mean lake levels in each of the hydrologic categories.

FIGURE 3 Historical record of level at the end of September for Upper Klamath Lake. Source: USBR.

Control of lake levels as a means of advancing the welfare of the endangered suckers raises more difficult scientific issues than the other requirements listed by the USFWS in its RPA. The recommendation for water- level control is based on concerns related to habitat (shoreline spawning areas and emergent vegetation), and water quality (low oxygen in summer, need for deep-water refugia in summer and fall, and possibility of adverse conditions under ice cover). Impairment of water quality, primarily through eutrophication of Upper Klamath Lake, is a cause of mortality and stress for sucker populations. As indicated above, the present scientific evidence for this association is credible. An essential premise of the lake-level recommendations is that the adverse water-quality conditions known to stress or kill the endangered suckers are associated with the lowest water levels within the recent historical range of levels (since 1990, when consistent documentation first began). Presumption of this connection, which is essential to the arguments for specific lake levels

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proposed in the RPA, is inconsistent with present information on Upper Klamath Lake. Control of phosphorus in Upper Klamath Lake offers the potential of suppressing population densities of algae, thus improving water quality in the lake (Welch and Burke 2001). No relationship between lake levels and population densities of algae (as shown by chlorophyll) is evident, however, in the 9-year water-quality monitoring record that has been fully analyzed (Figure4). Thus, the idea of relieving eutrophication through phosphorus dilution caused by higher lake levels is not consistent with the irregular relationship between chlorophyll and lake level. Also, lake level fails to show any quantifiable association with extremes of dissolved oxygen or pH (see data presented by Welch and Burke 2001). For example, the most extreme pH conditions recorded for the lake over the past 10 years occurred in 1995 and 1996, which were years of intermediate water level, and not in 1992 and 1994, when water levels were lowest. (These two years had the lowest recorded water levels since 1950.) Furthermore, a substantial mass mortality occurred in 1971, the year of highest recorded water levels since 1950 (USFWS 2001), and within the last ten years, mortality of adults was highest in 1995, 1996, and 1997, none of which were years of low water level. The absence of notable adult mortality in any year of low water during the 1990s might in fact suggest an association the reverse of the one postulated in the biological opinion, although the evidence is statistically inconclusive. The USFWS itself has found no association of mass mortality with lake levels (USFWS 2001, III.2.70). Intensified eutrophication now affects the characteristics of the lake every year, and thus may constitute a threat to the suckers regardless of interannual variation in water level. Higher water levels are potentially supported on the grounds of improved survival of fry or juveniles rather than suppression of adult mortality. Higher water levels could reduce the likelihood that spawning areas around the lake would be dewatered and could be favorable to fry or juveniles. Abundance of juvenile suckers has been monitored since 1991 on the basis of seining (Simon et al. 2000a). This information, which must be used cautiously because it is not quantitative, indicates low abundances of juveniles in the drought years 1992 and 1994 but not in drought year 1991. Abundances also were low in non- drought years 1997 and 1998. Simon et al. (2000a) have reported generally declining abundance during the non-drought interval 1995–1998. They have also shown (Simon et al. 2000a, b) that the abundance of age 1+ suckers consistently has been very low, suggesting a bottleneck at this life stage, but interpretation of the data is complicated by very low efficiency for catching fish older than one year. Overall, the study of young fish shows no clear pattern associated with lake level.

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FIGURE 4 Relationship of chlorophyll a and median August lake level in Upper Klamath Lake between 1991 and 1998. Chlorophyll data are averages reported by Welch and Burke (2001). Recruitment and mortality data are from USFWS (2001).

The most reliable current information on recruitment is through analysis of age-class structure of adult suckers (USFWS 2001, III. 2., page 43). This data record is not consistent with the underlying assumptions of proposals for maintenance of higher water levels. The strongest recruitment (as inferred from relative abundances of adult year classes) observed over the last ten years was for 1991 (Figure 5), which falls within the lowest 15% of lake levels since 1950. Furthermore, as shown by the continuing strength of the 1991 year class in 1995 and beyond, the year class showed good survival through the dry years of 1992 and 1994. While the use of emergent vegetation by fry is cited as a reason for maintaining high water levels, the combination of high recruitment in 1991 and low recruitment in other years (as inferred from year class data) casts doubt on the importance of this factor, at least within the operating range of the 1990s. Overall, the presumed causal connections between lake levels and recruitment of the sucker populations in Upper Klamath Lake do not have strong scientific support at present.

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FIGURE 5 Estimated age frequency distributions using opercles from Lost River suckers and shortnose suckers collected from 1997 fish kill in Upper Klamath Lake, Oregon. Estimates did not include all suckers collected, but were calculated using only suckers from which a length measurement (fork length) was obtained. Data are truncated from 1987 to 1994, additional information exists on other year classes of suckers. Source: USGS, unpublished data, 2001.

Mortality possibly could be caused by multiple factors that interact with lake level. For example, mortality of suckers is influenced by changes in water column stability; an extended period of stability leading to decline of oxygen near the bottom can be followed by sudden mixing of the entire water column associated with a change in weather (high wind velocity). Thus, interpretation of information on lake level is complicated by the influence of weather. There is no evidence as yet, however, that the significance of undesirable mixing events is higher when lake levels are low than when they are high. As a result, mixing as a cause of water quality conditions leading to mortality cannot be interpreted at this time in terms of lake level. Despite a monitoring record of substantial length, there is no clear evidence of a connection between the lake levels and the welfare of the two sucker species in Upper Klamath Lake. Lake levels cannot be reduced, however, below those observed in the past 10 years without risk of adverse occurrences that are not described in the detailed monitoring record (1990- present; analyses complete through 1998). A negative association between the wel

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fare of the species and the lake level could emerge if lake levels are reduced below those of recent historical experience. The absence of any empirical connection between the observed lake levels and the welfare of the endangered suckers cannot be taken as justification for continuous or frequent operation of the lake at the lowest possible levels, given that the effects of operating the lake at lower levels are undocumented. Thus, while the observational record contradicts important underlying assumptions of the RPA, it does not provide an endorsement for the lake levels proposed in the USBR biological assessment, which, if implemented, could take interannual mean lake levels well below those of recent historical observation. The potential benefits of higher lake levels in Clear Lake, Gerber Reservoir, and Tule Lake sump are more difficult to evaluate, because the record of analysis and observation for these water bodies is not as extensive as that for Upper Klamath Lake. These lakes have not suffered notable mass mortality in association with low lake levels, but Clear Lake populations showed poor body condition following severe drawdown in the early 1990s. The USFWS provides reasonable support for lake levels in Clear Lake no lower than the recent drought-related minimum (1992–1993:4,519 feet). The RPA reasonably adds a margin of two feet (4,521) to allow for water loss in the absence of withdrawals under drought conditions.

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3 EVALUATION OF THE BIOLOGICAL OPINION ON KLAMATH BASIN COHO SALMON Coho salmon enter the main stem of the Klamath River for spawning typically in their third year, primarily between October and December. Over most of this interval, main-stem flows below Iron Gate Dam often are high (about 2,500–3,000 cubic feet per second) (NMFS 2001). Thus, standard methods for observing and counting spawning fish are not easily applied, and the size of the spawning population is unknown. Approximations suggest that the entire ESU has about 10,000 spawning coho salmon of nonhatchery origin per year (Weitkamp et al. 1995). Only a small portion of that number is associated with the Klamath Basin, where several important tributary runs have been reduced to a handful of fish (NMFS 2001). Spawning coho in the Klamath Basin are restricted to use of tributaries that they can reach from the main stem up to Iron Gate Dam. Original spawning runs probably were largest in large tributaries but currently are restricted mainly to numerous small tributaries entering the main stem directly (Yurok Tribe 2001, as cited in NMFS 2001). Large tributaries have been severely degraded, show excessively high temperatures, and are dammed in critical places. Although a minor amount of spawning may occur in the main stem, the main stem serves adults primarily as a migration route. Fry appear in late fall or winter, when water levels are highest. Most fry probably remain in the tributaries but some move or are swept into the main

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stem, where they can be found in small numbers well into July. Juvenile coho become smolts and emigrate to the ocean between March and mid-June; peak migration occurs in mid-May (NMFS 2001). In general, juvenile coho can be expected to occupy places where summer temperatures are low (12–14°C appears to be optimal for growth). They are also favored by deep pools with complex cover, especially large woody debris, which is essential for survival over winter (Sandercock 1991). Such conditions exist primarily in tributary streams of the Klamath Basin. The reduction in stocks of native coho salmon in the Klamath River Basin has been caused by multiple interactive factors. Drastic reduction in spawning and juvenile habitat has occurred through impoundment and physical alteration of tributaries. Also, large numbers of smolts are released annually from the Iron Gate hatchery. Smolts, which are derived from a combination of Klamath Basin and Columbia River coho, likely compete with or have other negative effects on wild native coho at all stages of their life history, including the smoltification- emigration period, the ocean growth period, and spawning (Fleming and Gross 1993, Nielsen 1994, NRC 1996). Physical habitat in the main stem is a potential concern for the welfare of the coho in several life stages. The spawning run must have adequate flows for passage, which would be impaired by excessively shallow water (e.g., through amplification of predation losses). Access to tributaries is a related consideration for the spawning run, given that little spawning is likely in the main stem. Also, fry that enter the main stem must find cool, well-shaded pools, or return to a suitable tributary. Smolts moving downstream must find suitable temperature, flow, and habitat conditions compatible with their physiological transformation during migration (Wedemeyer et al. 1980). Habitat is an undeniable requirement for all life stages; however, assessment of habitat suitability is difficult and subject to considerable uncertainty. Numerical methods are now being applied to the estimation of habitat area in relation to flow (INSE 1999). These methods are commonly used in evaluating habitat, but in final form they require extensive field measurements that are not yet available. Initial modeling suggests that even though habitat for salmonids increases with higher flows, the percentage increase of habitat space corresponding to increases in flow during dry years is relatively small (INSE 1999, NMFS 2001). Water temperature is a major concern for the welfare of the Klamath Basin coho salmon. Summer temperatures appear to be especially critical. In the nearby Matolle River, which contains coho that are part of the SONCC ESU, the juvenile coho reside almost entirely in tributaries but do not persist when summer daily maximum temperatures exceed 18°C for more than a week (Welsh et al. 2001). Summer temperatures in the Klamath River main

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stem are, as judged by the literature on thermal tolerance, suboptimal or even lethal to juveniles (NMFS 2001). High temperatures are the result of reduced flow in the main stem and in tributaries as a result of diversions, warming of water in lakes prior to its flow to the main stem, and loss of shading. Climate variability, although probably responsible for some interannual thermal variation, is unlikely to be an important factor by comparison with changes in flow and loss of riparian vegetation. Modeling has shown that higher releases of water to the main stem can reduce water temperature slightly (Deas and Orlob 1999) while also reducing the amplitude of daily temperature fluctuations, provided that manipulation of flow itself does not raise the base temperature (see below). It is unlikely, however, that the small degree of cooling that could be accomplished in this way would affect survival of coho salmon because temperatures would continue to be suboptimal. Further modeling is in progress. The biological opinion issued by the NMFS for the Klamath Basin coho salmon states that the Klamath Project harms coho in the Klamath main stem (NMFS 2001). The NMFS presents an RPA with three components: (1) higher monthly minimum flows for the main stem of the Klamath River for April through November as a means of maximizing habitat space in the main stem and suppressing maximum water temperatures, (2) suppression of ramping rates below Iron Gate Dam, and (3) coordination involving other agencies. Figure 6 shows minimum flows given by NMFS as part of its RPA and shows minimum flows proposed by USBR as part of its biological assessment as well as historical low flows in dry and critical dry years (note that in selected months flows can be higher in critical dry years than in dry years because of water management practices). The NMFS RPA proposed low flows are well above historical operating conditions, which in turn are above the minimum flows proposed by USBR. The proposed low-flow limits on the Klamath River might not benefit the coho population significantly. Although the provision of additional flow seems intuitively to be a prudent measure for expanding habitat, the total habitat expansion that is possible with the limited water available in dry years is not demonstrably important to maintenance of the population. In wet years, any benefits from increased flow will be realized without special limits. Year classes that have high relative strength should have emerged from the wet years of the recent past flow regime if flow is limiting. This does not appear to have been the case in the past decade, however. Thus, factors other than dry-year low flows appear to be limiting to survival and maintenance of coho. Higher flows might work to the disadvantage of the coho population from July through September if the source of augmentation for flow is warmer than

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the water to which it is added. Flows in the main stem include not only water passing the Iron Gate Dam but also accruals from ungaged sources consisting of groundwater and small tributaries. Thus, the addition of larger amounts of water from the sequence of reservoirs above Iron Gate Dam might be disadvantageous to the fish. This issue apparently has not yet been studied in any rigorous manner, yet it is critical to the evaluation of higher flows in the warmest months.

FIGURE 6 Three flow regimes for the Klamath River below Iron Gate Dam: USBR (2001b, minima for dry and critical years) proposed minimum flows for dry and critical years, historical mean minimum flows for dry and critical dry years, and RPA minimum flows (NMFS 2001). Hydrologic categories used by USBR in its proposals (dry years and critical dry years) are explained in the text.

Increased flows also could have a detrimental effect on the availability of thermal refugia (mainly mouths of small tributaries). Thermal refugia may be most accessible and most extensive at low flows. Increase in flows might reduce the size of these refugia by causing more effective mixing of the small amounts of locally derived cool water with much larger amounts of warm water from points upstream.

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Progressive depletion of flows in the Klamath River main stem would at some point be detrimental to coho salmon through stranding or predation losses. Thus, incremental depletions beyond those reflected in the recent historical record could be accomplished only with increased risk to coho salmon. At the same time, the available information provides little support for benefits presumed to occur through the increase of flows beyond those of the past decade. While single-year or multiple-year averages of low-flow extremes beyond those presently reflected in the record cannot be supported, there also is presently little evidence of a scientific nature that increased low flows will improve the welfare of the coho salmon. Modeling of temperature and habitat might be useful, but convincing evidence of a relation between the welfare of the coho and environmental conditions must be drawn to some extent from direct observation. For example, when related to specific flow conditions, year of class strength, abundance of various life history stages, or other biological indicators of success would greatly improve the utility of modeling and other information. The small size and scattered nature of the present native coho population makes collection of such data difficult, however. The RPA requirements related to ramping rates and interagency coordination seem supportable. Given direct field observation of the stranding of coho at the current ramping rates (NMFS 2001) and the mortality that is implicit in these observations, reduction in ramping rates seems a reasonable and prudent measure for protection of coho. Coordination, a final requirement of the RPA, is an obvious necessity because of the need to optimize use of water for multiple purposes.

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CONCLUSIONS 26

4 CONCLUSIONS

The NRC Committee on Endangered and Threatened Fishes in the Klamath River Basin has studied the USBR biological assessment on the shortnose and Lost River suckers, the USFWS biological opinion with its reasonable and prudent alternative (RPA) on these same species, and supporting documentation. The committee also has heard oral presentations and open public comment on the issues related to these endangered fishes in the Klamath River Basin. The committee finds strong scientific support for the requirements of the RPA except for the requirement for specified minimum lake levels in Upper Klamath Lake. Extensive field data on the fish and environmental conditions in Upper Klamath Lake do not provide scientific support for the underlying premise of the RPA that higher lake levels will help maintain or lead to the recovery of these two species. At the same time, operation of Upper Klamath Lake at mean minimum levels below the recent historical levels (1990– 2000), as could occur through implementation of the USBR assessment, would pose unknown risks, because these conditions have not been observed over the last 10 years, the interval over which good environmental documentation is available. The present scientific record is consistent with use of operational principles in effect between 1990 and 2000. The NRC committee has studied the USBR biological assessment on the Southern Oregon/Northern California Coasts evolutionary significant unit of the coho salmon in the Klamath River Basin and the accompanying biological opinion prepared by NMFS, its RPA requirements, and supporting documentation. The committee also heard oral presentations of scientists contributing

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CONCLUSIONS 27

to research on this issue and open public testimony. The RPA contains requirements for minimum flows in the Klamath River below Iron Gate Dam, limitations on ramping rate below Iron Gate Dam, and interagency coordination. The committee finds reasonable scientific support for the suppression of ramping rates as given in the RPA and for coordination. The committee does not find scientific support for the proposed minimum flows as a means of enhancing the maintenance and recovery of the coho population. The proposal of USBR, however, as given in its biological assessment, could result in more extreme suppression of flows than has been seen in the past and cannot be justified. On the whole, there is no convincing scientific justification at present for deviating from flows derived from operational practices in place between 1990 and 2000. The conclusions of the NRC committee as presented above apply to interim management of the Klamath Project. The committee will make a separate analysis of the scientific evidence, including any new evidence, supporting various actions that might result in improvements in stocks of endangered suckers and coho salmon in the Klamath River Basin over the long term.

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REFERENCES 28

REFERENCES

Adams, R.M. and S.H.Cho. 1998. Agriculture and endangered species: an analysis of trade-offs in the Klamath basin, Oregon. Water Resour. Res. 34(10): 2741– 2749. Cooperman, M. and D.F.Markle. 2000. Ecology of Upper Klamath Lake Shortnose and Lost River Suckers. 2. Larval Ecology of Shortnose and Lost River Suckers in the Lower Williamson River and Upper Klamath Lake. Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR. 27 pp. Deas, M.L., and G.T.Orlob. 1999. Klamath River Modeling Project. Report No. 99– 04. Davis: Center for Environmental and Water Resources Engineering, Dept.of Civil and Environmental Engineering, Water Resources Modeling Group, University of California. December. Eilers, J.M., J.Kann, J.Cornett, K.Moser, A. St. Amand, and C.Gubala. 2001. Recent Paleolimnology of Upper Klamath Lake, Oregon. Prepared by J.C.Headwaters, Inc., for U.S. Bureau of Reclamation, Klamath Basin Area Office, Klamath Falls, OR. March 16, 2001. Fleming, I.A., and M.R.Gross. 1993. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecol. Appl. 3(2):230–245. INSE (Institute for Natural Systems Engineering). 1999. Evaluation of Interim Instream Flow Needs in the Klamath River: Phase I. Final report. Prepared for the Department of the Interior. August 1999. 53 pp.+appendices. Kann, J. 1998. Ecology and Water Quality Dynamics of a Shallow Hypereutrophic Lake Dominated by Cyanobacteria. Ph.D. Dissertation. University of North Carolina, Chapel Hill, NC. 110 pp. Markle, D.F., M.R.Cavalluzzi, T.E.Dowling, and D.Simon. 2000. Ecology of Upper Klamath Lake Shortnose and Lost River Suckers: 4. The Klamath Basin

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REFERENCES 29

Sucker Species Complex. Annual report: 1999. Prepared by Department ofFisheries and Wildlife, Oregon State University, Corvallis, OR, and Department of Biology, Arizona State University, Tempe, AZ, for U.S. Biological Resources Division, U.S. Geological Survey, Oregon State University, Corvallis, OR, and U.S. Bureau of Reclamation, Klamath Falls, OR. July 26, 2000. Moyle, P.B. 2002. Inland Fishes of California, revised and expanded. Berkeley: University of California Press. 502 pp. Nielsen, J.L. 1994. Invasive cohorts: impacts of hatchery-reared coho salmon on the trophic, developmental, and genetic ecology of wild stocks. Pp. 361–385 in Theory and Application in Fish Feeding Ecology, D.J.Stouder, K.L.Fresh, and R.J.Feller, eds. Columbia: University of South Carolina Press. NMFS (National Marine Fisheries Service). 2001. Biological Opinion. Ongoing Klamath Project Operations. National Marine Fisheries Service, Southwest Region, National Oceanic and Atmospheric Administration, Long Beach, CA. April 6, 2001. [Online]. Available: http:// swr.ucsd.edu/psd/kbo.pdf. [January 28, 2002]. NRC (National Research Council). 1996. Upstream: Salmon and Society in the Pacific Northwest. Washington, DC: National Academy Press. Perkins, D.J., J.Kann, and G.G.Scoppettone. 2000. The Role of Poor Water Quality and Fish Kills in the Decline of Endangered Lost River and Shortnose Suckers in the Upper Klamath Lake. Biological Resources Division, U.S. Geological Survey. Final Report. Contract 4–AA–29– 12160. Submitted to the U.S. Bureau of Reclamation, Klamath Falls Project Office, Klamath Falls, OR. September 2000. Saiki, M.K., D.P.Monda, and B.L.Bellerud. 1999. Lethal levels of selected water quality variables to larval and juvenile Lost River and shortnose suckers. Environ. Pollut. 105(1):37–44. Sandercock, F.K. 1991. Life history of coho salmon (Oncorhynchus kisutch). Pp. 397–445 in Pacific Salmon Life Histories, C.Groot, and L.Margolis, eds. Vancouver: University of British Columbia Press. Simon, D.C., M.R.Terwilliger, and D.F.Markle. 2000a. Ecology of Upper Klamath Lake Shortnose and Lost River Suckers: 3. Annual Survey of Abundance and Distribution of Age 0 Shortnose and Lost River Suckers in Upper Klamath Lake. Annual report: 1999. Oregon Cooperative Research Unit, Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR, for U.S. Biological Resources Division, U.S. Geological Survey, Oregon State University, Corvallis, OR, and U.S. Bureau of Reclamation, Klamath Falls, OR. March 31, 2000. 45 pp. Simon, D.C., M.R.Terwilliger, P.Murtaugh, and D.F.Markle. 2000b. Larval and Juvenile Ecology of Upper Klamath Lake Suckers: 1995–1998. Final report. Prepared by Oregon Cooperative Research Unit, Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR, for U.S. Bureau of Reclamation, Klamath Falls, OR. January 7, 2000. 108 pp. USBR (U.S. Bureau of Reclamation). 2001a. Biological Assessment of Klamath Project's Continuing Operations on the Endangered Lost River Sucker and Short

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nose Sucker. U.S. Bureau of Reclamation, Mid-Pacific Region, Klamath Basin Area Office, Klamath Falls, OR. February 13, 2001. [Online]. Available: http:// www.mp.usbr.gov/kbao/esa/34_final_sucker_bo_4_06_01.pdf. [August 18, 2001]. USBR (U.S. Bureau of Reclamation). 2001b. Biological Assessment of the Klamath Project's Continuing Operations on Southern Oregon/Northern California ESU Coho Salmon and Critical Habitat for Southern Oregon/Northern California ESU Coho Salmon. U.S. Bureau of Reclamation, Mid-Pacific Region, Klamath Basin Area Office, Klamath Falls, OR. January 22, 2001. [Online]. Available: http://www.mp.usbr. gov/kbao. [August 18, 2001]. USFWS (U.S. Fish and Wildlife Service). 1988. Endangered and threatened wildlife and plants: Determination of endangered status for the shortnose sucker and Lost River sucker. Fed. Regist. 53(137): 27130–27134. USFWS (U.S. Fish and Wildlife Service). 2001. Biological/Conference Opinion Regarding the Effects of Operation of the Bureau of Reclamation's Klamath Project on the Endangered Lost River Sucker (Deltistes luxatus), Endangered Shortnose Sucker (Chasmistes brevirostris), Threatened Bald Eagle (Haliaeetus leucocephalus), and Proposed Critical Habitat for the Lost River/Shortnose suckers. Klamath Falls, OR: Klamath Falls Fish and Wildlife Office. [Online]. Available: http://klamathfallsfwo.fws.gov. [August 18, 2001]. Vogel, D.A., K.R.Marine, and A.J.Horne. 2001. Protecting the Beneficial Uses of Waters of the Upper Klamath Lake: A Plan to Accelerate Recovery of the Lost River and Shortnose Suckers. Prepared for the Klamath Water Users Association, Klamath Falls, OR. March 2001. Walker, W.W. 2001. Development of a Phosphorus TMDL for Upper Klamath Lake, Oregon. Oregon Department of Environmental Quality. March 7, 2001. Wedemeyer, G.A., R.L.Saunders, and W.C.Clarke. 1980. Environmental factors affecting smoltification and early marine survival of anadromous salmonids. Mar. Fish. Rev. 42(6): 1–14. Weitkamp, L.A., T.C.Wainwright, G.J.Bryant, G.B.Milner, D.J.Teel, R.G.Kope, and R.S.Waples. 1995. Status Review of Coho Salmon from Washington, Oregon, and California. NOAA Technical Memorandum NMFS-NWFSC-24. Seattle, WA: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Northwest Fishers Science Center. 258 pp. [Online]. Available: http:// www.nwfsc.noaa.gov/pubs/tm/tm24/ tm24.htm. [December 13, 2001]. Welch, E.B., and T.Burke. 2001. Interim Summary Report: Relationship Between Lake Elevation and Water Quality in Upper Klamath Lake, Oregon. Prepared by R2 Resource Consultants, Inc., Redmond, WA, for the Bureau of Indian Affairs, Portland, OR. March 23, 2001. Welsh, H.H., G.R.Hodgson, B.C.Harvey, and M.F.Roche. 2001. Distribution of juvenile coho salmon in relation to water temperatures in tributaries of the Mattole River, California. N.Am.J.Fish Manage. 21(3):464–470.

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REFERENCES 31

Yurok Tribe. 2001. Letter to Irma Lagomarsino, NMFS, from Troy Fletcher regarding preliminary Yurok Tribe comments on draft biological opinion on ongoing Klamath Project operations. March 23, 2001.

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APPENDIX 32

APPENDIX STATEMENT OF TASK

The committee will review the government's biological opinions regarding the effects of Klamath Project operations on species in the Klamath River Basin listed under the Endangered Species Act, including coho salmon and shortnose and Lost River suckers. The committee will assess whether the biological opinions are consistent with the available scientific information. It will consider hydrologic and other environmental parameters (including water quality and habitat availability) affecting those species at critical times in their life cycles, the probable consequences to them of not realizing those environmental parameters, and the inter-relationship of these environmental conditions necessary to recover and sustain the listed species. To complete its charge, the committee will perform the following:

1. Review and evaluate the science underlying the Biological Assessments (Reclamation 2001) and Biological Opinions (USFWS 2001; NMFS 2001). 2 . Review and evaluate environmental parameters critical to the survival and recovery of listed species. 3 . Identify scientific information relevant to evaluating the effects of project operations that has become available since USFWS and NMFS prepared the biological opinions. 4 . Identify gaps in the knowledge and scientific information that are needed to develop comprehensive strategies for recovering listed species and provide an estimate of the time and funding it would require.

Copyright National Academy of Sciences. All rights reserved. EXHIBIT 2, Page 55 of 56 CaseScientific 3:18-cv-03078-WHO Evaluation of Biological Opinions Document on Endangered 50-5 and Threatened Filed 07/03/18Fishes in the Klamath Page River 56 ... of 56

APPENDIX 33

A brief interim report will be provided by January 31, 2002. The interim report will focus on the February 2001 biological assessments of the Bureau of Reclamation and the April 2001 biological opinions of the U.S.Fish and Wildlife Service and National Marine Fisheries Service regarding the effects of operations of the Bureau of Reclamation's Klamath Project on listed species. The committee will provide a preliminary assessment of the scientific information used by the Bureau of Reclamation, the Fish and Wildlife Service, and the National Marine Fisheries Service, as cited in those documents, and will consider to what degree the analysis of effects in the biological opinions of the Fish and Wildlife Service and National Marine Fisheries Service is consistent with that scientific information. The committee will identify any relevant scientific information it is aware of that has become available since the Fish and Wildlife Service and National Marine Fisheries Service prepared the biological opinions. The committee will also consider any other relevant scientific information of which it is aware. The final report, due March 30, 2003, will thoroughly address the scientific aspects related to the continued survival of coho salmon and shortnose and Lost River suckers in the Klamath River Basin. The committee will identify gaps in the knowledge and scientific information that are needed and provide approximate estimates of the time and funding needed to fill those gaps, if such estimates are possible. The committee will also provide an assessment of scientific considerations relevant to strategies for promoting the recovery of listed species in the Klamath Basin.

Copyright National Academy of Sciences. All rights reserved. EXHIBIT 2, Page 56 of 56 Case 3:18-cv-03078-WHO Document 50-6 Filed 07/03/18 Page 1 of 9

Transactions of the American Fisheries Society 128:953±961, 1999 American Fisheries Society 1999

NOTES Effects of Ambient Water Quality on the Endangered Lost River Sucker in Upper Klamath Lake, Oregon

BARBARA A. MARTIN AND MICHAEL K. SAIKI* U.S. Geological Survey, Biological Resources Division, Western Fisheries Research Center, Dixon Duty Station, 6924 Tremont Road, Dixon, California 95620, USA

Abstract.ÐPopulations of the Lost River sucker Del- operations processed ``enormous amounts'' of tistes luxatus have declined so precipitously in the Upper suckers into oil, dried ®sh, and other products (An- Klamath Basin of Oregon and California that this ®sh dreasen 1975). was recently listed for federal protection as an endan- gered species. Although Upper Klamath Lake is a major Over the past few decades, populations of the refuge for this species, ®sh in the lake occasionally ex- Lost River sucker have declined dramatically. Al- perience mass mortalities during summer and early fall. though the decline of this ®sh was recognized as This ®eld study was implemented to determine if ®sh early as the mid-1960s, it was not until 1985 that mortalities resulted from degraded water quality con- the severity of the situation was realized (USFWS ditions associated with seasonal blooms of phytoplank- 1993). Surveys of spawning Lost River suckers in ton, especially Aphanizomenon ¯os-aquae. Our results indicated that ®sh mortality did not always increase as Upper Klamath Lake yielded 23,123 ®sh in 1984 water temperature, pH, and un-ionized ammonia con- but only 11,860 ®sh in 1985 (Bienz and Ziller centration increased in Upper Klamath Lake. Little or 1987). Moreover, the snag ®shery harvest for all no mortality occurred when these water quality variables types of suckers (including Lost River, shortnose attained their maximum values. On the other hand, an Chasmistes brevirostris, and Klamath largescale inverse relation existed between ®sh mortality and dis- Catostomus snyderi) declined from approximately solved oxygen concentration. High mortality (Ͼ90%) occurred whenever dissolved oxygen concentrations de- 10,000 ®sh in 1968 (Golden 1969) to only 687 ®sh creased to 1.05 mg/L, whereas mortality was usually low in 1985 (Bienz and Ziller 1987). A ®sh kill in (Ͻ10%) when dissolved oxygen concentrations equaled Upper Klamath Lake during the summer of 1986 or exceeded 1.58 mg/L. Stepwise logistic regression also further reduced the populations of suckers and ap- indicated that the minimum concentration of dissolved parently eliminated many large adults (Buettner oxygen measured was the single most important deter- minant of ®sh mortality. and Scoppettone 1990). In 1988, these documented instances of population decline created a situation so dire that for purposes of federal protection, the The Lost River sucker Deltistes luxatus is en- Lost River sucker was listed as an endangered spe- demic to the Upper Klamath Basin of southern cies under the federal Endangered Species Act Oregon and northern California, including Upper (USFWS 1988). Klamath Lake and its tributaries, the Lost River The ®nal ruling that listed the Lost River sucker system, Tule Lake, Lower Klamath Lake, and as an endangered species offered the following Sheepy Lake (Moyle 1976). Historically, this ®sh reasons for its decline: damming of rivers, dredg- was widespread and abundant within the Upper ing and draining of marshes, water diversions, hy- Klamath Basin. Gilbert (1898) noted that the Lost bridization, competition with and predation by ex- River sucker was ``the most important food-®sh of otic species, insularization of habitat, and water the Klamath Lake region.'' At that time, spring quality problems associated with timber harvest, runs of this sucker were relied upon as a major removal of riparian vegetation, livestock grazing, food source by Klamath and Modoc Indians, and and agricultural practices (USFWS 1988). Over- the sucker was taken by local settlers for both hu- harvest and chemical contamination may also have man consumption and livestock feed (Cope 1879; contributed to the decline of the sucker. In addi- Coots 1965; Howe 1968). Suckers were so plen- tion, a shift toward hypereutrophication has been tiful that a cannery was established on the Lost River (Howe 1968), and several other commercial documented in Upper Klamath Lake (Miller and Tash 1967; Vincent 1968). During the warmer months of the year, massive blooms of phytoplank- * Corresponding author: michael࿞[email protected] ton, especially the cyanobacterium Aphanizome- Received March 16, 1998; accepted November 11, 1998 non ¯os-aquae, contribute to broad diurnal ¯uc- 953

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954 MARTIN AND SAIKI

FIGURE 1.ÐMap of the study area showing the locations of the eight sample sites. tuations in pH, dissolved oxygen, and other water is associated with high water temperature, high quality variables. Immediately following blooms, pH, high un-ionized ammonia concentration, low decaying phytoplankton and other organic matter dissolved oxygen concentration, or a combination can temporarily increase the concentration of un- of these variables. ionized ammonia and can reduce the concentration of dissolved oxygen to near zero. Because the Lost Description of the Study Area River sucker is a lake-dwelling ®sh that spawns Upper Klamath Lake, including the adjoining in tributary streams or springs, the U.S. Fish and Agency Lake, is located in Klamath County, Or- Wildlife Service (USFWS; 1988) suggested that egon, adjacent to the northern limits of Klamath reduction and degradation of lake and stream hab- Falls (Figure 1). The lake has a mean summer sur- itats are major factors responsible for the decline face area of 27,811 ha, a summer volume of 6.96 of this species. ϫ 108 m3, and a depth of 2.4 m (USACE 1978). The purpose of this study was to determine if The water column is intermittently strati®ed dur- ambient water quality conditions in Upper Klam- ing summer whenever winds are calm for several ath Lake during summer and early fall can be lethal days. The lake is usually ice covered during the to juveniles of the Lost River sucker. This study winter. sought to achieve the following objectives: (1) to Summertime water temperatures in Upper determine whether ®sh mortality varies among Klamath Lake can approach 30ЊC near the surface, sites, test dates (tests), or both, within Upper and temperatures of 22±24ЊC are common in the Klamath Lake and (2) to determine if ®sh mortality upper 1±2 m of water (Bortleson and Fretwell

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NOTES 955

1993). At the same time, dissolved oxygen con- exception occurred during the ®rst four tests with centrations are often photosynthetically supersat- the hatchery control, when only one cage was urated in the upper part of the water column, but available. concentrations of less than 2.0 mg/L may occur At each sampling site, live cages were suspend- near the bottom. Occasionally, dissolved oxygen ed from styrofoam buoys with 1-m lengths of gal- concentration can drop to as low as 0.2 mg/L vanized cable. The styrofoam buoys were attached throughout the water column in response to a com- to a large air-®lled main buoy which, in turn, was bination of high temperature, algal senescence, attached by a cable to a 45.4-kg cement anchor. A and stagnant water (no mixing of the water column Hydrolab Datasonde 1 multiprobe logger (Hydro- by wind). During periods of peak algal productiv- lab Corporation, Austin, Texas) was attached to ity, pH values commonly reach 9 or 10 and may the cable of the main buoy so that the probes were even exceed 10. Although recent measurements of at a depth of 1 m. In order to prevent the probes ammonia are not available, concentrations during from ¯oating, the Hydrolab probe protector was the winter of 1981±1982 averaged 1±2 mg/L as attached to the cable with a plastic cable tie. ammonia nitrogen (1.2±2.4 mg/L as total ammo- Test ®sh were obtained from the Braymill nia). Hatchery and transported to sampling sites in an aerated 45.4-L insulated ice chest. After discarding Methods deformed or unhealthy ®sh, 10 juveniles of similar Selection of ®eld sites.ÐEight 4-d-long tests size (mean Ϯ SD, 0.22 Ϯ 0.19 g) were selected at were performed during summer 1995 at eight sites random and placed into each cage. Test ®sh sub- in Upper Klamath Lake (Figure 1). The tests oc- sisted on food products that passed through the curred on the following days: June 20±24 (test 1), screens. June 25±29 (test 2), July 3±7 (test 3), July 7±11 During each test, live cages were monitored dai- (test 4), August 14±18 (test 5), August 28±Sep- ly, and dead ®sh were removed and debris was tember 1 (test 6), September 4±9 (test 7), and Sep- cleaned from screens. Dead ®sh were counted, tember 11±15 (test 8). In addition, a ``hatchery weighed, measured, and then preserved in 70% control'' was maintained at the Klamath Tribe's ethanol. After each test was terminated, all sur- Braymill Hatchery (near Chiloquin, Oregon) in or- viving ®sh were weighed, measured, and preserved der to monitor the quality of different batches of in 70% ethanol. ®sh used in the tests. Water quality measurements.ÐEach Hydrolab Field sites and their geospatial coordinates were Datasonde 1 was calibrated and programmed to as follows: (1) Rocky Point, 42Њ28.44ЈNby take hourly measurements of pH, dissolved oxy- 122Њ05.09ЈW; (2) inlet from Agency Lake (Agen- gen, and water temperature. The accuracy of Hy- cy), 42Њ29.78ЈN by 122Њ00.37ЈW; (3) mouth of drolab measurements was veri®ed by comparing Odessa Creek (Odessa), 42Њ26.12ЈNby 122Њ02.81ЈW; (4) mouth of Williamson River these measurements with daily grab-sample mea- (Williamson), 42Њ27.75ЈN by 121Њ57.33ЈW; (5) surements from a YSI model 57 temperature-dis- Ball Bay (Ball), 42Њ25.07ЈN by 122Њ00.18ЈW; (6) solved oxygen meter (Yellow Springs Instrument Shoalwater Bay (Shoalwater), 42Њ24.10ЈNby Company, Yellow Springs, Ohio) and a Cole- 121Њ57.30ЈW; (7) Hagelstein Park (Hagelstein), Parmer model 5994 Digisense pH meter (Cole- 42Њ23.00ЈNby121Њ48.95ЈW; and (8) Howard Bay Parmer Instrument Company, Chicago, Illinois). (Howard), 42Њ18.72ЈN by 121Њ56.69ЈW. Results Total ammonia was monitored by collecting 1-L from long-term hydrological monitoring studies (J. grab samples of water from each site. Samples were Kann, aquatic ecologist, Ashland, Oregon, unpub- vacuum ®ltered through a 1.6-␮m binder-free glass- lished data) were used to guide the selection of ®ber pre®lter, triple ®ltered through a 0.4-␮m±pore sites so as to encompass a range of water quality size polycarbonate track-etched membrane ®lter, conditions. and then stored on ice. A Nalgene polyvinyl chlo- Experimental setup and procedure.ÐWith one ride vacuum hand pump was used to create the vac- exception, each ®eld site and the hatchery control uum. In the laboratory, samples were analyzed for contained three live cages constructed from 18.9- total dissolved ammonia with an Orion SA 270 con- L plastic buckets that were modi®ed to allow water centration meter and an Orion No. 951201 ammonia circulation (by cutting two 34-cm ϫ 23±cm open- gas-sensing electrode (Orion Research, Inc., Cam- ings in the sides and then by covering the openings bridge, Massachusetts). Total ammonia concentra- with 0.043-cm±mesh stainless-steel screen). The tions were converted to un-ionized ammonia con-

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956 MARTIN AND SAIKI

TABLE 1.ÐCumulative mortality (%) of juvenile Lost River suckers held for 4 days in live cages at various sites within Upper Klamath Lake during eight separate tests. Values are angular-transformed means (ranges in parentheses).

Testa Site 12345678 Agency 0.0 0.0 16.4 0.0 13.0 1.1 0.0 0.0 (0±0) (0±0) (0±50) (0±0) (10±20) (0±10) (0±0) (0±0) Ball 0.0 1.1 1.1 0.0 0.0 4.5 4.5 1.1 (0±0) (0±10) (0±10) (0±0) (0±0) (0±10) (0±10) (0±10) Hagelstein 1.1 0.0 9.3 10.0 0.0 0.0 0.0 1.1 (0±10) (0±0) (0±20) (10±10) (0±0) (0±0) (0±0) (0±10) Howard 0.0 0.0 100.0 100.0 0.0 1.1 100.0 100.0 (0±0) (0±0) (100±100) (100±100) (0±0) (0±10) (100±100) (100±100) Odessa 0.0 10.0 52.1 9.3 1.1 98.9 100.0 1.1 (0±0) (10±10) (10±80) (0±20) (0±10) (90±100) (100±100) (0±10) Rocky Point 0.0 1.1 0.0 1.1 0.0 0.0 0.0 2.4 (0±0) (0±10) (0±0) (0±10) (0±0) (0±0) (0±0) (0±20) Shoalwater 5.1 0.0 1.1 1.1 0.0 0.0 0.0 ND (0±40) (0±0) (0±10) (0±10) (0±0) (0±0) (0±0) Williamson 5.1 ND 1.1 10.0 0.0 1.1 0.0 0.0 (0±40) (0±10) (10±10) (0±0) (0±10) (0±0) (0±0) a ND ϭ no data. centrations by using the formula described in Em- large Wald chi-square statistic is more important erson et al. (1975). than are other independent variables represented Statistical comparisons.ÐResults were summa- by smaller Wald chi-square statistics. rized by analysis of variance, Pearson's product- moment correlation, and stepwise logistic regres- Results sion using the SAS statistical package (SAS Insti- Fish Mortality tute, Cary, North Carolina). Raw data were either Cumulative mortality varied over sites and tests, logarithmically transformed or, if expressed in per- with some mortality occurring at all sites during centages, angular transformed before undergoing one or more of the tests (Table 1). Complete mor- statistical summarization and comparisons. Null hy- tality occurred only at Howard and Odessa, where- potheses of statistical comparisons were rejected if as relatively low mortality (Յ5.1%) was recorded P was less than or equal to 0.05. at Ball, Rocky Point, and Shoalwater. Juvenile For comparisons employing Pearson's correla- suckers at the remaining sites (Agency, Hagelstein, tions and stepwise logistic regression where ®sh and Williamson) experienced moderate amounts of mortality was the dependent variable, only the fol- mortality (10.0±16.4%) on at least one occasion. lowing water quality (independent) variables were Mortality did not occur among the hatchery con- considered: maximum temperature, maximum pH, trols. minimum dissolved oxygen concentration, and maximum un-ionized ammonia concentration. Water Quality Available studies indicated that ®sh mortality in Pearson product-moment correlations indicated Upper Klamath Lake was likely to be in¯uenced that all water quality variables (temperature, dis- by these water quality variables (Andreasen 1975; solved oxygen, pH, and un-ionized ammonia) were Coleman et al. 1988; Falter and Cech 1991; Bor- signi®cantly intercorrelated (P Յ 0.05). With one tleson and Fretwell 1993; Castleberry and Cech exception, intercorrelations resulted from diurnal 1993; Wood et al. 1996; Saiki et al. 1999). In step- patterns. Un-ionized ammonia was exceptional be- wise logistic regression, the Wald chi-square sta- cause it was measured only once daily, so a diurnal tistic from the last step of the stepwise algorithm pattern was not identi®ed. is proportional to the amount of explanatory power Temperature.ÐWater temperatures varied from that would be lost by removing a given variable 7.86ЊC at Rocky Point to 26.65ЊC at Odessa (Table from the model (N. H. Willits, Division of Statis- 2). Signi®cant two-way interaction (F46,5593 ϭ tics, University of California, personal commu- 88.87; P Ͻ 0.0001) indicated that thermal patterns nication). Thus, if ®sh mortality is the dependent were not consistent among sites and tests. variable in a logistic regression model, a water Dissolved oxygen concentration.ÐDissolved ox- quality (independent) variable represented by a ygen concentrations varied from 0 mg/L at Howard

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NOTES 957

TABLE 2.ÐSummary of water temperature, dissolved oxygen concentration, pH, and un-ionized ammonia concentra- tion during eight tests at various sites in Upper Klamath Lake. Values are geometric means (ranges in parentheses).

Dissolved oxygen Un-ionized Temperature concentration ammonia Site (ЊC) (mg/L) pH (mg/L) Agency 19.72 7.70 9.41 0.04894 (14.57±24.71) (2.60±20.00) (8.39±10.78) (0.00892±0.18457) Ball 19.58 8.49 9.11 0.06028 (14.36±26.19) (1.94±20.00) (8.51±9.96) (0.00349±0.37635) Hagelstein 15.47 11.64 8.81 0.00395 (10.98±24.63) (5.90±20.00) (7.39±9.79) (0.00034±0.01723) Howard 20.37 8.26 9.42 0.06823 (14.91±26.23) (0.00±20.00) (6.46±10.55) (0.01058±0.65179) Odessa 18.04 6.59 8.08 0.01637 (12.88±26.65) (0.34±20.00) (6.09±10.60) (0.00003±0.14445) Rocky Point 15.76 9.50 7.16 0.00011 (7.86±24.63) (6.32±12.97) (6.57±8.64) (0.00004±0.00053) Shoalwater 19.53 9.55 8.90 0.03531 (17.11±23.44) (3.55±20.00) (8.46±10.24) (0.00535±0.24430) Williamson 17.97 10.79 8.58 0.00404 (13.73±24.37) (7.22±17.47) (7.32±9.86) (0.00040±0.16163) to 20 mg/L (the maximum concentration measur- Agency (Table 2). Moreover, on at least one oc- able with the Hydrolab Datasonde 1) at Agency, casion, pH values temporarily exceeded 10.5 at Ball, Hagelstein, Howard, Rocky Point, and Shoal- Agency, Odessa, and Howard. Signi®cant two-way water (Table 2). On several occasions, dissolved interaction (F46,5593 ϭ 123.81; P Ͻ 0.0001) indi- oxygen concentrations declined to less than or cated that pH did not exhibit a consistent pattern equal to 1.05 mg/L at both Howard and Odessa. among sites and tests.

However, signi®cant two-way interaction (F46,5591 Un-ionized ammonia concentration.ÐUn-ionized ϭ 50.95; P Ͻ 0.0001) indicated that dissolved ox- ammonia concentrations varied from 0.00003 mg/ ygen patterns were not consistent among sites and L at Odessa to nearly 0.652 mg/L at Howard (Table tests. 2). Signi®cant two-way interaction (F46,234 ϭ 4.08; Hydrogen-ion concentration.ÐExtreme values P Ͻ 0.0001) indicated this variable did not exhibit for pH varied from 6.09 at Odessa to 10.78 at a consistent pattern among sites and tests.

TABLE 3.ÐRelation of selected water quality variables Relation of Fish Mortality to Selected Water to mean mortality of ®sh caged for four days in Upper Quality Variables Klamath Lake; N, sample size; r, Pearson correlation co- ef®cient; P, probability. According to an adjusted Bonferroni ␣ of 0.004 (to account for 12 simultaneous comparisons), Mean mortality minimum and mean dissolved oxygen concentra- Water quality variable Na rPtions were signi®cantly correlated with ®sh mor- Maximum temperature 60 0.199 0.1274 tality (Table 3). However, if coef®cient of deter- Mean temperature 60 0.224 0.0857 mination (r2) values are considered, only minimum Minimum temperature 60 0.199 0.1278 dissolved oxygen concentration (r2 ϭ 60.1%) ac- Maximum dissolved oxygen counted for more than one-half of the variation in concentration 60 0.244 0.0604 Mean dissolved oxygen ®sh mortality. concentration 60 Ϫ0.617 Ͻ0.0001 When three-dimensional (3-D) plots of the data Minimum dissolved oxygen were examined, mortality of caged ®sh exhibited concentration 60 Ϫ0.775 Ͻ0.0001 Maximum pH 60 0.196 0.1333 little or no relationship with maximum tempera- Mean pH 60 Ϫ0.025 0.8519 ture, maximum pH, and maximum un-ionized am- Minimum pH 60 Ϫ0.255 0.0495 monia concentration (Figure 2a, 2c). Although Maximum un-ionized ammonia 60 0.334 0.0092 Mean un-ionized ammonia 60 0.298 0.0209 some tests exhibited high (Ͼ90%) ®sh mortality Minimum un-ionized ammonia 60 0.187 0.1520 at temperatures of 20.4±25.0ЊC, pHs of 9.4±10.5, a Data on ®sh mortality and water quality were not available during and un-ionized ammonia concentrations of 0.140± Test 2 at Williamson and Tests 1, 2, and 8 at Shoalwater. 0.370 mg/L, other tests exhibited little or no ®sh

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958 MARTIN AND SAIKI

FIGURE 2.ÐThree-dimensional plots of mean mortality in juvenile Lost River suckers exposed for4dtoselected water quality variables in Upper Klamath Lake; (a) minimum dissolved oxygen concentration and maximum pH; (b) minimum dissolved oxygen concentration and maximum temperature; and (c) minimum dissolved oxygen concentration and maximum un-ionized ammonia concentration. Dashed lines represent a dissolved oxygen con- centration of 1.58 mg/L. mortality under similar or more extreme condi- 1.58 mg/L. One exceptional event occurred when tions. By comparison, the 3-D plots indicated that ®sh mortality averaging 52% coincided with a high mortality (Ͼ90%) of caged ®sh occurred minimum dissolved oxygen concentration of 2.35 whenever dissolved oxygen concentrations mg/L. dropped to less than or equal to 1.05 mg/L, where- Stepwise logistic regression was used to further as mortality was usually low (Ͻ10%) when dis- assess the relationship between ®sh mortality and solved oxygen concentrations equaled or exceeded water quality variables. When main effects were

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NOTES 959

TABLE 4.ÐRelative importance of selected water qual- erage 30.51ЊC (95% con®dence interval, 29.99± ity variables as determinants of mortality in juvenile Lost 31.04ЊC). By comparison, the highest temperature River suckers caged in Upper Klamath Lake for four days. measured in Upper Klamath Lake during our study Wald ␹ 2 scores are computed from stepwise logistic re- gression.a was 26.65ЊC (Table 2). Moreover, Saiki et al. (1999) indicated that 96-h LC50 values for un- Variable Wald ␹ 2 scores ionized ammonia averaged 0.78 mg/L (95% con- Minimum dissolved oxygen (DO) 95.86*** ®dence interval, 0.70±0.86 mg/L). This lethal con- Maximum pH (pH) 33.21*** centration exceeded the maximum of 0.652 mg/L b Maximum un-ionized ammonia (NH3) 0.51 measured in lake water during our study (Table 2). Maximum temperature (Temp) 11.28*** DO*pH 56.69*** Collectively, these ®ndings indicate that water DO*NH3 37.23*** temperatures and un-ionized ammonia concentra- DO * Temp c tions in Upper Klamath Lake were not directly pH*NH3 26.59*** pH * Temp c responsible for the occasional episodes of high ®sh NH3 * Temp 6.72** mortality that we documented in our live cages. DO*pH*NH3 53.06*** Falter and Cech (1991) determined that the crit- DO*pH*Temp c c ical pH maximum for juvenile Lost River suckers DO*NH3 * Temp c pH*NH3 * Temp was 9.55 Ϯ 0.43 (mean Ϯ SD), whereas Saiki et DO*pH*NH3 * Temp 17.67*** al. (1999) indicated that 96-h LC50 values for this a Codes: * P Յ 0.05; ** P Յ 0.01; *** P Յ 0.001. species and life stage averaged 10.30 (95% con- b Variable was forced into the logistic regression model to enable ®dence interval, 9.94±10.67). During our study, analysis of higher-level interactions. pH values approaching or exceeding these labo- c Variable did not meet the entry criterion of P Յ 0.05. ratory-derived tolerance limits were recorded on several occasions in Upper Klamath Lake (Table examined, the large Wald chi-square statistic as- 2). However, when pH averaged 10.30 and peaked sociated with minimum dissolved oxygen concen- at 10.78 during Test 1 at Agency, there were no tration indicated that this variable was consistently mortalities among our caged ®sh (Table 1). These the single most important determinant of ®sh mor- ®ndings suggest either that the high pH was not tality (Table 4). Certain combinations of maximum sustained long enough to kill ®sh or that other pH, maximum un-ionized ammonia concentration, environmental conditions may have ameliorated maximum temperature, and minimum dissolved the lethal effects of high pH. oxygen concentration also contributed to ®sh mor- Saiki et al. (1999) reported that juvenile Lost tality, but they did so to a lesser degree than did River suckers succumbed to hypoxia at 96-h LC50 minimum dissolved oxygen concentration alone values that averaged 1.62 mg/L (95% con®dence (Table 4). interval, 1.41±1.86 mg/L). During our study, dis- solved oxygen concentrations in Upper Klamath Discussion Lake fell below 1.62 mg/L on several occasions Several laboratory studies have examined the at Howard and Odessa (Table 2). However, high tolerances of the Lost River sucker and related mortalities (Ͼ90%) occurred in caged ®sh from species (Klamath largescale sucker and shortnose these sites only when dissolved oxygen concen- sucker) in Upper Klamath Lake to high water tem- trations declined to 1.05 mg/L or less (Table 1; perature (Castleberry and Cech 1993; Saiki et al. Figure 2). The high mortalities also coincided with 1999), low dissolved oxygen concentration (Cas- the occurrence of senescent algae that imparted a tleberry and Cech 1993; Saiki et al. 1999), high yellow±brown color to lake water (B. A. Martin, pH (Falter and Cech 1991; Saiki et al. 1999), and unpublished observations). By comparison, only a high un-ionized ammonia concentration (Saiki et few ®sh died (9.3% mortality) on the one occasion al. 1999). Unlike these laboratory studies, the pres- when minimum dissolved oxygen concentration ent study exposed juvenile Lost River suckers to equaled 1.58 mg/L (Figure 2). Although some vari- ambient summertime conditions in the lake in or- ation was present, ®sh typically experienced little der to determine whether one or more of these or no mortality as minimum dissolved oxygen con- variables could account for occasional ®sh kills. centrations approached or exceeded 2.0 mg/L. According to Saiki et al. (1999), the 96-h median (Note: One exceptional event occurred during test lethal concentrations (LC50 values; also referred 3 at Odessa when 52% mortality coincided with a to as median tolerance limits) of juvenile Lost Riv- minimum dissolved oxygen concentration of 2.35 er suckers exposed to high water temperature av- mg/L.) Overall, these observations suggest that

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960 MARTIN AND SAIKI mortality of caged juvenile Lost River suckers in Acknowledgments Upper Klamath Lake resulted from hypoxia as- We thank B. Bellerud, D. Follansbee, and L. sociated with a dissolved oxygen de®cit. These Whitley for assisting with ®eld work; M. Buettner, ®ndings are supported by results from stepwise L. Dunsmoor, and J. Kann for providing technical logistic regression that identi®ed minimum dis- and logistical support; and K. Brenneman, R. Fritz- solved oxygen concentration as the single most sche, and T. Hassler for reviewing an early version important water quality variable in¯uencing ®sh of this manuscript. The Klamath Tribe is acknowl- mortality (Table 4). edged for providing the test animals used in this Although less important than minimum dis- study. This manuscript is based on a thesis sub- solved oxygen concentration, our results from mitted in partial ful®llment of a MS degree by the stepwise logistic regression indicated that inter- senior author to the faculty of Humboldt State Uni- actions among several combinations of water qual- versity. ity variables were signi®cantly associated with mortality of caged ®sh (Table 4). Other investi- References gators reported that low concentrations of dis- Alabaster, J. S., D. G. Shurben, and G. Knowles. 1979. solved oxygen could reduce the tolerance of ®sh The effect of dissolved oxygen and salinity on the to elevated un-ionized ammonia concentrations toxicity of ammonia to smolts of salmon, Salmo (Downing and Merkens 1955; Merkens and Down- salar L. Journal of Fish Biology 15:705±712. ing 1957; Alabaster et al. 1979; Thurston et al. Andreasen, J. K. 1975. Systematics and status of the family Catostomidae in southern Oregon. Doctoral 1981). Moreover, water temperature can in¯uence dissertation. Oregon State University, Corvallis. the onset of hypoxia in ®sh, with higher temper- Bienz, C. S., and J. S. Ziller. 1987. Status of three la- atures generally increasing the lower lethal thresh- custrine sucker species (Catostomidae). U.S. Fish old for dissolved oxygen concentration (Heath and Wildlife Service, Completion Report, Sacra- 1987). Although not identi®ed as a cause of ®sh mento, California. mortality, Serafy and Harrell (1993) demonstrated Bortleson, G. C., and M. O. Fretwell. 1993. A review of possible causes of nutrient enrichment and de- that supersaturated dissolved oxygen conditions cline of endangered sucker populations in upper allowed banded killi®sh Fundulus diaphanus, blue- Klamath Lake, Oregon. U.S. Geological Survey, gill Lepomis macrochirus, and striped bass Morone Water-Resources Investigations Report 93±4087, saxatilis to tolerate high pH levels that would oth- Portland, Oregon. erwise be lethal to these species. Serafy and Har- Buettner, M., and G. Scoppettone. 1990. Life history rell suggested that high pH reduced the ability of and status of catostomids in upper Klamath Lake, Oregon. U.S. Fish and Wildlife Service, National ®sh to transpire oxygen and that supersaturated Fisheries Research Center, Completion Report, Se- dissolved oxygen conditions helped to nullify this attle. effect. Castleberry, D. T., and J. J. Cech, Jr. 1993. Critical Although beyond the scope of our study, re- thermal maxima and oxygen minima of ®ve ®shes source managers have hypothesized that algal tox- from the upper Klamath basin. California Fish and ins might also contribute to mortality of the Lost Game 78:145±152. River sucker (Bortleson and Fretwell 1993; Wood Coleman, M. E., J. Kann, and G. G. Scoppettone. 1988. Life history and ecological investigations of catos- et al. 1996). Under certain nutrient conditions, A. tomids from the upper Klamath basin, Oregon. U.S. ¯os-aquae can produce a toxic strain that has Fish and Wildlife Service, National Fisheries Re- caused ®sh kills in other waters (English et al. search Center, Annual Report, Seattle. 1993). However, the toxic strain has not been re- Coots, M. 1965. Occurrences of Lost River sucker, Del- ported from Upper Klamath Lake (J. Kann, aquatic tistes luxatus (Cope), and shortnose sucker, Chas- ecologist, Ashland, Oregon, personal communi- mistes brevirostris (Cope), in northern California. California Fish and Game 51:68±73. cation). Moreover, Microcystis aeruginosa (anoth- Cope, E. D. 1879. The ®shes of Klamath Lake, Oregon. er toxin-producing cyanobacterium implicated in American Naturalist 13:784±785. ®sh kills) occurred for the ®rst time at high den- Downing, K. M., and J. C. Merkens. 1955. The in¯uence sities in Upper Klamath Lake during the summer of dissolved-oxygen concentration on the toxicity of 1996 (Kann, personal communication). Clearly, of un-ionized ammonia to rainbow trout (Salmo much still remains to be learned about the envi- gairdnerii Richardson). Annals of Applied Biology 43:243±246. ronmental variables that can affect long-term sur- Emerson, K., R. C. Russo, R. E. Lund, and R. V. Thurs- vival of Lost River sucker populations in Upper ton. 1975. Aqueous ammonia equilibrium calcu- Klamath Lake and elsewhere in the Upper Klamath lations: effect of pH and temperature. Journal of the Basin. Fisheries Research Board of Canada 32:2379±2383.

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NOTES 961

English, W. R., T. E. Schwedler, and L. A. Dyck. 1993. larval and juvenile Lost River and shortnose suck- Aphanizomenon ¯os-aquae, a toxic blue-green alga ers. Environmental Pollution 105:37±44. in commercial channel cat®sh, Ictalurus punctatus, Serafy, J. E., and R. M. Harrell. 1993. Behavioural re- ponds: a case history. Journal of Applied Aquacul- sponse of ®shes to increasing pH and dissolved ox- ture 3:195±209. ygen: ®eld and laboratory observations. Freshwater Falter, M. A., and J. J. Cech, Jr. 1991. Maximum pH Biology 30:53±61. tolerance of three Klamath basin ®shes. Copeia Thurston, R. V., G. R. Phillips, R. C. Russo, and S. M. 1991:1109±1111. Hinkins. 1981. Increased toxicity of ammonia to rainbow trout (Salmo gairdneri) resulting from re- Gilbert, C. H. 1898. The ®shes of the Klamath basin. duced concentrations of dissolved oxygen. Cana- U.S. Fish Commission Bulletin 17:1±13. dian Journal of Fisheries and Aquatic Sciences 38: Golden, M. P. 1969. The Lost River sucker, Catostomus 983±988. luxatus (Cope). Oregon State Game Commission, USACE (U.S. Army Corps of Engineers). 1978. Klam- Central Region, Administrative Report 1-69, Port- ath River basin, Oregon, reconnaissance report. land. USACE, San Francisco District, San Francisco. Heath, A. G. 1987. Water pollution and ®sh physiology. USFWS (U.S. Fish and Wildlife Service). 1988. En- CRC Press, Boca Raton, Florida. dangered and threatened wildlife and plants; short- Howe, C. B. 1968. Ancient tribes of the Klamath Coun- nose and Lost River suckers, Houghton's goldenrod try. Binford and Mort, Portland, Oregon. and Pitcher's thistle; ®nal rules. U.S. Of®ce of the Merkens, J. C., and K. M. Downing. 1957. The effect Federal Register 53(18 July 1988):27130±27134. of tension of dissolved oxygen on the toxicity of USFWS (U.S. Fish and Wildlife Service). 1993. Lost un-ionized ammonia to several species of ®sh. An- River (Deltistes luxatus) and shortnose (Chasmistes nals of Applied Biology 45:521±527. brevirostris) sucker recovery plan. USFWS, Port- Miller, W. E., and J. C. Tash. 1967. Interim report, upper land, Oregon. Vincent, D. T. 1968. The in¯uence of some environ- Klamath Lake studies, Oregon. Federal Water Pol- mental factors on the distribution of ®shes in upper lution Control Administration, Paci®c Northwest Klamath Lake. Master's thesis. Oregon State Uni- Water Laboratory, Water Pollution Control Re- versity, Corvallis. search Series WP-20-8, Corvallis, Oregon. Wood, T. M., G. J. Fuhrer, and J. L. Morace. 1996. Re- Moyle, P. B. 1976. Inland ®shes of California. Univer- lation between selected water-quality variables and sity of California Press, Berkeley. lake level in upper Klamath and Agency lakes, Or- Saiki, M. K., D. P. Monda, and B. L. Bellerud. 1999. egon. U.S. Geological Survey, Water-Resources Re- Lethal levels of selected water quality variables to port 96-4079, Portland, Oregon.

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ENVIRONMENTAL POLLUTION

Environmental Pollution 105 (1999) 37±44

Lethal levels of selected water quality variables to larval and juvenile Lost River and shortnose suckers

M.K. Saiki*, D.P. Monda1, B.L. Bellerud2 US Geological Survey, Biological Resources Division, Western Fisheries Research Center-Dixon Duty Station, 6924 Tremont Road, Dixon, CA 95620, USA

Received 19 March 1998; accepted 16 November 1998

Abstract Resource managers hypothesize that occasional ®sh kills during summer±early fall in Upper Klamath Lake, Oregon, may be linked to unfavorable water quality conditions created by massive algal blooms. In a preliminary e€ort to address this concern, short-term (96-h-long) laboratory tests were conducted with larval and juvenile Lost River (Deltistes luxatus) and shortnose (Chas- mistes brevirostris) suckers to determine the upper median lethal concentrations (LC50s; also referred to as median tolerance limits) for pH, un-ionized ammonia, and water temperature, and the lower LC50s for dissolved oxygen. The mean LC50s varied among species and life stages as follows: for pH, 10.30±10.39; for un-ionized ammonia, 0.48±1.06 mg litre1; for temperature, 30.35± 31.82C; and for dissolved oxygen, 1.34±2.10 mg litre1. Comparisons of 95% con®dence limits indicated that, on average, the 96-h

LC50s were not signi®cantly di€erent from those computed for shorter exposure times (i.e., 24 h, 48 h, and 72 h). According to two- way analysis of variance, LC50s for the four water quality variables did not vary signi®cantly ( p>0.05) between ®sh species. However, LC50s for pH (exposure times of 24 h and 48 h) and dissolved oxygen (exposure times of 48 h, 72 h, and 96 h) di€ered signi®cantly ( p0.05) between life stages, whereas LC50s for un-ionized ammonia and water temperature did not exhibit signi®cant di€erences. In general, larvae were more sensitive than juveniles to high pH and low dissolved oxygen concentrations. When com- pared to ambient water quality conditions in Upper Klamath Lake, our results strongly suggest that near-anoxic conditions asso- ciated with the senescence phase of algal blooms are most likely to cause high mortalities of larval and juvenile suckers. Published by Elsevier Science Ltd. Keywords: Dissolved oxygen; Water quality; Upper Klamath Lake; Chasmistes brevirostris; Deltistes luxatus

1. Introduction these suckers are more numerous now than in 1984± 1985, young individualsÐespecially the 1991 and 1993 The Lost River sucker (Deltistes luxatus) and the year classesÐdominate adult populations today whereas shortnose sucker (Chasmites brevirostris) are endemic to older individuals were dominant in the earlier surveys the Klamath Basin of northern California and southern (M. Buettner, US Bureau of Reclamation, Klamath Oregon. Field studies indicate that populations of these Falls, Oregon, personal communication). suckers have declined precipitously, with only 11,860 Upper Klamath Lake is the primary refuge for Lost Lost River and 2650 shortnose suckers counted during River and shortnose suckers (Buettner and Scoppettone, surveys of spawning adults in 1984 and 1985 (Bienz and 1990). Over the past several decades, water quality in this Ziller, 1987). On July 18, 1988, the US Fish and Wildlife large (surface area, 27,811 ha), shallow (mean depth, Service listed the two suckers as endangered species. 2.4 m) lake has deteriorated as agricultural activities Although studies conducted in 1995±1997 suggest that increased in the Klamath Basin. During August 1986, mass mortalities of Lost River suckers and smaller num- * Corresponding author. Fax: +1-707-6785039; e-mail: michael_ bers of shortnose suckers were attributed to unsuitable [email protected]. water quality (Scoppettone, 1986). Since then, ®sh mor- 1 Present address: 3700 North First Avenue, Apt. 2008, Tucson, tality events have occurred sporadically throughout the Arizona 85719, USA. 2 Present address: Oregon Department of Fish and Wildlife, Fish lake, with the most recent incident occurring in August± Research and Development, 211 Inlow Hall, Eastern Oregon State September 1997 (J. Kann, aquatic ecology consultant, College, 1410 L Avenue, LaGrande, Oregon 97850, USA. Ashland, Oregon, personal communication). Although

0269-7491/99/$Ðsee front matter Published by Elsevier Science Ltd. PII: S0269-7491(98)00212-7

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38 M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 the precise cause of the die-o€s has not been deter- both with a dry ration (90% krill:10% Spirulina) used by mined, high pH, high water temperatures, high un- the Braymill Fish Hatchery. In tests conducted during ionized ammonia concentrations, and low dissolved 1994, the dry ration was supplemented with live brine oxygen concentrations are possible contributing factors shrimp nauplii. After tests were initiated, larvae were fed (Bortleson and Fretwell, 1993; Wood et al., 1996). ad libitum twice daily whereas juveniles were not fed. The purpose of this study was to determine if extreme Larval tests were initiated when test animals were 35 water quality conditions could explain the occasional days old, whereas juvenile tests were initiated when test die-o€s of Lost River and shortnose suckers that occur animals were 3±7 months old. during summer±early fall in Upper Klamath Lake. Speci®cally, several acute (96-h-long) toxicity tests with 2.1. Experimental design larval and juvenile suckers were conducted to estimate the upper median lethal concentrations (LC50s; also Our tests were designed speci®cally to estimate the referred to as median tolerance limits) for pH, un- upper LC50s for pH, un-ionized ammonia, and water ionized ammonia, and water temperature, and the lower temperature, and the lower LC50s for dissolved oxygen LC50s for dissolved oxygen. during 96-h-long exposure times. Except when manipu- lated upwards or downwards as part of a test, target levels of the four variables were as follows: temperature, 2. Methods 20C; pH, 8.0; dissolved oxygen, 7.6 mg litre1 (100% saturation); and un-ionized ammonia, <0.01 mg litre1. Tests were conducted in 1992±1994 with larval and Each test consisted of ®ve levels of the speci®ed vari- juvenile Lost River and shortnose suckers obtained from able and one control (Table 1). Although test chambers The Klamath Tribe's Braymill Fish Hatchery near were duplicated, the water (test medium) came from a Chiloquin, OR. The ®sh were reared from eggs and common source (i.e., each pair of duplicate test cham- sperm collected during March±June from wild adults bers contained treatments that were not independent captured in the Williamson River or in Sucker and from each other). Thus, data from duplicate treatments

Ouxy springs along the northeastern shore of Upper were pooled when estimating the LC50 values. Klamath Lake. All test animals were transported to our laboratory in Klamath Falls in 45.4-litre insulated ice Table 1 Summary of control and ®ve treatment levels of pH, un-ionized chests aerated with compressed oxygen. ammonia (NH3), water temperature (TEMP), and dissolved oxygen Test animals were acclimated to laboratory condi- (DO) in 96-h tests with Lost River and shortnose suckers tions (water temperature, 20C; photoperiod, 16-h light:8-h dark) in a round 250-litre ®berglass tank for at Variable Control or treatment Mean (95% con®dence interval) least 4 days prior to experimentation. The tank received approximately two changes of reconstituted water per pH Control 7.85 (7.80±7.91) day from a gravity-fed ¯ow-through system. 1 8.75 (8.44±9.07) 2 9.30 (9.02±9.60) Reconstituted water was prepared in batches of 500± 3 9.87 (9.62±10.11) 1800 litre by adding reagent-grade sodium bicarbonate 4 10.57 (10.39±10.76) . (NaHCO3), calcium sulfate dihydrate (CaSO4 2H2O), 5 11.17 (11.04±11.30) . hydrated magnesium sulfate (MgSO4 7H2O), and potas- NH (mg L1) Control 0.0026 (0.0020±0.0033) sium chloride (KCl) to deionized water (city tapwater 3 1 0.075 (0.052±0.11) treated by charcoal ®ltration and reverse osmosis). 2 0.17 (0.14±0.21) Sodium hydroxide (5 M NaOH) was used to adjust the 3 0.36 (0.29±0.46) pH of reconstituted water to about 8.0. This recon- 4 0.86 (0.69±1.1) stituted water mimicked the ionic composition occurring 5 2.3 (1.8±3.1) during springtime in Upper Klamath Lake (M.K. Saiki, TEMP (C) Control 20.28 (19.48±21.12) unpublished data) when the water quality is assumed to 1 25.22 (23.68±26.86) be non-lethal to suckers. Target values for selected water 2 27.36 (26.25±28.52) quality variables in the reconstituted water were as fol- 3 29.14 (28.51±29.77) lows: pH, 8.0‹0.2; calcium hardness, 18.3‹1.8 mg 4 31.01 (30.23±31.81) 1 1 5 33.14 (32.47±33.83) litre as CaCO3; total hardness, 34.7‹2.4 mg litre as 1 CaCO3; total alkalinity, 38‹3.8 mg litre as CaCO3; DO (mg L1) Control 7.94 (7.45±8.47) 1  speci®c conductance, 200‹40 mScm @25C; total 1 3.83 (3.08±4.75) ammonia, <0.01 mg litre1; sulfates, 40‹4 mg litre1; 2 2.77 (2.14±3.57) and chlorides, 2.0‹0.2 mg litre1. 3 2.20 (1.69±2.88) During the 4-day acclimation period, larvae were fed 4 1.56 (1.25±1.94) 5 0.83 (0.64±1.06) three times daily whereas juveniles were fed once daily,

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M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 39

Each 25-litre glass aquarium (test chamber) contained ammonia concentration, with an Orion1 portable ion/ 40 larval or 10 juvenile test animals. Test chambers were pH meter and an Orion1 model 95-12 ammonia elec- randomly placed in water baths so that each water bath trode; water temperature and dissolved oxygen concen- accommodated one complete set of duplicate treatments tration, with a YSI1 model 57 oxygen meter; and speci®c and controls. To maintain a constant temperature of conductance, with a Cole-Parmer1 model 9100-00 con- 20C, water baths were equipped with Visitherm1 ther- ductivity meter. The fraction of total ammonia in un- mostatically controlled immersion heaters and Little ionized form was calculated from equations in Emerson Giant1 submersible pumps (for water circulation). Each et al. (1975). Water quality measurements in test test chamber received periodic deliveries of water from a chambers were discontinued after ®sh mortalities proportional diluter (Mount and Brungs, 1967), with reached 100%. displaced water over¯owing the test chamber through a Other water quality variables were measured from screened outlet. The diluter cycled at 15-min intervals, pooled water samples of each treatment at the beginning thus delivering about two changes of water per day to and end of each test. The variables and analytical pro- each test chamber. Prior to ¯owing into the test cham- cedures were as follows: total and calcium hardness, by ber, water from the diluter was temporarily stored in a titration with a HACH1 model HAC-DT Total-and- 4-litre head tank. The head tank was partially immersed Calcium Hardness kit; sulfates, by the HACH1 Sulfa- in the water bath to help equilibrate incoming water to Ver 4 method; total alkalinity, by potentiometric titra- thermal conditions in the test chamber. tion (APHA et al., 1981); and chlorides, by HACH1 For tests with pH and un-ionized ammonia, a Micro- mercuric nitrate buret titration or HACH1 silver medic1 precision metering pump was used to inject chloride buret titration. (Note: During this study, the stock solutions of either NaOH or ammonium chloride method for measuring chlorides was changed from

(NH4Cl) into the proportional diluter. By using recon- mercuric nitrate buret titration to silver nitrate buret stituted water, the diluter automatically created and titration to avoid production of mercuric chloride, a dispensed a control and ®ve progressively lower con- hazardous waste.) centrations (treatments) of test solution into individual At daily or weekly intervals, or at the beginning and test chambers. end of each test, water quality measurements were sub- For tests with temperature and dissolved oxygen, jected to quality control procedures that included use of proportional diluters were used only to deliver recon- calibration blanks, reference standards, and compar- stituted water to the test chambers. A series of ®ve pro- isons of instrument readings with measurements taken gressively higher water temperatures was achieved by with a general purpose precision thermometer (water placing thermostatically controlled aquarium heaters temperature) or the Winkler method with azide mod- into chambers separated by 2.5-cm-thick Styrofoam1 i®cation (dissolved oxygen; APHA et al., 1981). The sheets within the water baths. Five progressively lower resolution of various analytical procedures was not dissolved oxygen concentrations were achieved by bub- measured during this study. bling appropriate volumes of nitrogen gas, compressed air, or both, through water in the head tanks. The tops 2.3. Data analysis of test chambers were modi®ed for dissolved oxygen tests by attaching plexiglass covers to reduce di€usion Water quality data were logarithmically transformed by atmospheric oxygen. before computing means and tests of statistical sig- Treatments were checked for mortalities 1 h after ni®cance (e.g., analysis of variance, ANOVA) with the test was started, 24 h after the test was started, SAS1 System software (SAS Institute Inc., 1987). Point and at 24-h intervals thereafter. Test animals were estimates of 96-h LC50 values were calculated with considered dead if they showed no response to physi- computer software (lc50.bas, modi®ed 5/16/86) written cal contact (i.e., prodding with a glass rod). After 96 h, by the US Environmental Protection Agency (USEPA, survivors were euthanized by overdosing with MS-222, Environmental Monitoring and Support Laboratory, then discarded. In juvenile tests, individual test ani- Cincinnati, OH). The software also computed the 95% mals (both dead and alive) were weighed prior to con®dence interval for each point estimate. Methods for being discarded. interpolating LC50 values were selected after considering assumptions of the various statistical models (Gelber et 2.2. Water quality measurements al., 1985). The binomial method was used if none of the treatments resulted in partial mortalities. If partial The following water quality variables were measured mortalities occurred in at least one treatment, then twice daily in each test chamber: pH; total ammonia; either the binomial or the Spearman-Karber methods temperature; dissolved oxygen; and speci®c con- were used. If two or more treatments had partial mor- ductance. Hydrogen-ion concentration (pH) was meas- talities, the probit method was also used. When two or 1 2 ured with a Cole-Parmer Digisense pH meter; total more methods were suitable for estimating the LC50

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40 M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 value, we prioritized the reporting of results as follows: (Table 2). During the ®rst 48 h of exposure, mean LC50s binomial

Although mean LC50s computed for pH, un-ionized the gill area and eyes, and several had eyes that were ammonia, and temperature generally decreased with seemingly ruptured. If dead ®sh were not removed from increasing exposure time (i.e., 24, 48, 72, and 96 h), the highest pH treatments within 1±2 h following death, whereas those for dissolved oxygen increased with time, their body tissues began to dissolve or liquefy. Fish the corresponding 95% con®dence intervals typically surviving the 96-h tests did not bleed from the gills or exhibited broad overlap (Table 2). Such temporal com- eyes, or secrete noticeable amounts of mucous. parisons are not statistically valid due to pseudor- eplication (lack of independence); however, they suggest 3.2. Un-ionized ammonia tests that the LC50s were mostly determined by mortalities occurring during the ®rst 24 h. These ®ndings are con- Mean LC50s for un-ionized ammonia varied from a sistent with our observations that regardless of the low of 0.48 mg litre1 in larval Lost River suckers after water quality variable being tested, most ®sh exposed to 96 h of exposure to a high of 1.29 mg litre1 in larval lethal conditions died during the 24-h interval immedi- shortnose suckers after 24 h of exposure (Table 2). Jud- ately following initiation of the tests. ging from our data, the toxicity of un-ionized ammonia did not vary among ®sh species or life stages (Table 3). 3.1. Hydrogen-ion (pH) tests At higher concentrations, dying ®sh bled from the gills and some juveniles (but not larvae) exhibited signs of

In pH tests, mean LC50s for larval and juvenile Lost hyperactivity. Even at non-lethal concentrations, juve- River and shortnose suckers varied from 10.30 to 10.69 niles were seemingly hyperactive but in good condition

Table 2

Upper median lethal concentrations (LC50s) for pH, un-ionized ammonia (NH3), and water temperature (TEMP), and lower LC50s for dissolved oxygen (DO) to larval and juvenile Lost River (LR) and shortnose (SN) suckers at 24-h exposure intervals during 96-h-long tests

Variable Species and Weight (g) Mean LC50 values (95% con®dence intervals) after each exposure time: life stagea 24 h 48 h 72 h 96 h pH LR larva NWb 10.42 (10.38±10.47) 10.39 (10.32±10.46) 10.36 (10.27±10.46) 10.35 (10.26±10.45) LR juvenile 0.28±0.49 10.66 (10.59±10.74) 10.62 (10.54±10.71) 10.39 (10.12±10.67) 10.30 (9.94±10.67) SN larva NWb 10.38 (10.31±10.46) 10.38 (10.31±10.46) 10.38 (10.31±10.46) 10.38 (10.31±10.46) SN juvenile 1.01±1.11 10.69 (10.61±10.77) 10.66 (10.61±10.72) 10.58 (10.56±10.61) 10.39 (10.22±10.56)

1 b c c c c NH3 (mg L ) LR larva NW 0.56 (0.52±0.61) 0.51 (0.47±0.55) 0.49 (0.45±0.54) 0.48 (0.44±0.52) LR juvenile 0.49±0.80 1.02 (1.01±1.04) 0.92 (0.82±1.04) 0.89 (0.77±1.04) 0.78 (0.70±0.86) SN larva NWb 1.29 (0.83±2.00) 1.24 (0.82±1.88) 1.19 (0.79±1.78) 1.06 (0.73±1.53) SN juvenile 0.53±2.00 0.51 (0.30±0.87) 0.48 (0.28±0.82) 0.54 (0.35±0.82) 0.53 (0.34±0.82)

TEMP (C) LR larva NWb 31.93 (31.82±32.04)c 31.85 (31.69±32.01)c 31.77 (31.58±31.96)c 31.69 (31.47±31.91)c LR juvenile 0.48±0.86 30.76 (30.04±31.50) 30.76 (30.04±31.50) 30.65 (30.04±31.27) 30.51 (29.99±31.04) SN larva NWb 31.85 (31.75±31.96) 31.85 (31.75±31.96) 31.85 (31.75±31.96) 31.82 (31.75±31.90) SN juvenile 0.54±0.64 31.07 (29.44±32.80) 30.35 (29.44±31.28) 30.35 (29.44±31.28) 30.35 (29.44±31.28)

DO (mg L1) LR larva NWb 2.01 (1.90±2.13) 2.10 (2.07±2.13) 2.10 (2.07±2.13) 2.10 (2.07±2.13) LR juvenile 0.39±0.86 1.58 (1.35±1.86) 1.58 (1.35±1.86) 1.62 (1.41±1.86) 1.62 (1.41±1.86) SN larva NWb 1.92 (1.89±1.96) 2.04 (1.90±2.18) 2.09 (1.90±2.29) 2.09 (1.90±2.29) SN juvenile 0.39±1.15 1.14 (0.84±1.55) 1.34 (1.15±1.55) 1.34 (1.15±1.55) 1.34 (1.15±1.55)

a When tests were initiated, larvae were 35 days old whereas juveniles were 3±7 months old. b NW, test animals were not weighed. c This test was not repeated; the 95% con®dence interval was calculated from statistical procedures used to estimate the LC50 value.

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M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 41

Table 3

Results of two-way analysis of variance, as F-values and signi®cance levels, for median lethal concentrations (LC50s) of larval and juvenile life stages of Lost River and shortnose suckers exposed for as long as 96 h to pH, un-ionized ammonia (NH3), water temperature (TEMP), and dissolved oxygen (DO)

Variable Source Degrees of freedom Exposure time (h)

24 48 72 96 pH Species 1 0.01 0.06 0.49 0.08 Lifestage 1 15.16* 12.75* 0.56 0.02

Species*Lifestage 1 0.22 0.10 0.32 0.02 Error MS 4 0.00001662 0.00001774 0.00007940 0.00015652

NH3 Species 1 0.02 0.07 0.22 0.29 Lifestage 1 0.13 0.16 0.06 0.08

Species*Lifestage 1 2.94 3.02 3.17 2.42 Error MS 3 0.05997679 0.05971162 0.04636936 0.04290837 TEMP Species 1 0.01 0.07 0.02 0.00 Lifestage 1 0.66 2.79 3.21 3.60

Species*Lifestage 1 0.03 0.08 0.07 0.05 Error MS 3 0.00043933 0.00018770 0.00016752 0.00015346 DO Species 1 1.13 0.78 0.79 0.79 Lifestage 1 4.71 9.35* 9.89* 9.89*

Species*Lifestage 1 0.66 0.37 0.69 0.69 Error MS 4 0.01160837 0.00498678 0.00475048 0.00475048

*p0.05. at the end of the tests. By comparison, dying larvae were than juveniles to hypoxia (Table 2). Fish experiencing not hyperactive. Instead, the dying larvae generally hypoxia had diculty swimming, and exhibited a gasp- became listless (comatose) and, except for brief bursts of ing behavior while struggling to remain near the surface. rapid swimming, settled to the bottom of the test By comparison, ®sh in treatments containing higher chamber where they remained motionless until ®nally concentrations of dissolved oxygen swam freely succumbing to treatment e€ects. throughout the water column without exhibiting the gasping behavior. 3.3. Temperature tests

Mean LC50s for water temperature varied from 4. Discussion 30.35C among juvenile shortnose suckers after 48±96 h of exposure to 31.93C among larval Lost River suckers During summer and early fall, water quality condi- after 24 h of exposure (Table 2). However, two-way tions in Upper Klamath Lake ¯uctuate considerably in ANOVA failed to detect statistically signi®cant di€er- response to natural cycles in productivity of algae, ences among ®sh species and life stages (Table 3). Fish especially the cyanobacterium known as Aphanizomenon exposed to the highest treatments died shortly after ¯os-aquae (Bortleson and Fretwell, 1993; Wood et al., placement into test chambers (i.e., within 1 h). Dead ®sh 1996). Resource managers have long hypothesized that typically exhibited bloated abdomens and ¯oated at the stressful water quality conditions associated with algal surface. On several occasions, the abdomens of dead blooms or algal senescence were responsible for occa- ®sh were ruptured. Fish dying in intermediate treatment sional reports of ®sh kills in the lake (Scoppettone, levels typically succumbed at later time intervals (48 or 1986). Wood et al. (1996) mentioned that toxins pro- 72 h after tests were initiated) and did not exhibit duced by A. ¯os-aquae might also contribute to mor- bloated abdomens. talities. In addition, Microcystis aeruginosa (another toxin- producing cyanobacterium) occurred for the ®rst 3.4. Dissolved oxygen tests time at high densities in portions of Upper Klamath Lake during summer 1996 (J. Kann, unpublished data).

Mean LC50s for dissolved oxygen varied from 1.14 However, toxins associated with A. ¯os-aquae and mg litre1 to 2.10 mg litre1 (Table 2). During the ®rst M. aeruginosa have not yet been detected in lake 24 h of exposure, our data failed to demonstrate di€er- water (J. Kann, personal communication). This study ences among life stages and ®sh species (Table 3). was a preliminary attempt to link excessively high pH, However, beginning at 48 h and extending through 96 h, un-ionized ammonia concentration, and water tempera- a signi®cant ( p<0.05) di€erence occurred among life ture, and excessively low dissolved oxygen concentration stages. In general, larvae were seemingly more sensitive with mortality of larval and juvenile Lost River and

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42 M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 shortnose suckers. As such, the study focused on indi- (1997) reported un-ionized ammonia concentrations as vidual water quality variables rather than on the com- high as 0.61 mg litre1 in Upper Klamath Lake, the bined e€ects of multiple variables. Because the study mean concentrations never exceeded 0.21 mg litre1. design did not allow for gradual acclimation of ®sh to Moreover, on two occasions when un-ionized ammonia extreme test conditions, our test results should be concentrations in lake water temporarily approached or viewed as conservative estimates of the lethal levels of exceeded LC50 values computed by the present study, these water quality variables. juvenile Lost River suckers held in cages did not experi- Our results indicated that larval and juvenile Lost River ence mortality (Martin, 1997). Collectively, these results and shortnose suckers acclimated to presumably innoc- suggest that un-ionized ammonia concentrations may uous water quality conditions±pH, 8.0; un-ionized not remain elevated suciently long in Upper Klamath ammonia, <0.01 mg litre1; temperature, 20C; and Lake to cause ®sh mortality. 1 dissolved oxygen, 7.6 mg litre or 100% saturation± During our study, the LC50s for water temperature experienced high mortality (50% or more of ®sh died) varied from 29.4C to 31.9C (Table 2). Bortleson and when exposed for as long as 96 h to pH averaging Fretwell (1993) mention that summertime water tem- 10.30±10.69 (Table 2). Although the average daily pH peratures in Upper Klamath Lake usually ¯uctuate from in Upper Klamath Lake rarely exceeds 10.0 during 21Cto25C, although temperatures as high as 30C summer±early fall, maximum pH frequently surpasses might occur in surface algal scums. However, ®eld stud- 10.5 (Martin, 1997). However, Martin (1997) noted that ies by several other investigators indicate that surface mortalities did not occur among juvenile Lost River temperatures rarely exceed 26±27C (Coleman et al., suckers caged during a series of 4-day-long ®eld tests in 1988; Martin, 1997; J. Kann, unpublished data). During Upper Klamath Lake even when pH momentarily summer±early fall 1995, the maximum temperature peaked at 10.78. Our results, combined with ®ndings reported by Martin (1997) was 26.65C. These results from Martin (1997), suggest that high pH may not be suggest that elevated water temperatures are not expec- sustained long enough in Upper Klamath Lake to ted to be an immediate source of mortality among suck- directly cause ®sh mortality. In addition, peaks in pH ers inhabiting Upper Klamath Lake. Moreover, suckers typically coincided with supersaturated dissolved oxy- can avoid overly warm water temperatures by seeking gen concentrations (Martin, 1997). Serafy and Harrell cooler water near the bottom (if adequately oxygen- (1993) demonstrated that ®sh may remain in high-pH ated), or at river mouths and underground springs habitats (such as aquatic weed beds) if dissolved oxygen where cooler temperatures generally prevail (Buettner is supersaturated even though the pH exceeds thresh- and Scoppettone, 1990). Although not examined during olds that ordinarily trigger avoidance behavior. Accord- our study, summertime temperatures in Upper Klamath ing to Serafy and Harrell (1993), high pH adversely Lake could exert sublethal in¯uences on suckers by a€ects the proper functioning of ®sh gills (by damaging altering their spatial distribution (and hence, their access surface tissues and precipitating excessive mucous to suitable cover and forage) and increasing their meta- secretion) and severely impairs oxygen uptake. How- bolic requirements. ever, some of the lost gill function may be nulli®ed The mean LC50s of juvenile shortnose suckers to high under conditions of oxygen supersaturation. temperature (30.35±31.07C) were 1.6±2.4C lower than During our study, the pH tests with juvenile short- the critical thermal maximum of 32.7C reported by nose suckers yielded mean LC50s that varied from 10.39 Castleberry and Cech (1993). The procedure used (96-h exposure) to 10.69 (24 h exposure), whereas Falter by Castleberry and Cech was expected to yield a ther- and Cech (1991) reported that the critical pH maximum mal value that was higher than our LC50s because, as for this species and life stage is 9.55‹0.43. Although the mentioned earlier for pH, the critical maximum tends to di€erent values were not compared for statistical sig- exceed the tolerance level determined by longer-duration ni®cance, we had expected an opposite pattern where tests such as those used for calculating LC50 values LC50 values are lower than the critical pH maximum. (Becker and Genoway, 1979). In other words, if the According to Becker and Genoway (1979), the critical exposure period is suciently brief (seconds or minutes), maximum (determined by gradually increasing pH or ®sh can endure high water temperatures that would be some other variable away from acclimation over a per- fatal if the exposure period was longer (hours or days). iod of several minutes or a few hours until loss of equi- Our estimates of the LC50s for minimum dissolved librium or death occurs) usually exceeds the tolerance oxygen concentrations varied from 1.14 mg litre1 to 1 level determined by the LC50 because most animals do 2.10 mg litre (Table 2). In Upper Klamath Lake, dis- not immediately lose equilibrium or die when exposed solved oxygen concentrations ¯uctuate widely in to conditions that can eventually cause death. response to weather conditions and cycles in algal pro-

In our tests with un-ionized ammonia, mean LC50s ductivity (Coleman et al., 1988; Martin, 1997). Oxygen varied from 0.48 mg litre1 to 1.29 mg litre1 for expo- concentrations less than 4.5 mg litre1 can persist lake- sure times of 24 h to 96 h (Table 2). Although Martin wide for several weeks to a month, and concentrations

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M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44 43 less than 0.5 mg litre1 can occur for more than 6 h Lost River and shortnose suckers in Upper Klamath either throughout the water column in isolated areas of Lake. Overall, temporary episodes of near-anoxic con- the lake or near bottom when the lake is thermally ditions occurring during the senescence phase of algal strati®ed (Coleman et al., 1988; Kann and Smith, 1993; blooms are perhaps the most serious threat to ®sh. Martin, 1997). Collectively, these observations indicate Although pH and un-ionized ammonia in lake water that low dissolved oxygen concentrations in Upper can temporarily approach or exceed lethal levels pre-

Klamath Lake may occasionally exceed the lethal con- dicted by our LC50s, the exposure duration may be too centrations for Lost River and shortnose suckers. Martin brief to directly cause ®sh kills. Mortality of larvae and (1997) demonstrated that juvenile Lost River suckers juveniles from exposure to lethal water quality during caged in Upper Klamath Lake experienced high mor- summer and early fall could explain why the numbers of talities (>90%) whenever dissolved oxygen concentra- adult suckers in Upper Klamath Lake are low even tions temporarily fell below 1.40 mg litre1, whereas though these adults spawn successfully in tributaries only a few ®sh died (<10% mortality) when dissolved such as the Williamson and Sprague rivers (and possibly oxygen concentrations equalled or exceeded 1.58 mg the Wood River and Crooked Creek) and in Sucker litre1. Although suckers in the lake can avoid asphyx- Springs on the eastern shore of the lake (Stubbs and iation if they ®nd better-oxygenated waters at the White, 1993). Future studies should attempt to estimate mouths of rivers or outlets to underground springs lethal thresholds for multiple combinations of water (Buettner and Scoppettone, 1990), such actions must be quality variables to better approximate the conditions taken within 1±2 h or less (the time span during which occurring in Upper Klamath Lake. dissolved oxygen concentrations in large portions of Upper Klamath Lake can plummet from 5 mg litre1 to 0 mg litre1; Martin, 1997). Acknowledgements Castleberry and Cech (1993) reported a critical dis- solved oxygen minimum of 0.44±0.98 mg litre1 for We thank M. Brooks, D. Cappellini, D. Haapala, M. juvenile shortnose suckers. By comparison, the esti- Jordan, M. Keeler, B. Martin, R. Renfro, L. Whitley, mated LC50s for juvenile shortnose suckers averaged and L. Wilcox for assisting with the laboratory work; 1.14±1.34 mg litre1. We expected the critical dissolved M. Buettner, S. Campbell, J. Conner, L. Dunsmoor, oxygen minimum computed by Castleberry and Cech and J. Kann for reviewing early drafts of this manu- (determined by gradually decreasing the dissolved oxy- script and providing other technical assistance; and The gen concentration over a period of minutes or hours Klamath Tribe for supplying the test animals. This until loss of equilibrium or death occurs) to be lower study was funded by the US Bureau of Reclamation than our estimated LC50s because most animals do not through several reimbursable agreements with the Bio- immediately lose equilibrium or die when exposed to logical Resources Division (US Geological Survey) or conditions that can eventually cause death (Becker and its predecessor agencies. Genoway, 1979). Although beyond the scope of our study, Martin (1997) noted that several combinations of high pH, high References un-ionized ammonia concentrations, high temperature, and low dissolved oxygen concentrations were see- American Public Health Association (APHA), American Water Works Association and Water Pollution Control Federation, 1981. mingly associated with increased mortality of juvenile Standard Methods for the Examination of Water and Wastewater, Lost River suckers caged in Upper Klamath Lake. These 15th Edition, 1980. American Public Health Association, Washing- ®ndings are consistent with studies indicating that phy- ton, DC, USA. siological stress (as evidenced by increased mucous pro- Becker, C.D., Genoway, R.G., 1979. Evaluation of critical thermal maxima for determining thermal tolerance of freshwater ®sh. duction and hemorrhaging in gills and other tissues) in Environmental Biology of Fishes 4, 245±256. ®sh exposed to high pH is exacerbated by high tem- Bienz, C.S., Ziller, J.S., 1987. Status of three lacustrine sucker species peratures, low dissolved oxygen concentrations, or both (Catostomidae). Completion report, The Klamath Tribes. Fish and (Serafy and Harrell, 1993; Svobodova et al., 1993); that Wildlife Department, Chiloquin, OR. toxicity of un-ionized ammonia increases at higher pH Bortleson, G.C., Fretwell, M.O., 1993. A review of possible causes of nutrient enrichment and decline of endangered sucker populations levels and lower dissolved oxygen concentrations in Upper Klamath Lake, Oregon (U.S. Geological Survey, Water (Russo, 1985; Svobodova et al., 1993); and that oxygen Resources Investigations Report 93-4087). Portland, OR. requirements of ®sh generally increase at higher tem- Buettner, M., Scoppettone, G., 1990. Life history and status of peratures or higher pH levels (Svobodova et al., 1993). Catostomids on Upper Klamath Lake, Oregon. Completion report. In conclusion, results from our study support the U.S. Fish and Wildlife Service, National Fisheries Research Center, Seattle, WA. contention of resource managers that extreme water Castleberry, D.T., Cech, J.J., 1993. Critical thermal maxima and oxy- quality conditions encountered during an algal bloom gen minima of ®ve ®shes from the Upper Klamath Basin. California or the ensuing senescence phase can be acutely lethal to Fish and Game 78, 145±152.

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44 M.K. Saiki et al. / Environmental Pollution 105 (1999) 37±44

Coleman, M.E., Kann, J., Scoppettone, G.G., 1988. Life history and Russo, R.C., 1985. Ammonia, nitrite, and nitrate. In: Rand, G.R., ecological investigations of catostomids from the Upper Klamath Petrocelli, S.R. (Eds.), Fundamentals of Aquatic Toxicology, Basin, Oregon. Completion report. U.S. Fish and Wildlife Service, Chapter 15. Taylor and Francis (Hemisphere Publishing Corpora- National Fisheries Research Center, Seattle, WA. tion), Bristol, PA, pp. 455±71. Emerson, K., Russo, R.C., Lund, R.E., Thurston, R.V., 1975. Aqueous SAS Institute Inc., 1987. SAS/STAT User's Guide, release 6.03 Edi- ammonia equilibrium calculations: e€ects of pH and temperature. tion. Cary, NC. Journal of the Fisheries Research Board of Canada 32, 2379±2383. Scoppettone, G.G., 1986. Upper Klamath Lake, Oregon, Catostomid Falter, M.A., Cech, J.J., 1991. Maximum pH tolerance of three Kla- research. Completion report. U.S. Fish and Wildlife Service, math Basin ®shes. Copeia 4, 1109±1111. National Fisheries Research Center, Seattle, WA. Gelber, R.D., Lavin, P.T., Mehta, C.R., Schoenfeld, D.A., 1985. Sta- Serafy, J.E., Harrell, R.M., 1993. Behavioural response of ®shes to tistical analysis. In: Rand, G.R., Petrocelli, S.R. (Eds.), Funda- increasing pH and dissolved oxygen: ®eld and laboratory observa- mentals of Aquatic Toxicology, Chapter 5. Taylor and Francis tions. Freshwater Biology 30, 983±988. (Hemisphere Publishing Corporation), Bristol, PA, pp. 110±23. Svobodova, Z., Lloyd, R., Machova, J., Vykusova B., 1993. Water Kann, J., Smith, V.H., 1993. Chlorophyll as a predictor of elevated quality and ®sh health (European Inland Fisheries Advisory Com- pH in a hypertrophic Lake: estimating the probability of exceeding mission Technical Paper No. 54). Food and Agriculture Organiza- critical values for ®sh success (Klamath Tribes Research Report tion of the United Nations, Rome, Italy. KT-93-02). The Klamath Tribes. Chiloquin, OR. Stubbs, K., White, R., 1993. Lost River (Deltistes luxatus) and Short- Martin, B.A., 1997. E€ects of ambient water quality on the endan- nose (Chasmistes brevirostris) Sucker recovery plan. U.S. Fish and gered Lost River sucker (Deltistes luxatus) in Upper Klamath Wildlife Service, Portland, OR. Lake, Oregon. MSc Thesis, Humboldt State University, Arcata, Wood, T.M., Fuhrer, G.J., Morace, J.L., 1996. Relation between CA. selected water-quality variables and lake level in Upper Klamath Mount, D.I., Brungs, W.A., 1967. A simpli®ed dosing apparatus for and Agency Lakes, Oregon (U.S. Geological Survey, Water ®sh toxicology studies. Water Research 1, 21±29. Resources Investigations Report 96-4079) Portland, OR.

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