- a- " - TR EL-82-4 Biological Report 82(11.86) January 1989 Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Northwest) SEA-RUN CUTTHROAT TROUT Fish and Wildlife Service -US. Department of the Interior Coastal Ecology Group Waternays Experiment Station US. Army Corps of Engineers B i o l o g i c a l Report 82(ll. 86) TR EL-82-4 January 1989 Species P r o f i l e s : L i f e H i s t o r i e s and Environmental Requirements o f Coastal Fishes and I n v e r t e b r a t e s ( P a c i f i c Northwest) SEA-RUN CUTTHROAT TROUT G i l b e r t B. Pauley, Kevin Oshima, Karen L. Bowers, and G. L - Thomas Washington Cooperative F i s h e r y Research U n i t School o f F i s h e r i e s U n i v e r s i t y o f Washington S e a t t l e , WA 98195 Project O f f i c e r David Moran U.S. F i s h and W i l d l i f e S e r v i c e N a t i o n a l Wetlands Research Center 1010 Gause Boulevard S l i d e l l , LA 70458 This study was conducted i n cooperation w i t h Coastal Ecology Group Waterways Experiment S t a t i o n U. S. Army Corps o f Engineers Vicksburg, MS 39180 Performed f o r U.S. Department of the I n t e r i o r F i s h and W i l d l i f e S e r v i c e Research and Development N a t i o n a l Wetlands Research Center Was h i ngton , DC 20240 T h i s series may be referenced as follows F i s h and Wildlife Service. 1983-19 . Species profiles: life histories and environmental requirements of coastal fishes and invertebrates. U. S. Fish W i l d l . Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers TR EL-82-4. U.S. This profile may be cited as follows: Pauley, G. 0. , K. Oshima, K. C. Bowers, and G. L. Thomas. 1989, Species prof'i l e s : 1 i fe hi stories and environmental requirements of coastal fishes and invertebrates ( P a c i f i c Northwest)--sea-run cutthroat trout. U.S. Fish Wi ld. Serv. Biol. Rep. 82(11.86). U.S. Army Corps o f Engineers TR EL-82-4. 21 PP - PREFACE This species profile i s one of a series on coastal aquatic organisms, principally f i s h , o f sport, commercial, or ecological importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profile has sections on taxonomy, life history, ecological role, environmental requirements, and economic importance, i f applicable. A three-ring binder is used for this series so that new profiles can be added as they are prepared. This project is jointly planned and financed by the U. S. Army Corps of Engineers and the U.S. Fish and Wildlife Service. Suggestions or questions regarding this report should be directed to one of the following addresses. Information Transfer Specialist National Wetlands Research Center U.S. Fish and Wildlife Service NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA 70458 U. S . Army Engineer Waterways Experiment Statior Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 CONVERSION TABLE M e t r i c t o U.S. Customary Tc - Mu1 t i p l y Obtain m i l l i m e t e r s (mm) c e n t i m e t e r s (cm) meters (m) meters (m) k i lometers (km) k i lometers (km) inches inches feet fathoms statute miles nautica miles square meters ( m 2 ) square k i lometers (km2) h e c t a r e s (ha) square f e e t square m i l e s acres l i t e r s (1) c u b i c meters ( m 3 ) c u b i c meters (m3) ga 1 1 ons cubic f e e t acre- f e e t m i l l i g r a m s (mg) grams ( g ) kilograms (kg) m e t r i c tons ( t ) m e t r i c tons ( t ) ounces ounces pounds pounds s h o r t tons kilocalories (kcal) C e l s i u s degrees ("C) B r i t i s h thermal u n i t s F a h r e n h e i t degrees U. S. Customary t o Metric inches inches feet ( f t ) fathoms statute miles ( m i ) n a u t i c a l m i l e s (nmi) mi 11 i m e t e r s centimeters meters meters kilometers kilometers square f e e t ( f t ' ) square m i l e s ( m i 2 ) acres square meters square k i l o m e t e r s hectares gallons (gal) cubic feet ( f t 3 ) acre- f e e t 1i t e r s cubic meters cubic meters ounces (02) ounces ( o z ) pounds ( l b ) pounds ( I b ) s h o r t tons ( t o n ) m i 1 1 igrams B r i t i s h thermal u n l t s ( B t u ) F a h r e n h e ~ tdegrees ( O r ) grams k i lograms m e t r i c tons metric tons 0.2520 0.5556 iv (OF kilocalories C e l s i u s degrees CONTENTS Page PREFACE . . . . . CONVERSION TABLE . ACKNOWLEDGMENTS . . . . . . . . . . . . . . . . . . .;. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . NOMENCLATURE/TAXONOMY/RANGE iii iv vi . . MORPHOLOGY/IDENTIFICATION AIDS . REASON FOR INCLUSION IN SERIES . LIFE HISTORY . . . . . . . . . . Spawning . . . . . . . . . Eggs and.Fecundity . . . . Alevins and Fry . . . . . . Juveniles and Smolting . . Saltwater L i f e . . . . . . AGE AND GROWTH CHARACTERISTICS . ECOLOGICAL ROLE . . . . . . . . ENVIRONMENTAL REQUIREMENTS . . . Temperature . . . . . . . . Substrate and Cover . . . . Dissolved Oxygen . . . . . Turbidity . . . . . . . . . Water Velocity . . . . . . Water Depth . . . . . . . . . . . . . . . . . . . . . . . . Timber Harvest THE FISHERY LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 ACKNOWLEDGMENTS The authors thank James Johnston of the Washington Department o f Game and Howard Fuss of t h e Washington Department o f Fisheries f o r t h e i r reviews and technical suggestions. Figure 1. Sea-run c u t t h r o a t t r o u t . SEA-RUN CUTTHROAT TROUT S c i e n t i f i c name.. .. Salmo c l a r k i c l a r k i Preferred common name. . . .Sea-run cutthroat t r o u t (Figure 1) Other common names . . . .Coastal cutt h r o a t t r o u t , red-throated t r o u t , sea t r o u t , sea-run c u t t h r o a t , searun c u t , bl ueback t r o u t , autumn t r o u t , harvest t r o u t , ye1 lowbel ly. Class . . . . . . . . . . . . . . . . . . . . .Osteichthyes Order .................... Salmoniformes Family ...................... Salmon idae --- + Geographic range: In coastal streams from northern Cal i f o r n i a through Oregon and B r i t i s h Columbia and i n t o southeastern Alaska (Figures 2 and 3). Rarely found more than 160 km inland. MORPHOLOGY/IDENTIFICATION A I D S The following information i s taken from Hart (1973), Scott and Crossman (1973), and Wydoski and Whitney (1978). Morphological differences between coastal cut. t h r o a t and inland c u t t h r o a t were outlined by Qadri (1959) and T r o t t e r (1987). Morphology: Fin rays--dorsal fin anal 8-12, p e c t o r a l s 13, and pelvic 9. Adipose, small, fleshy and s l e n d e r , pelvics abdominal i n position with a free- tipped f 1 eshy appendage above i n s e r t i o n . Cycloid s c a l e s , 120180 a t l a t e r a l l i n e ; g i l l rakers 1522, rough and widely spaced on f i r s t g i l l arch. Body elongate, s l i g h t l y compressed. 8-11, I d e n t i f i c a t i o n aids: Red o r orange s t r e a k along the inner edge o f lower jaw in fresh specimens. Small t e e t h a t the back of the tongue between the second g i l l c l e f t s . Teeth well developed on upper and lower jaws. Maxi l lary usual ly extends beyond the p o s t e r i o r margin o f the eye in f i s h longer than 10 cm. Back i s greenishblue tending toward m e t a l l i c blue in f r e s h sea-run f i s h ; s i d e s s i l v e r y ; d i s t i n c t black spots on b a c k , head, CANADA '. WASHINGTON OREGON F i g u r e 2. Map showing m a j o r r i v e r s t h a t s u p p o r t sea-run c u t t h r o a t t r o u t i n t h e P a c i f i c Northwest. I t s h o u l d be n o t e d t h a t many small streams t o o numerous t o be shown on t h i s map a l s o s u p p o r t sea-run c u t t h r o a t s . An a c t i v e s p o r t f i s h e r y 2 OREGON / MILES 0 0 0 50 100 KILOMETERS e x i s t s f o r these f i s h i n s a l t w a t e r throughout as in t h e rivers. Puget Sound and Hood Canal as well ALASKA , CANADA ITEO STATES Figure 3. Sea-run c u t t h r o a t t r o u t coastal d i s t r i b u t i o n from northern California t o southeastern Alaska and the general l i m i t s of inland d i s t r i b u t i o n in North America ( a f t e r Johnston and Mercer 1976). anal f i n , t a i l , and sides (extending well below l a t e r a l l i n e ) . S i z e range o f adults i s 0 . 5 to 2.7 kg. REASON FOR INCLUSION IN SERIES Although not quite a s well-known a s i t s larger cousin the steelhead (Salmo g a i rdneri ) , the sea-run cutthroat t r o u t i s extensively sought by anglers through the rivers and small streams of Northern Cal if o r n i a , Oregon, Washington, and B r i t i s h Columbia t h a t flow into saltwater. The freshwater s p o r t f i s h e r y takes place in the f a l l , an extensive saltwater sport f i s h e r y exists y e a r round in inland waters such as Puget Sound. The searun c u t t h r o a t i s highly sought a f t e r by fly-fishermen. while LIFE HISTORY Spawni ng Sea-run c u t t h r o a t t r o u t have a variable l i f e h i s t o r y , which will be discussed below. Like salmon, sexually mature sea-run c u t t h r o a t t r o u t r e t u r n t o t h e i r n a t a l streams t o spawn. Spawning sea-run c u t t h r o a t t r o u t home precisely t o specific t r i b u t a r i e s , w h i l e immature f i s h do n o t always r e t u r n t o t h e i r home stream t o feed o r when seeking an overwinter h a b i t a t (Johnston 1982). Homing o f n a t i v e sea-run c u t t h r o a t i s extremely p r e c i s e (Campton l98O), a1though hatcheryp l a n t e d f i s h may s t r a y as much as 30%. which makes s u r v i v a l r a t e s impossible t o determine (Johnston and Mercer 1976). Gel electrophoretic studies have shown t h a t c u t t h r o a t p o p u l a t i o n s are g e n e t i c a l l y d i s c r e t e a t t h e small stream l e v e l (Campton 1980), a1 though there i s l i m i t e d hybridization w i t h hatchery-propagated steelhead t r o u t (Campton and U t t e r 1985). The time o f r e t u r n t o freshwater f o r spawning i s f a i r l y c o n s i s t e n t among c u t t h r o a t t r o u t from t h e same stream, b u t v a r i e s widely by geographical l o c a t i o n (Johnston and Mercer 1976; Johnston 1982). In Oregon and Washington, c u t t h r o a t t r o u t r e - e n t e r freshwater anytime from J u l y through 1952; Anderson and March (Sumner Narver 1975). Very few o v e r w i n t e r i n s a l t w a t e r (Johnston 1982). A spring r u n has been observed o n l y i n Alaska (Jones 1972). I n a d d i t i o n , a small percentage o f the immigrants a r e sexuall y immature i n d i v i d u a l s (Jones 1973-76; Johnston 1982; T i p p i n g 1986). I n most coastal r i v e r s o f Washington and Oregon, the stocks o f f i s h are s e x u a l l y mature a t f i r s t r e t u r n t o freshwater; however, a l a r g e percentage of c u t t h r o a t females i n t h e Columbia River, Puget Sound, B r i t i s h Columbia, and Alaska do n o t spawn during the winter o f t h e i r f i r s t r e t u r n t o freshwater (Johnston 1982). Anadromous c u t t h r o a t t r o u t travel f a r t h e r upstream toward t h e headwaters of t h e watershed than e i t h e r steelhead t r o u t o r coho salmon and r e a r sympatr i c a l l y w i t h r e s i d e n t c u t t h r o a t popul a t i o n s (Johnston 1982; Michael 1983). Spawning g e n e r a l l y takes p l a c e i n l a t e w i n t e r and spring, b u t t i m i n g v a r i e s by geographic l o c a t i o n (Dymond 1932; Cramer 1940; S c o t t and Crossman 1973). I n Puget Sound and southern B r i t i s h Columbia, coastal c u t t h r o a t t r o u t e x h i b i t two d i s t i n c t m i g r a t i o n (1) f i s h r e t u r n i n g t o l a r g e times: r i v e r systems b e g i n e n t e r i n g i n J u l y and peak i n September-October ( e a r l y e n t r y f i s h ) and (2) f i s h r e t u r n i n g t o small streams d r a i n i n g d i r e c t l y t o s a l t w a t e r begin e n t e r i n g i n November and peak i n January-February ( l a t e e n t r y f i s h ) (Johnson 1982; Mercer 1982). This d i f f e r e n c e i s most l i k e l y due t o food avai 1a b i 1i t y , s a l t w a t e r t o 1 erance, o r stream flows (Johnston 1982). Throughout t h e i r range, sear u n c u t t h r o a t t r o u t spawn i n small t r i b u t a r i e s o f large o r small sti-eams w i t h a drainage area l e s s than 13 km and may thereby a v o i d c o m p e t i t i o n f o r r e a r i n g area w i t h steelhead (sea-run rainbow t r o u t . Salmo a a i r d n e r i 1 and coho salmon ' ( ~ n c ~ rnh m i u t c h ) (Cramer 1940; Sumner 19 2; OeWitt 1954; Glova and Mason 1977; Johnston 1982). Cutthroat trout prefer with small -tospawning habitat moderate-size gravel (Hunter 1973). A Spawning behavior i n c u t t h r o a t t r o u t i s t y p i c a l o f o t h e r stream-spawning salmonids such as steelhead and salmon (Smith 1941; Cope 1957; S c o t t and Crossman 1973). The female d i g s a r e d d i n t h e g r a v e l and deposits her eggs. Simultaneously, t h e male covers t h e eggs w i t h m i l t , The female then b u r i e s t h e eggs under a l a y e r o f gravel. A1 though c u t t h r o a t t r o u t a r e r e p e a t spawners, post-spawni ng mort a l i t i e s are sometimes h i g h due t o weight l o s s and t h e r i g o r s o f spawning (Cramer 1940; Sumner 1952; Giger 1972; S c o t t and Crossman 1973). The f i s h t h a t s u r v i v e m i g r a t e downstream i n e a r l y t o l a t e spring. K e l t s (spent a d u l t s ) e n t e r s a l t w a t e r e a r l i e r than smol t i ng j u v e n i 1es (Giger 1972; Jones 1975). Sumner (1952) s t a t e d t h a t 39% o f the c u t t h r o a t survived the f i r s t spawning i n Oregon, 17% t h e second, and 12% t h e t h i r d . These d a t a are r e p r e s e n t a t i v e o f c o n d i t i o n s where a t a r g e t e d c u t t h r o a t f i s h e r y does n o t exist. Sportfishing pressure reduces the survival of spawning adults following the first migration (Giger 1972). Eggs and Fecundity The spherical eggs of cutthroat trout range in size from 4.3 to 5.1 mm in diameter. Fecundity depends on the sire and age of the female. In Alaska, fish ranging in sire from 34 to 46 cm produced 486 to 2,286 eggs 1975). In per female (Jones Washington, females ranging from 20 to 43 cm produced 226 to 4,420 eggs (Johnston and Mercer 1976). Average number of eggs was between 1,100 and 1,700 for females of all sizes pooled (Scott and Crossman 1973). Mercer (1982) showed a strong linear correlation between increased length and increased number of eggs, with an "r" value of -97. Eggs normally hatch in 28 to 40 days, but may require 50 days or more (Merriman 1935; Cope 1957; Scott and Crossman 1973). Hatching is temperature-dependent (Merriman 1935). Alevins and Fry ~ e w l hatched ~ yolk-sac larvae, called alevins, remain in the gravel in the redd for 1 to 2 weeks before they emerge as fry, The fry are less than 3 cm long and commonly live in the shallow, low-velocity stream margins (Johnston and Mercer 1976). However, their distribution is often governed by the presence of other salmonid species (see Ecological Role section). The first scales are formed when the fish are 3.5 cm in fork ( F L)--a length normal ly length attained during the first summer of life (Cooper 1970; Giger 1972). The fry are sensitive to many kinds of environmental changes. Logging (Mori ng and Lantz l974), increased temperatures (Go1 den l975), loss of cover (Lowery l965), reductions in food supply, and s i 1 tation (Bustard and Narver 1975) can all increase larval mortality. Johnston and Mercer (1976) listed a number of natural sources of mortality, such as interspecific competition with other salmonids, intraspecific competition, and crowding induced by low summer flows, but they indicated that habitat a1 teration was probably the greatest threat to cutthroat trout stocks. Juveni 1 es and Smolti ng Prior to smolting and entering saltwater, juveniles (parr) may migrate up- and downstream several times (Sumner 1953; Moring and Lantz 1975). This migration can result i n confusion for someone trying to count or estimate the number of smolting cutthroat trout migrating downstream and out into the marine environment. The age at which smolting first occurs is extremely variable, being somewhat size-dependent and occurring between Age I and Age IV (Moring and Lantz 1975) or even later (Johnston and Mercer 1976; Jones 1977; Fuss 1978). A good working average is between Ages I11 and IV. Fuss (1978, 1982) noted that most initial migrants are Aged I11 or IV and outlined several reasons why initial migration occurs at different ages, Fish in these age groups average 20-25 cm FL. In Washington and Oregon, downstream movement of smolts takes place from March to June, but peaks in mid-May (Johnston and Mercer 1976). A division of migration types seems to exist between cutthroat trout entering the protected water of areas such as Puget Sound and those entering areas of surf-pounded coast. Cutthroat trout entering protected waters are, on average, younger and smaller (Age 11, 16 cm FL) (Michael 1983; Johnston 1980), while those entering surf zones are, on average, Age IV and 21 cm (Fuss 1982). Regardless o f age or habitat, cutthroat trout school before entering the marine environment and remain schooled throughout their saltwater migrations (Gi ger 1972)--a behavior which probably has s u r v i v a l value. Saltwater L i f e Although t h e r e have been few s t u d i e s o f c u t t h r o a t t r o u t movements a t sea, i t appears t h a t t h e y overwinter i n t h e marine environment and s t a y c l o s e t o shore. A1 though c u t t h r o a t remain a t sea v a r y i n g lengths o f time, they r e t u r n t o freshwater i n the same y e a r i n which they migrated o u t t o sea (Giger 1972; Anderson and Narver 1975; Jones 1976; Johnston and Mercer 1976). I n Puget Sound, Washington, the anadromous c u t t h r o a t t r o u t feed and m i g r a t e along t h e beaches, m o s t l y i n water l e s s than 3-m deep (Johnston 1980). I n general, t h e i r movement a l o n g t h e c o a s t i s b e l i e v e d t o be c o r r e l a t e d w i t h onshore ocean c u r r e n t s , w i t h the f i s h staying close t o the c o a s t l i n e (Giger 1972; Johnston and Mercer 1976). Regardless o f age, anadromous c u t t h r o a t t r o u t are b e l i e v e d t o b e g i n schooling j u s t b e f o r e they e n t e r i n t o the marine h a b i t a t (Johnston 1980). I n Alaska, c u t t h r o a t t r o u t were r e l u c t a n t t o cross bodies o f water 3 t o 8 km wide', and p r e f e r r e d t o f o l l o w t h e shore1 i n e (Jones 1976). - Many c u t t h r o a t t r o u t have been captured w i t h scars from p r e d a t o r attack while a t sea, and that p r e d a t i o n i s b e l i e v e d t o represent a major source o f marine m o r t a l i t y (Giger 1972). Estimates from Oregon s t u d i e s i n d i c a t e 20%-40% s u r v i v a l o f i n i t i a l migrant f i s h i n the marine environment (Giger 1972). Tipping (1986) noted that survival was considerably h i g h e r f o r hatchery sear u n c u t t h r o a t t r o u t t h a t were r e l e a s e d as smolts >21 cm FL (12.8% r e t u r n s ) t h a n f o r f i s h l e s s than t h a t l e n g t h (2.3% r e t u r n s ) . AGE AND GROWTH CHARACTERISTICS Growth i n c u t t h r o a t t r o u t v a r i e s according t o the type of freshwater and sal twater envi ronments they inhabit. Johnston and Mercer (1976) summarized growth o f a d u l t c u t t h r o a t trout in the Pacific Northwest (Oregon, B r i t i s h Columbia, and Alaska) and s t a t e d t h a t f i s h t h a t l i v e i n the lower stream grow more r a p i d l y than f i s h t h a t r e s i d e upstream and t h a t a1 so basic stream productivity influences growth. N i c k e l son and Larson (1974) i n d i c a t e d t h a t c o a s t a l c u t t h r o a t t r o u t n o t o n l y l o s t weight when f o o d i n streams was scarce, b u t t h a t t h e y a l s o decreased i n length. It i s n o t known how f r e q u e n t l y t h i s occurs. Giger (1972) c i t e d problems . t h a t were i n h e r e n t i n d e t e r m i n i n g c u t t h r o a t t r o u t age and growth due t o v a r i a b l e 1i f e c y c l e s and d i f f e r e n t m i g r a t i o n strategies. An unusual p a t t e r n o f age-length d i s t r i b u t i o n s e x i s t s f o r i n i t i a l outmigrants because t h e f i s h which spend more time i n streams p r i o r t o o u t m i g r a t i o n grow slower. This slower growth may be a r e s u l t o f the greater competition for food in streams t h a n occurs i n t h e marine environment. The r e s u l t i s t h a t f i s h the same system entering from s a l t w a t e r f o r t h e f i r s t t i m e a t Age I V are o n l y s l i g h t l y l a r g e r than i n i t i a l o u t m i g r a n t s of Age 11, and are much s m a l l e r t h a n Age I V f i s h t h a t have spent one o r more summers a t sea. (1978) i n d i c a t e d t h a t backFuss c a l c u l a t e d l e n g t h s f o r young f i s h ( l e s s t h a n 2 years o l d ) may be i n error, but the calculations f o r f i s h 2 y e a r s o l d o r o l d e r a r e probably a c c u r a t e because of the near l i n e a r r e l a t i o n s h i p between f i s h l e n g t h and s c a l e r a d i u s f o r the o l d e r f i s h . According t o Fuss (l978), Age I+ c u t t h r o a t ranged from 69 t o 121 mm and Age 11+ c u t t h r o a t from 140 t o 180 mm. These figures may be m i s l e a d i n g because of t h e a d d i t i o n a l (+) growth. I n a subsequent paper, Fuss (1982) i n d i c a t e d more a c c u r a t e l y t h a t Age I f i s h were 70-77 mm and Age I1 f i s h The l a t e r work were 100-113 mm. involved more data and better techniques. The growth r a t e i n more northern streams i s slower than in the more southern streams (Johnston and Mercer 1976). Cutthroat t r o u t can l i v e a maximum of 10 years (Jones 1976), b u t most adult f i s h runs are made up of individuals between 3 and 6 years of age (Johnston and Mercer 1976; Fuss 1982). Repeat spawners returning f o r a fourth o r f i f t h spawning a t t a i n lengths of 17 to 19 inches (Giger 1972; Johnston and Mercer 1976). and Northcote 1971; Northcote 1972). Schultz and Habitat s e l e c t i o n by juveni 1e cutt h r o a t t r o u t appears t o be governed by the presence of o t h e r salmonid species. In t h e absence of competing s p e c i e s , c u t t h r o a t t r o u t have been found t o p r e f e r pools (Glova and Mason 1976; Johnston and Mercer 1976; Glova 1984). When sympatric with coho salmon, social dominance i s asserted by the coho salmon due t o t h e i r e a r l i e r emergence and l a r g e r body s i z e (Glova and Mason 1976; Johnston 1982). ECOLOGICAL ROLE The use of i s o l a t e d headwater streams by cutthroat trout for spawning and the f i r s t years of rearing serves t o reduce hybridization interactions with other salmonids, primarily steelhead t r o u t and coho salmon, which t y p i c a l l y spawn and r e a r further downstream from nursing zones of young-of-the-year and Age I cutthroat trout (Johnston 1982). Hybridization between c u t t h r o a t t r o u t and hatchery rainbow t r o u t does not occur in the few areas of overlap (Sumner 1972; Campton 1980; Campton and Utter 1985). Sympatric and a l l o p a t r i c populations of r e s i d e n t and anadromous c u t t h r o a t t r o u t may a l s o l i v e within a watershed (DeWitt 1954; Royal 1972; Scott and Crossman 1973; Moring and Lantz 1975; Jones 1979; Johnston 1982). Another s e l e c t i v e advantage for i s o l a t i o n of coastal cutthroat t r o u t , according t o Johnston (1982), i s in the outcome of social interactions with anadromous steelhead t r o u t and coho salmon. Competitive interactions f o r food and space between anadromous c u t t h r o a t t r o u t and steelhead trout or coho salmon juveniles a r e most frequently decided i n favor of the steelhead t r o u t o r t h e coho salmon (Glova and Mason 1976, 1977; Johnston 1982; Glova 1984). Coastal cutthroat t r o u t in B r i t i s h Columbia also have evolved both a spatial and feeding segregation where they cohabit with the predacious Dolly Varden (Salvelinus malma) (Andrusak Cutthroat and rainbow t r o u t coexist i n the same reaches of many streams (Moring and Youker 1979; Nicholas 1978), and occasionally they hybridize (Campton and Utter 1985). The upper reaches of the stream a r e usually dominated by c u t t h r o a t t r o u t , while rainbow t r o u t dominate the lower sections (Hartman and Gill 1968; Nicholas 1978). I areas where anadromous rainbow t r o u t (steelhead) e x i s t , t h i s arrangement does not always occur because the steelhead juveniles emigrate a t Age I and Age I1 i n the spring, which leaves available the pools t h a t the o l d e r c u t t h r o a t - t r o u t seem t o p r e f e r i n a l l sections of t h e streams (Johnston and Mercer 1976). I t has been pointed out t h a t t h e r e a r e t h r e e d i s t i n c t 1i fe history types of coastal c u t t h r o a t t r o u t : only one of these i s anadromous, while the o t h e r two types spend t h e i r e n t i r e 1ives in freshwater (Thomasson 1978; Michael 1983; Fuss 1982, 1984). The r e s u l t of i n t e r a c t i o n with e i t h e r coho salmon o r rainbow t r o u t tends t o be the displacement of the juvenile young-of-the-year cutthroat t r o u t from the preferred pools to the The cutthroat trout riffles. generally return t o the pools a f t e r f a l l i n g water temperatures reduce aggressive i n t e r a c t i o n with these other species and the heavier winter flows threaten displacement from the r i f f l e s (Bustard and - Narver 1975; Glova and Mason 1976). Cutthroat t r o u t are opportunistic feeders (Behnke and Zarn 1976; Wydoski and Whitney 1978). Aquatic i n s e c t s , general l y t h e most a v a i l a b l e food i n streams, a r e t h e dominant i t e m i n most c u t t h r o a t t r o u t d i e t s (Dimick and Mote 1934; Lowery 1966; Allen 1969; Carlander 1969; B a x t e r and Simon 1970; S c o t t and Crossman 1973; Hunter 1973; G r i f f i t h 1975; Glova 1984). Other foods, such as zooplankton (McAfee 1966; Carlander 1969; T r o j n a r and Behnke 1974 ; Hickman 1977), t e r r e s t r i a l i n s e c t s (Lowry 1966; Glova 1984; Martin 1984), and fish (Carlander 1969; Hunter 1973; M a r t i n 1984) are important 1ocal l y or seasonally. Many o f t h e coastal c u t t h r o a t stocks t i m e t h e i r runs t o c o i n c i d e w i t h t h e a v a i 1a b i 1it y of salmon eggs i n t h e streams (Johnston 1982). Cutthroat trout become more p i s c i v o r o u s as t h e y increase i n s i z e ( I d y l l 1942; McAfee 1966; Carlander 1969; B a x t e r and Simon 1970; Wydoski and Whitney 1978). I n freshwater, c u t t h r o a t t r o u t p r e y on threespine s t i c k l e b a c k (Gasterosteus aculeatus), sockeye salmon (Oncorhynchus nerka), and coho salmon (Lowery 1966; Armstrong 1971). There i s considera b l e s i m i l a r i t y and o v e r l a p i n t h e d i e t o f young c u t t h r o a t t r o u t and young coho salmon, b u t d i r e c t c o n f l i c t may be avoided by h a b i t a t p a r t i t i o n i n g (Glova 1984). Wilzbach (1985) i n d i cated t h a t food a v a i l a b i l i t y i s rather i m p o r t a n t i n d e t e r m i n i n g abundance and microhabi t a t d i s t r i b u t i o n o f a d u l t c u t t h r o a t t r o u t w i t h i n a stream. Freshwater p r e d a t o r s o f c u t t h r o a t f r y i n c l u d e rainbow t r o u t , brook t r o u t (Sa'lvelinus f o n t i n a l i s ) , D o l l y Varden, and shorthead sculpins (Cottus confusus), as w e l l as a d u l t c u t t h r o a t t r o u t (Horner and B j o r n n 1976). Other possible predators are great blue be 1 t e d herons (krdea herodias j , k i n g f i s h e r s (Ceryle a l c y o n ) , and mink (Mustela v i son) a c c o r d i n g t o Horner and B j o r n n (1976). I n t h e marine environment, c u t t h r o a t t r o u t feed on gammarid amphipods, sphaeromid isopods, callianassid shrimp, immature c r a b s , and v a r i o u s fish, including chum salmon ; (0ncorh;nchus, gorbu&ha) and P a c i f i c sand lance (Ammodytes hexapterus) .. (Armstrong 1971; Simenstad and K i nney 1978); h e r r i n g and s c u l p i n s a r e a l s o eaten. Predators of sea-run c u t t h r o a t t r o u t i n t h e marine environment i n c l u d e P a c i f i c hake ( M e r l u c c i u s productus), spiny dogfish (Squalus a c a n t h i a s ) , harbor seal (Phoca v i t a m a d a d u l t salmon ( ~ i T 1 9 7 7 ENVIRONMENTAL REQUIREMENTS Temperature Unusual stream temperatures can l e a d t o disease o u t b r e a k s i n m i g r a t i n g f i s h , a l t e r e d t i m i n g o f m i g r a t i o n , and accelerated o r retarded maturation (Reiser and B j o r n n 1979). Most stocks o f anadrornous salmonids have evolved to take advantage o f temperature p a t t e r n s i n t h e i r home streams, and s i g n i f i c a n t a b r u p t d e v i a t i o n s from t h e normal p a t t e r n can a d v e r s e l y a f f e c t t h e i r survival, Cutthroat t r o u t are not usually found i n waters where t h e maximum temperature c o n s i s t e n t l y exceeds 22 OC, although they tolerate brief p e r i o d s o f daytime w a t e r temperatures as h i g h as 26 O C i f c o n s i d e r a b l e c o o l i n g takes p l a c e a t n i g h t (Behnke and Zarn 1976). Be1 1 (1973) r e p o r t e d p r e f e r r e d temperatures o f 9 t o 12 O C f o r c u t t h r o a t t r o u t , w i t h spawning temperatures f o r c u t t h r o a t t r o u t t h a t range from 6 t o 17 O C (Hunter 1973). egg i n c u b a t i o n The duration o f v a r i e s and i s dependent upon temperature (Merriman 1935); t h e optimum temperature f o r i n c u b a t i o n i s approxi m a t e l y 10 t o 11 O C (Merriman 1935; Snyder and Tanner 1960). Egg s u r v i v a l ' POPULATION CONSUMPTION TOTALSTOCK BIOMASS 0 WATER TEMPERATURE FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DATE F i g u r e 4. Sea-run c u t t h r o a t t r o u t p o p u l a t i o n consumption and t o t a l s t o c k biomass (wet weight) versus temperature i n a 500-m l o n g s e c t i o n o f Bear Creek, Washington (modi f i e d from M a r t i n 1984). is a l s o dependent upon (Smith e t a l . 1983). temperature Metabolic r a t e s o f j u v e n i l e c u t t h r o a t t r o u t are h i g h e s t between 11 and 2 1 "C (Owyer and Kramer 1975). According t o M a r t i n (1984), s t o c k biomass remained n e a r l y c o n s t a n t i n a g i v e n s e c t i o n o f a small stream i n t h e Pacific Northwest from July to September, b u t the water temperature rose and food consumption increased p a r a l l e l t o t h i s r i s e ( F i g u r e 4). The optimal temperature f o r j u v e n i l e s i s 15 O C , and e q u i l i b r i u m and a b i l i t y t o swim i s l o s t between 28 and 30 C' (Heath 1963). I n a s t u d y i n southeast Alaska, outm i g r a t i o n o f c u t t h r o a t t r o u t peaked a t w a t e r temperatures between 4 t o 6 C' and i n m i g r a t i o n began a t 10 t o 12 O C and peaked a t 9 t o 10 O C (Jones 1977). S u b s t r a t e and Cover There i s c o n s i d e r a b l e v a r i a t i o n i n size (gravel suitable substrate diameter) f o r c u t t h r o a t t r o u t ; they have been observed spawning i n g r a v e l o f 2 t o 5 cm (June 1981), 0.6 t o 10.2 cm (Hunter 1973), and 0.16 l o 0.64 cm (Hooper 1973). Hanson (1977) found 1to 4-year-old cutthroat trout in streams a s s o c i a t e d w i t h s u b s t r a t e s o f 5 t o 30 cm. I r v i n g and Bjornn (1984) found t h a t survival o f cutthroat embryos increased as t h e percentage o f f i n e sediments decreased. The s i z e o f a d u l t c u t t h r o a t t r o u t a f f e c t s gravels i z e s e l e c t i o n and t h e t a i l s o f pools i n small t r i b u t a r y streams a r e u s u a l l y chosen f o r spawning (Gohnston and Mercer 1976). Cover f o r f i s h can be provided by overhanging vegetation, undercut banks, and submerged o b j e c t s such as l o g s and rocks, as w e l l as by water depth and t u r b u l e n c e (Giger 1972; Bustard and Narver 1975; Reiser and Bjornn 1979). Cover p r o t e c t s t h e f i s h from d i s t u r b a n c e and p r e d a t i o n and a l s o p r o v i d e s shade. The biomass o f c u t t h r o a t t r o u t i n streams increases w i t h t h e amount o f cover present (Wesche 1974; Lantz 1976; Nickelson and R e i s e n b i c h l e r 1977). O i s s o l ved Oxygen .. Ooudoroff and Shumway (1970) r e p o r t e d t h a t salmonids t h a t hatched a t low dissolved-oxygen l e v e l s tended t o be weak and s m a l l , t o develop s l o w l y , and t o show increased abnormal it i e s . C u t t h r o a t t r o u t general l y a v o i d water w i t h l e s s t h a n 5 mg/l o f dissolved oxygen in the summer (Tro j n a r 1972; Sekul ic h 1974). Doudoroff and Shumway (1970) a1so demonstrated t h a t b o t h swimming speed and growth r a t e s o f salmonids d e c l i n e d as d i ssol ved-oxygen l e v e l s decreased. Turbidity According to Reiser and Bjornn (1979), salmonid f i s h e s cease movement o r m i g r a t i o n i n streams w i t h very h i g h s i l t loads (>4,000 mg/l). Because more r a d i a t i o n i s absorbed by t u r b i d water than by c l e a r water, a thermal b a r r i e r t o movement and m i g r a t i o n may develop (Reiser and B j o r n n 1979). Bachman (1958) r e p o r t e d t h a t c u t t h r o a t t r o u t a d u l t s stopped f e e d i n g and moved t o cover a t t u r b i d i t i e s above 35 ppm. Suspended sediment l e v e l s exceeding 103 ppm, combined w i t h d i s s o l v e d oxygen c o n c e n t r a t i o n s l e s s than 6.9 mg/l and v e l o c i t i e s i n t h e redd o f less than 55 cm/h, can reduce egg s u r v i v a l t o below 10% ( B i a n c h i 1963). F r y emergence from t h e g r a v e l may be hindered by excessive sand and s i l t (Reiser and B j o r n n 1979). These conditions may also limit the production o f benthic invertebrates necessary for optimum r e a r i n g o f juvenile fish (Reiser and B j o r n n 1979). Water V e l o c i t y F r y i n stream environments p r e f e r shal lower water and slower v e l o c i t i e s than o t h e r c u t t h r o a t trout 1if e stages ( M i l l e r 1957; Horner and B j o r n n 1976). V e l o c i t i e s o f l e s s t h a n 0.30 m/s a r e p r e f e r r e d and t h e optimum i s l e s s than 0.08 m/s (Horner and B j o r n n 1976). Juveni l e c u t t h r o a t t r o u t i n streams a r e most o f t e n found i n v e l o c i t i e s o f 0.25 t o 0.50 m/s (T.E. Nickelson, unpubl. data). Thompson (1972) found young-of-the-year and l - y e a r - 0 1 d c u t throat t r o u t i n water v e l o c i t i e s o f 0.5 t o 0.6 m/s. Hanson (1977) found l - y e a r - o l d f i s h i n v e l o c i t i e s averaging 0.10 m/s. Hanson (1977) a l s o found 2-, 3-, and 4 - y e a r - o l d c u t t h r o a t t r o u t a t mean v e l o c i t i e s o f 0.14, 0.20, and 0.14 m/s, respectively. C u t t h r o a t t r o u t have been found spawning i n small streams w i t h f l o w r a t e s as low as 0.01-0.03 m3/s (Dimick and M a r r y f i e l d 1945; Wyatt 1959; Moring and Youker 1979). Spawning may occur i n s e l e c t e d areas o f l a r g e streams (Moring and Youker 1979). C u t t h r o a t t r o u t spawning occurs i n water v e l o c i t i e s r a n g i n g from 0 . 1 1 t o 0.56 m/s i n Washington (Hunter 1973) and 0.30 t o 0.90 m/s i n northern Cal if o r n i a (Haoper 1973). Coastal c u t t h r o a t t r o u t p r e f e r t o spawn i n the headwater t r i b u t a r i e s o f l a r g e r streams. Summer water f l o w r a t e s i n these n a t a l creeks seldom exceed 0.28 m3/s during low flow and most average 0.12 m3/s (Johnston r e a r i n g h a b i t a t (Moring 1974; Moring 1975). Water Depth THE FISHERY C u t t h r o a t t r o u t spawn a t t h e t a i l ends of pools a t depths a s shallow a s 10-15 cm (June 1981) o r a s deep a s 1 m (Johnston and Mercer 1976). Hunter (1973) found c u t t h r o a t spawning i n water 6-40 cm deep. June (1981) found some redds i n r i f f l e a r e a s . Angling f o r sea-run c u t t h r o a t t r o u t o c c u r s i n both t h e marine and freshwater environments (Hi s a t a 1973; Johnston and Mercer 1976; Washington 1977; Mercer 1982). Although only l i m i t e d d a t a a r e a v a i l a b l e on t o t a l annual h a r v e s t and f i s h i n g p r e s s u r e , t h e sea-run c u t t h r o a t t r o u t f i s h e r y r e p r e s e n t s an important component of t h e s p o r t f i s h e r i e s of the Pacific Northwest. 1982 ) . In streams, a d u l t c u t t h r o a t t r o u t a r e found i n t h e deeper pools and slower v e l o c i t y w a t e r , while t h e f r y a r e found i n shallower, f a s t e r a r e a s (Gri f f i t h 1972; June 1981). Timber Harvest Logging p r a c t i c e s can d i r e c t l y and indirectly change the physical environments in t h e small headwater streams that are the important spawning and r e a r i n g a r e a s f o r c o a s t a l cutthroat trout. Such changes can s i g n i f i c a n t l y a1 t e r c u t t h r o a t t r o u t populations (Hall and Lantz 1969; Moring 1975; Lantz 1976). Poor logging p r a c t i c e s i n t h e v i c i n i t y of small streams can cause a v a r i e t y of changes t h a t a r e detrimental t o t h e c u t t h r o a t trout. These include (1) an i n c r e a s e i n both t h e maximum and t h e d i u r n a l f 1u c t u a t i o n o f t h e water temperature (Hal 1 and Lantz 1969; Moring 1975; R i n g l e r and Hall 1975; Chamberlin 1982); (2) decreases i n d i s s o l v e d oxygen i n both s u r f a c e and i n t r a g r a v e l w a t e r (Hall and Lantz 1969; Moring 1975; Ringler and Hall 1975; Chamberl i n 1982); ( 3 ) increased s e d i ments i n t h e stream ana cnanges i n t h e amount of f i n e s p r e s e n t i n t h e gravel (Hall and Lantz 1969; Moring 1975; Chamberlin 1982); (4) increased stream flow r e s u l t i n g i n v e l o c i t y and depth changes (Moring 1975; Chamberl i n 1982); ( 5 ) a l t e r a t i o n of t h e amount of woody d e b r i s in t h e stream causing pH changes (Chamberlin 1982); and ( 6 ) 1 o s s of terrestrial and a q u a t i c v e g e t a t i o n used a s food sources and and Lantz Angler s u r v e y s conducted by t h e U. S . National Marine F i s h e r i e s Service i n 1974 i n t h e marine a r e a of Puget Sound, Washington, indicated t h a t c u t t h r o a t t r o u t were second only t o salmon i n p o p u l a r i t y among s a l t w a t e r a n g l e r s (Johnston and Mercer 1 9 7 6 ) . An e a r l i e r Washington Department of Game survey i n 1964 showed a t o t a l of 43,700 a n g l e r s f i s h e d f o r anadromous cutthroat in marine areas of Washington S t a t e (Johnston and Mercer 1976). The average per capita e x p e n d i t u r e was $25 p e r day i n 1974, with t h e t o t a l value t o the S t a t e ' s economy e s t i m a t e d a t $8,850,000 per year. Although t h i s d a t a may have been b i a s e d , t h e r e i s no reason t o b e l i e v e a d e c l i n e i n p a r t i c i p a t i o n has o c c u r r e d i n t h e marine f i s h e r y . The t o t a l p a r t i c i p a t i o n i n and value of t h e f r e s h w a t e r f i s h e r y f o r anadromous c u t t h r o a t i n Washington has never been evaluated. However, over 1,800 man-days were expended in 1970 pursuing c u t t h r o a t t r o u t in saltwater on s o u t h e r n Hood Canal (Johnston and Mercer 1976). I n Oregon i n 1972, a n g l e r s caught an e s t i m a t e d L26,000 a d u l t c u t t h r o a t , 86% from c o a s t a l streams and 14% from the Columbia River and i t s t r i b u t a r i e s (Johnston and Mercer 1976). They e s t i m a t e d t h e annual value of t h e stream c u t t h r o a t t r o u t f i s h e r y t o Oregon's economy was $3,750,000, based on an average p e r c a p i t a e x p e n d i t u r e estimated a t $30 p e r day i n Washington S t a t e by O l i v e r e t a l . (1975). The p r o j e c t e d c a t c h i n Washington and i t s impact on t h e S t a t e ' s economy has been assumed t o be g r e a t e r t h a n i n Oregon due t o grea'ter numbers o f streams i n Washington S t a t e t h a t c o n t a i n c u t t h r o a t t r o u t and a l a r g e r human p o p u l a t i o n c l o s e t o these streams (Johnston and Mercer 1976). Tipping (1981) i n d i c a t e d t h a t 84% o f a l l sea-run c u t t h r o a t t r o u t were caught by anglers t a r g e t i n g on them, w h i l e 16% were caught i n c i d e n t a l t o salmon and steelhead. Most a n g l i n g e f f o r t was by p l u n k e r s , w i t h b o a t e r s second, and d r i f t fishermen t h i r d (Tipping and S p r i n g e r 1980; T i p p i n g 1981). Overall cutthroat trout a n g l i n g e f f o r t i s h i g h e s t i n August ( T i p p i n g and S p r i n g e r 1980), b u t t h e h i g h e s t catch was i n t h e f i r s t 2 weeks of September ( T i p p i n g 1981). Most anglers caught one f i s h . o r l e s s p e r trip (Tipping and S p r i n g e r 1980; T i p p i n g 1981). The m a j o r i t y o f f i s h caught were s e x u a l l y mature ( T i p p i n g 1981). The amount of a n g l e r p a r t i c i p a t i o n 1 i k e l y double between 1972 and 1990 i n Oregon and a s i m i l a r t r e n d may occur i n Washington (Johnston and will The v a l u e o f t h e Mercer 1976). c u t t h r o a t f i s h e r y t o b o t h Oregon and Washington economies i s expected t o i n c r e a s e along w i t h f i s h i n g pressure; however, i t i s a n t i c i p a t e d t h a t t h e n a t i v e stocks o f sea-run c u t t h r o a t w i 11 n o t increase and may d e c l jne due t o h a b i t a t d e g r a d a t i o n and p o s s i b l e overfishing (Johnston and Mercer 1976). New r e s t r i c t i v e r e g u l a t i o n s ( 2 f i s h p e r day and 1 2 - i n c h minimum size) w i t h i n B r i t i s h Columbia and Washington, which were designed t o increase spawner escapement, have s i g n i f i c a n t l y increased t h e abundance of older, larger fish in many watersheds (J. M. Johnston, unpubl . data). Johnston (1980) i n d i c a t e d t h a t a two-fish l i m i t i n the Stillaguamish R i v e r would reduce t h e t o t a l h a r v e s t o f sea-run c u t t h r o a t t r o u t by about 20%. A 1 4 - i n c h minimum s i z e l i m i t on sea-run cutthroat caught i n a1 1 marine waters and s e l e c t e d f r e s h w a t e r streams was put in force in Washington S t a t e i n 1987. Concurrent w i t h h a b i t a t p r o t e c t i o n , agencies r e s p o n s i b l e f o r t h e rnanagement o f sea-run c u t t h r o a t p o p u l a t i o n s have attempted t o spread t h e h a r v e s t a b l e p o r t i o n o f t h e s t o c k s t o more a n g l e r s b y imposing r e s t r i c t i o n s on c a t c h and possession, s i z e , season, and gear. LITERATURE CITED 1969. L i m i t a t i o n s on A l l e n , K.R. p r o d u c t i o n i n salmonid p o p u l a t i o n s i n streams. Pages 3-18 i n T.G. Northcote, ed. Symposium on-salmon and trout in streams. .. - H. R. MacMi 1l a n Lectures i n F i s h e r i e s , University of 0 r i t i s h Columbia, Vancouver. B,G. , and 0.W. Narver. Anderson, 1975. F i s h p o p u l a t i o n s o f C a r n a t i o n Creek and other B a r k l e y Sound s treams--1974: d a t a r e c o r d and progress r e p o r t . F i s h Res. Board Can. MS Rep. No. 1351. 73 pp. Andrusak, H. , and T.G. Northcote. 1971. 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Oregon State sity, Corvallis. 60 pp. R., and R.W. Whitney. 1978. Inland f i s h e s of Washington. Universi ty of Washington Press, Seattle. 220 pp. Wydoski, Wyatt, B. 1959. Observations on the movements and reproduction of the Cascade form of cutthroat trout. Univer- 5 4. Title and Subtitle Species P r o f i l e s : L i f e H i s t o r i e s and Environmental Requirements o f Coastal fishes and I n v e r t e b r a t e s ( P a c i f i c ~ o r t h w e s t ) - - S e a - r u n cutthroat trout ncwn 0.t. January 1989 L 7. AutMttr) G i l b e r t 0. Paulev. Kevin Oshima, Karen L. Bowers. and Garv L. Thomas ( 10. 9. h r l o r m i y Omanltmtion Heme and Address Washington Cooperative Fishery Research U n i t U n i v e r s i t y o f Washington S e a t t l e , WA 98195 I Z ~ m n r o & a OmmlUtlon Nam* PmtuWTasklWw* Und N ; 11. Conrmct(C> or CramlGI No. (Cl and Addnss U.S. Department o f t h e I n t e r i o r F i s h and W i l d l i f e Service Research and Development N a t i o n a l Wetlands Research Center Washington, DC 20240 U.S. Army Corps o f Engineers Watarways Experiment S t a t i o n P.O. Box 6 3 1 Vicksburg, MS 39180 I 15. Supplmumary Notn *U.S. Army Corps o f Engineers Report No. TR EL-82-4 Species p r o f i l e s are 1 i t e r a t u r e summaries o f t h e taxonomy, morphology, range, 1i f e h i s t o r y , and environmental requirements o f c o a s t a l a q u a t i c species. They a r e designed t o a s s i s t w i t h environmental impact assessments. The sea-run c u t t h r o a t t r o u t (Salmo c l a r k i c l a r k i ) i s anadromous and i s found i n most c o a s t a l streams from Southeast Alaska t o A t h r i v i n g s p o r t f i s h e r y e x i s t s b o t h i n these freshwater streams Northern C a l i f o r n i a . and i n p r o t e c t e d marine waters. Many o f t h e small streams c o n t a i n i n g c u t t h r o a t t r o u t can be adversely a l t e r e d by poor timber operations. Optimum temperature f o r i n c u b a t i n g i s 10 t o II O C and eggs hatch i n 28-40 days. Optimum temperature f o r j u v e n i l e s i s 15 O C and a b i l i t y t o swim i s l o s t a t about 28 OC. A d u l t s p r e f e r f a i r l y slow-moving water w i t h p l e n t y o f cover. Spawning occurs i n smal 1 diameter g r a v e l . L i k e s t e e l head t r o u t , they may r e t u r n from s a l t w a t e r t o spawn several times. Temperature Estuaries Animal migrations 0. Identlhm/+n.End~a Feeding h a b i t s Trout L i f e cycles Tqrrn. Sea-run c u t t h r o a t t r o ut Coastal c u t t h r o a t t r o u t Salmo c l a r k i c l a r k i --- Ecological r o l e Environmental requirements Timber h a r v e s t