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REFERENCE COPY Do Not Remove from the Library U. S. Fish and Wildlife Service IYQTI:?~~~ kVetlands Kesearch Center -rR EL-82-4 Biological Report 82(11.88) January 1989 7G.3 Caiun Dome Boulevard La favette, Louisiana 70506 Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Northwest) PINK SALMON Fish and Wildlife Service Coastal Ecology Group Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers

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Page 1: SALMON - USGS National Wetlands Research Center

REFERENCE COPY Do Not Remove from the Library

U. S. Fish and Wildl i fe Service I Y Q T I : ? ~ ~ ~ kVetlands Kesearch Center -rR EL-82-4 Biological Report 82(11.88)

January 1989 7G.3 Caiun Dome Boulevard La favette, Louisiana 70506

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Northwest)

PINK SALMON

Fish and Wildlife Service Coastal Ecology Group

Waterways Experiment Station

U.S. Department of the Interior U.S. Army Corps of Engineers

Page 2: SALMON - USGS National Wetlands Research Center

B i o l o g i c a l Report 82(11.88) 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 F ishes and I n v e r t e b r a t e s ( P a c i f i c Northwest)

PINK SALMON

S c o t t A. Bonar, G i l b e r t B. Pauley, and G. L. Thomas Washington Cooperat ive F ishery 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

P r o j e c t O f f i c e r David Moran

U.S. F i s h and W i l d l i f e Serv ice Nat iona l Wetlands Research Center

1010 Gause Boulevard S l i d e l l , LA 70458

Performed f o r Coastal Ecology Group

Waterways Experiment S t a t i o n U. S. A r m y Corps o f Engineers

Vicksburg, MS 39180

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 Serv ice Research and Development

Na t i ona l Wetlands Research Center Washington, DC 20240

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T h i s s e r i e s may be re fe renced as f o l l ows :

U. S. F i s h and W i l d l i f e Serv ice. 1983-19-. Species p r o f i l e s : l i f e h i s t o r i e s and envi ronmenta l requi rements o f coas ta l f i s h e s and i n v e r t e b r a t e s . U. S. F i s h W i l d l . Serv. B i o l . Rep. 82(11). U.S. Army Corps o f Engineers, TR EL-82-4.

T h i s p r o f i l e may be c i t e d as f o l l o w s :

Bonar, S.A., G.B. Pauley, and G.L. Thomas. 1989. Species p r o f i l e s : l i f e h i s t o r i e s and envi ronmenta l requi rements o f coas ta l f i s h e s and i n v e r t e b r a t e s ( P a c i f i c Nor thwest) - -p ink salmon. U. S. F i s h W i l d l . Serv. B i o l . Rep. 82(11.88). U.S. Army Corps of Engineers, TR EL-82-4. 18 pp.

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PREFACE

Th is species p r o f i l e i s one o f a se r i es on coasta l aquat ic organisms, p r i n c i p a l l y f i s h , o f spor t , commercial, o r eco log ica l importance. The p r o f i 1 es are designed t o prov ide coasta l managers, engineers, and b i o l o g i s t s w i t h a b r i e f comprehensive sketch o f t he b i o l o g i c a l c h a r a c t e r i s t i c s and environmental requirements o f t h e species and t o descr ibe how popu la t ions o f ' t h e species may be expected t o r e a c t t o environmental changes caused by coasta l development. Each p r o f i l e has sect ions on taxonomy, 1 i f e h i s t o r y , eco log ica l r o l e , environmental requirements, and economic importance, i f appl i cab le . A t h ree - r i ng b inder i s used f o r t h i s ser ies so t h a t new p r o f i l e s can be added as they are prepared. Th is p r o j e c t i s j o i n t l y planned and f inanced by t h e U. S. Army Corps o f Engineers and t h e U.S. F i sh and W i l d l i f e Service.

Suggestions o r questions regarding t h i s r e p o r t should be d i r e c t e d t o one of t he f o l l owing addresses.

I n fo rma t ion Trans fer S p e c i a l i s t Nat ional Coastal Ecosystems Team U.S. F i sh and W i l d l i f e Serv ice NASA-Sl i d e l 1 Computer Complex 1010 Gause Boulevard S l i d e l l , LA 70458

U. S. Army Engineer Waterways Experiment S t a t i o n A t ten t i on : WESER-C Post O f f i c e Box 631 Vicksburg, MS 39180

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CONVERSION TABLE

M e t r i c t o U. S . Customary

To Obta in M u l t i p l y

m i l l i m e t e r s (mm) cent imeters (cm) meters (m) meters (m) k i 1 ometers (km) k i 1 ometers (km)

inches inches f e e t fathoms s t a t u t e m i l e s n a u t i c a l m i l e s

square meters (m2) 10.76 square k i 1 ometers ( km2) 0 .3861 hectares (ha) 2 .471

square f e e t square m i l e s acres

l i t e r s (1) cub ic meters (m3) cub ic meters (m3)

ga l l ons cub ic f e e t a c r e - f e e t

m i l l i g r a m s (mg) grams (g) k i 1 ograms (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

B r i t i s h thermal u n i t s Fahrenhei t degrees

k i 1 o c a l o r i e s (kca l ) Ce ls ius degrees ( O C )

U . S . Customary t o M e t r i c

25.40 2 .54 0.3048 1.829 1.609 1.852

inches inches f e e t ( f t ) fathoms s t a t u t e m i l e s ( m i ) n a u t i c a l m i l e s (nmi)

m i 11 imete rs cen t imete rs meters meters k i 1 ometers k i 1 ometers

square f e e t ( f t 2 ) square m i l e s (mi2) acres

square meters square k i lomete rs hectares

gal 1 ons (ga l ) cubic f e e t ( f t 3 ) ac re - fee t

l i t e r s cub ic meters cub ic meters

ounces (02) ounces (02) pounds ( l b ) pounds ( l b ) s h o r t tons ( ton )

m i l l i g r a m s grams k i 1 ograms m e t r i c tons m e t r i c tons

B r i t i s h thermal u n i t s (Btu) Fahrenhei t degr,ees (OF)

k i l o c a l o r i e s Ce ls ius degrees

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CONTENTS

Page

CONVERSION TABLE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii PREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i v ACKNOWLEDGMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v i

NOMENCLATURE/TAXONOMY/RANGE . . . . . . . . . . . . . . . . . . . . . . . . MORPHOLOGY/IDENTIFICATION AIDS . . . . . . . . . . . . . . . . . . . . . . REASON FOR INCLUSION I N SERIES . . . . . . . . . . . . . . . . . . . . . . LIFE HISTORY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Spawning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F e c u n d i t y . Eggs. A l e v i n s . and F r y . . . . . . . . . . . . . . . . . . . . M a r i n e s t a g e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

GROWTH CHARACTERISTICS . . . . . . . . . . . . . . . . . . . . . . . . . . THE FISHERY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ECOLOGICAL ROLE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 ENVIRONMENTAL REQUIREMENTS . . . . . . . . . . . . . . . . . . . . . . . . 1 0

Water Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 S a l i n i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 D i s s o l v e d O x y g e n . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 S u b s t r a t e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Water Depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Water V e l o c i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 T u r b i d i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

LITERATURECITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3

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ACKNOWLEDGMENTS

We g r a t e f u l l y acknowledge reviews by Robert F. Donnelly and Steven S. Parker , both of the F isher ies Research I n s t i t u t e , U n i v e r s i t y o f Washington, S e a t t l e .

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F igu re 1. The p i n k salmon, showing extreme sexual dimorphism o f t h e male ( lower f i s h ) . Repr in ted w i t h permiss ion o f S c o t t and Crossman (1973).

PINK SALMON

S c i e n t i f i c name . . . . . . . . . . . Oncorh nchus gorbuscha (Wal baum) &

P r e f e r r e d common name.. ... P ink salmon .. Other common names.. humpback, hump-

back salmon, humpy ( S c o t t and Cross- man 1973)

Class . . . . . . . . . . . . . . . . . . . . . Oste ichthyes Order . . .................. Salmoniformes Fami ly . . . . . . . . . . . . . . . . . . . . . . Salmonidae

Geographic range: Anadromous i n r i v - e r s and smal l streams from no r t he rn C a l i f o r n i a n o r t h t o t h e a r c t i c waters o f Alaska, Canada, and t h e S o v i e t Union. Washington S t a t e appears t o be a t t h e southern end o f t h e range f o r streams t h a t suppor t c o n s i s t e n t l y e x p l o i t a b l e spawning runs o f p i n k salmon. Popu la t ions i n As ia occur as f a r south as Hondo I s l a n d i n Japan. P ink salmon have been i n t r oduced i n t o t h e Great Lakes. Major r i v e r s suppo r t i ng p i n k salmon runs i n t h e P a c i f i c Northwest a r e shown i n F i gu re 2. South o f B r i t i s h Columbia, o n l y a few r i v e r s

i n t h e S ta te o f Washington t h a t empty i n t o Puget Sound suppor t ma- j o r p i n k salmon runs. M i g r a t i o n p a t t e r n s o f f i s h e n t e r i n g B r i t i s h Columbia and Washington waters a r e shown i n F i gu re 3. Tagging s t u d i e s have i n d i c a t e d t h a t t h e m a j o r i t y o f a d u l t p i n k salmon des t i ned f o r Puget Sound r i v e r s m ig ra te th rough t h e S t r a i t o f Juan de Fuca a t t h e south- e r n end o f Vancouver I s l a n d , w h i l e those f i s h r e t u r n i n g t o southern B r i t i s h Columbia m ig ra te p r i m a r i l y th rough t h e S t r a i t o f Georg ia on t h e eas te rn s i d e o f Vancouver I s l a n d , w i t h some r e t u r n i n g th rough t h e S t r a i t o f Juan de Fuca (F i gu re 3).

MORPHOLOGY/IDENTIFICATION AIDS

Morphology and i d e n t i f i c a t i o n a i d s a re from H a r t (1973) and S c o t t and Crossman (1973). Dorsal f i n 10- 15 rays; adipose sma l l , s lender , f l eshy ; anal 13-17 rays ; p e l v i c s 10 rays , abdominal, w i t h a f r e e - t i p p e d

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Marine dlstributlon:

commonly abundant

occasionally present

M I L E S

K I L O M E T E R S

1 .-- CALIFORNIA

OREGON

Figure 2. Major r i v e r s i n t he P a c i f i c Northwest support ing p i n k salmon runs.

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B R I T 1 S I I

L U M I 3 I A

F igu re 3. P r i n c i p a l m i g r a t i o n r ou tes (ar rows) o f p i n k salmon e n t e r i n g Puget Sound and B r i t i s h Columbia waters ( f r om Washington S t a t e Department o f F i s h e r i e s 1959).

f l e s h y appendage above i n s e r t i o n ; marg in o f o r b i t . Males on spawning p e c t o r a l s 15 rays. C y c l o i d sca les m i g r a t i o n develop a prominent h i g h sma l l , 150 t o 205 on l a t e r a l l i n e , hump immediate ly beh ind t h e head which e a s i l y d i s t i n g u i s h e s p i n k (F i gu re 1). Juven i l es have no p a r r salmon f rom o t h e r spec ies o f salmon. marks, a r e b l u e t o g reen i sh a l ong back G i l l r a k e r s 26 t o 34 on f i r s t g i l l and s i l v e r y on s ides . D i s t i n g u i s h a b l e arch. Body e longa te and somewhat l a t - f rom o t h e r salmon by smal l sca les . era1 l y compressed.

I d e n t i f i c a t i o n a i ds : Ta i 1 (caudal REASON FOR INCLUSION I N SERIES f i n ) moderate ly fo rked . Numerous l a r g e , b l ack , ova l spots on upper P i n k salmon a re t h e most abundant s ides and back and a l l o f caudal f i n . o f t h e spec ies o f P a c i f i c salmon. Upper jaw reaches beyond p o s t e r i o r They spawn i n No r t h American and As ian

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streams b o r d e r i ng t h e P a c i f i c and Arc- t i c Oceans. T h e i r range i n No r t h Amer- i c a i s ex tens i ve ; t h e y a re t h e most numerous salmon i n t h e commercial salmon f i s h e r i e s w i t h a ve r y h i g h r a t e o f e x p l o i t a t i o n by Canadian, Japanese, Russian, and U n i t e d S ta tes commercial f i shermen (A1 exande rsdo t t i r and Math i sen 1983).

LIFE HISTORY

Spawni ng

P ink salmon have a 2-year l i f e c y c l e , which i s so i n v a r i a b l e t h a t f i s h r unn ing i n odd-numbered c a l endar years a r e e f f e c t i v e l y i s o l a t e d f rom even-year f i s h so t h a t no gene f l o w occurs between them ( L a r k i n and R icker 1964; H a r t 1973; Donne l l y 1983). It i s t h e s i m p l e s t and l e a s t v a r i e d l i f e h i s t o r y o f any salmon. Because o f t h i s s i m p l i c i t y , t h e l i f e c y c l e may be diagrammed as a c i r c l e w i t h ve r y 1 it- t l e ove r l ap o f t h e va r i ous stages (F i gu re 4) . Usual age a t m a t u r i t y i s 2 years ; however, 3 -year -o ld specimens have been d iscovered on occas ion (Anas 1959; Turner and B i l t o n 1968; A1 exande rsdo t t i r and Math isen 1983).

The salmon r e t u r n t o t h e i r stream o f o r i g i n t o spawn i n t h e f a l l when t h e wate r temperature ranges f rom 8 t o 14 O C ( P r i t c h a r d 1939; Hunter 1959), and u s u a l l y e n t e r t h e r i v e r on h i g h wate r f r e s h e t s ( P r i t c h a r d 1939). Some p i n k salmon spawn severa l m i l e s up- stream f rom s a l t w a t e r i n a few r i v e r systems ( S c o t t and Crossman 1973), b u t spawning genera l l y takes p l a c e e i t h e r i n f r eshwa te r c l o s e t o t h e sea o r i n t h e i n t e r t i d a l zones. P i n k salmon a r e cons idered t h e most s p e c i a l i z e d o f t h e salmon i n t h e genus .Oncorhynchus be- cause t h e y a r e t h e l e a s t dependent on f reshwate r , r e g u l a r l y spawning i n t h e i n t e r t i d a l areas (Mi 11 e r and Brannon 1982; A1 exande rsdo t t i r and Math i sen 1983).

Males a r e l a r g e r than females. I n genera l , l a r g e r f i s h , p redominan t l y

Dec , Jan

F i gu re 4. Two-year l i f e c y c l e o f p i n k salmon from Kodiak A rch i pe l ago. Dates may vary i n d i f f e r e n t areas a l ong t h e P a c i f i c coast . Shaded areas show ove r l aps i n t i m e ( a f t e r Donne l l y 1983).

males, e n t e r t h e streams f i r s t ( H a r t 1973). Runs may be a l t e r n a t e l y l a r g e and smal l i n consecu t i ve yea rs (Mer re l 1 1962). Spawning usual 1 y oc- cu r s i n l a t e August th rough e a r l y October i n mucn o f t h e range (Sher idan 1962). The female d i g s a depress ion i n t h e g rave l on t h e stream bo t tom i n an area averag ing 0 .6 t o 0 . 9 m i n w i d t h (Wel ls and McNeil 1970), and i n wa te r w i t h an average depth o f 0.15-0.53 m and v e l o c i t y o f 21-100 cm/s (Col l i n g s 1972). The t a i l i s used t o f o r c e wate r down on t h e g rave l t o remove f i n e sediments (W icke t t 1959a). A dominant male guards t h e female d u r i n g t h e d i g g i n g process, a t t a c k i n g compe t i t o r s o r i n t r u d e r s .

Once t h e nes t ( redd) i s dug, t h e female drops i n t o t h e m idd le o f t h e deepest p a r t w i t h a dominant male (Chebanov 1982), and t h e d e p o s i t i o n o f eggs and m i l t occurs . Up t o s i x males have been recorded spawning w i t h a s i n g l e female (W icke t t 1959a).

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Chebanov (1980) observed t h a t , com- pared t o l a r g e r males, sma l l e r males showed l e s s spawning a c t i v i t y and per - formed more movements w i t h i n t h e spawning ground. A f t e r a b r i e f m i l l - i n g , t h e spawning procedure i s repea t - ed upstream f rom t h e nes t , and t h e new d i gg i ngs cover t h e eggs i n t h e p r e v i - ous l y dug redd. The female s tays i n t h e area, p r o t e c t i n g her redds f rom o t h e r d i g g i n g females (Chebanov 1980). When d e n s i t i e s o f spawners a r e h i gh , nes ts a r e sometimes superimposed upon one ano ther and t h e t o t a l number o f eggs s u r v i v i n g i s reduced (McNeil 1967).

Fecund i t y , Eggs, A l ev i ns , and F r y

Mature female p i n k salmon c o n t a i n between 1,500 and 1,900 eggs ( P r i t c h a r d 1937; H a r t 1973). Fecundi- t y i s r e l a t e d p o s i t i v e l y t o s i z e (Foe rs te r and P r i t c h a r d 1941). The eggs a r e l a r g e (about 6 mm i n diame- t e r ) and orange-red. Bo th e x t r a g r a v e l and i n t r a g r a v e l chemical and phys i ca l ( i n c l u d i n g h y d r a u l i c ) f a c t o r s c o n t r o l t h e success o f egg s u r v i v a l . Water p e r c o l a t i n g th rough t h e g rave l p ro - v i des oxygen. Du r i ng t he i n c u b a t i o n pe r i od , s u b s t a n t i a l m o r t a l i t y i n . redds may r e s u l t from f r eez i ng , f l o w f l u c t u - a t i o n s , dewater ing, oxygen reduc t i on , p reda t i on , and m i c r o b i a l i n f e s t a t i o n . I n some ins tances , t h e t o t a l egg mor- t a l i t y may be as h i g h as 75% t o 90% (Krueger 1981).

The l e n g t h o f t h e i n c u b a t i o n p e r i o d i s dependent on wate r temperature (Car l e t a l . 1959). Under n a t u r a l c o n d i t i o n s i n B r i t i s h Columbia, eggs ha tch f rom l a t e December t o l a t e Feb- r u a r y (Neave 1966). Once hatched, t h e a l e v i n s remain i n t h e g rave l f o r sev- e r a l weeks w h i l e t h e yo1 k sac i s ab- sorbed and i nco rpo ra ted i n t o t h e body. Swimming f r y emerge f rom t h e g rave l i n B r i t i s h Columbia streams as e a r l y as l a t e February, b u t peak emergence i n most l o c a l i t i e s occurs d u r i n g A p r i l o r May (Neave 1966).

P i nk salmon m i g r a t e t o t h e sea soon a f t e r emerging f rom t h e g rave l and spend most o f t h e i r l i v e s i n s a l t - water . F r y were abundant i n marsh a rea t i d a l channels o f t h e F raser R i ve r Es tuary f rom March t o June, w i t h peak numbers o c c u r r i n g i n A p r i l and e a r l y May (Levy and Nor thco te 1982). U n l i k e ch inook and chum salmon f ry which r e s i d e t e m p o r a r i l y i n t h e marsh be fo re m i g r a t i n g i n t o t h e P a c i f i c Ocean, p i n k salmon f ry appear t o be o n l y t r a n s i e n t r e s i d e n t s of t h e es tu - a r i n e marsh a rea as t hey make t h e i r r a p i d and a c t i v e m i g r a t i o n s downstream (Levy and No r t hco te 1982). The f ry swim a t t h e su r face , c r e a t i n g r i p p l e s d u r i n g t h e i r downstream m i g r a t i o n when c r u i s i n g speeds o f 0.6 f t / s a r e reached a t temperatures o f 6 t o 7 OC (W icke t t 1959b).

Downstream m i g r a t i o n s u s u a l l y beg in a t n i g h t . B r e t t and A l d e r i c e (1958) r e p o r t e d t h a t m i g r a t i o n s r a r e l y com- menced be fo re t h e i n c i d e n t l i g h t i n t e n s i t y had f a1 l e n below 0 . 1 f o o t cand le a t about 8: 30 t o 8: 45 p.m. , w i t h a peak m i g r a t i o n between 9: 00 and 10: 30 p.m. , and n e a r l y no movement by 12:30 a.m. A s t r o n g l i g h t avoidance r e a c t i o n i s shown i n i t i a l l y by i n d i v i d u a l f r y , b u t s choo l i ng f r y acc l ima te t o t h e l i g h t i n about 15 min and no l onge r a v o i d i t (Hoar 1956).

F r y m i g r a t i n g t o s a l t w a t e r u s u a l l y do n o t feed, b u t i f t h e d i s t a n c e i s g r e a t , t hey may feed on l a r v a l i n s e c t s (McDonald 1960). Both t h e d i s t a n c e t r a v e l e d by t h e f i s h and t h e constancy o f d i r e c t i o n a r e g r e a t e r f o r l a r g e schools than f o r smal l ones (Hoar 1958). The behav io r o f f r y i n l a r g e schools appears t o be more o r d e r l y and p r e c i s e t han i n sma l l schools (Hoar 1958).

E a r l y ha tchery f r y m ig ra te down- s t ream about 35 days ahead o f l a t e ha tchery f r y and about 55 days ahead o f w i l d f r y i n B r i t i s h Columbia ( T a y l o r 1980). L a t e r m i g r a t i n g f r y have cons i de rab l y h i ghe r mar i ne

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s u r v i v a l than e a r l y m ig ra t i ng f r y , poss ib l y because low water tempera- t u res encountered by e a r l y migrat- i n g f r y may slow t h e i r growth i n t he estuary and make them more vu l herabl e t o predators (Tay lor 1980).

A f t e r 1 eavi ng freshwater , t h e young tend t o remain c lose inshore dur ing t h e i r f i r s t summer (Manzer 1956). I n t h e i n t e r t i d a l regions, t he young salmon p r e f e r feeding i n water o f r e l a t i v e l y low s a l i n i t y . Pre fer red food inc ludes var ious i nve r teb ra te eggs, amphipods, and copepods i n Puget Sound (Gerke and Kaczynski 1972; Kaczynski e t a l . 1973), a l though o the r p lank ton i c organisms are eaten e l se- where (Cooney e t a l . 1981). Adu l t p i n k salmon feed p r imar i l y on amphipods, euphausi i d s , and f i s h (LeBrasseur 1966). Godin (1981a) conducted l abo ra to ry t e s t s which showed t h a t 68% o f t h e j u v e n i l e f i s h were most a c t i v e du r ing t h e day. Stomach analyses from f r y showed t h a t most feeding occurs dur ing day1 i g h t hours i n t h e l i t t o r a l zone (Godin 1981b).

M ig ra t i on i n t he sea i s s a l t a t o r y , w i t h sho r t per iods o f a c t i v e m ig ra t i on in te rspersed w i t h 1 onger per iods when the f r y do n o t migra te (Healey 1967). Theories f o r t he c o n t r o l o f marine m ig ra t i on i n c l ude movement toward h igher sa l i n i t i e s (Baggerman 1960; McInerney 1964; Leggett 1977) and the use o f t i d a l f l o w and food grad ien ts (Hurley and Woodall 1968; Leggett 1977). The f i s h begin moving o f f sho re i n l a t e summer (Manzer 1956), i n a manner t h a t appears t o be gradual o r i r r e g u l a r , and 1 eave the young-of- the-year a t a d is tance o f 6-12 m i from the nearest land i n the f a l l (Neave 1966).

Marine Stages

A t the beginning o f t h e i r ocean migra t ions , p i n k salmon from Washing- ton, Oregon, and Southern B r i t i s h Co- lumbia move northward from t h e i r

streams o f o r i g i n i n a narrow b e l t a long the coast (Hart 1973). Substan- t i a l numbers o f f i s h from B r i t i s h Columbia stocks move up t o 500 m i o f fshore , m ig ra t i ng r a p i d l y t o a t t a i n t h a t d is tance (Har t 1973). T ida l cur- ren ts may s t rong l y i n f l uence the d i - r e c t i o n o f movement, as w e l l as o the r o r i e n t a t i o n cues (Leggett 1977). Stocks from southern l oca t i ons tend t o en te r t he ocean f i r s t , fo l lowed by stocks from more nor thern and western l oca t i ons as the season progresses. I n w i n t e r and e a r l y spr ing , t he d i s - t r i b u t i o n o f stocks i s more souther ly i n t he western North P a c i f i c than i n t he eastern North P a c i f i c . This may r e f l e c t t he i n f l uence o f co lde r waters i n t he western suba rc t i c gyre (Royce e t a1 . 1968; Burgner 1980). Ocean migra t ions o f p i n k salmon vary g r e a t l y between populat ions from d i f f e r e n t geographical areas; t he m ig ra t i on o f stocks o f t he eastern North P a c i f i c i s much more r e s t r i c t e d than t h a t o f oth- e r stocks (Burgner 1980).

The v e r t i c a l d i s t r i b u t i o n o f p i n k salmon i n t he ocean i s no t we1 1 known. G i l l n e t and l o n g l i n e catches on which knowledge o f ho r i zon ta l d i s t r i - b u t i o n i s based are usua l l y made w i t h i n 6 t o 7 m o f t he surface. I n t he Gul f o f Alaska, some f i s h have been caught a t depths o f 24 t o 36 m (Manzer and LeBrasseur 1959; Neave 1953, 1966).

A f t e r spending about 18 months a t sea, adu l t s r e t u r n t o t he spawning streams i n p r e d i c t a b l e and h i g h l y segregated even-numbered-year and odd-numbered-year runs. Salmon spawn- i n g migra t ions are thought t o be guided by environmental cues such as 01 f a c t i o n , cur ren ts , temperature, and s a l i n i t y , as we l l as by c e l e s t i a l nav iga t ion o r magnetic o r i e n t a t i o n (Royce e t a l . 1.968; Leggett 1977; Brannon 1982), b u t the exact cause o f the migra t ions i s no t known. The t i m i n g o f t he migra t ions i s va r i ab le . The r a t e o f s t r a y i n g i s much h igher i n p i n k salmon than i n o ther species o f salmon. Th is s t r a y i n g could con-

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s t i t u t e a s u r v i v a l s t r a t e g y t h a t p i n k salmon have evolved e i t h e r t o r e c o l o- n i z e streams which have l o s t a yea r c l a s s due t o env i ronmenta l ca tas t rophe o r t o ensure t h a t a l l f i s h headed f o r an un inhab i t ab le stream a r e n o t e r a d i - cated. Th i s s t r a y i n g mechanism cou ld have evolved w i t h t h e normal tendency o f p i n k salmon t o spawn i n sma l l , env i ronmenta l l y uns tab le streams. Legget t (1977) and Brannon (1982) d iscussed i n d e t a i l t h e numerous t h e o r i e s t h a t have been advanced t o e x p l a i n salmonid homing.

GROWTH CHARACTERISTICS

The use o f sca le annulus f o rma t i on t o determine age i n p i n k salmon has been d iscussed by Pearson (1966). When t h e f ry ha tch and leave t h e sub- s t r a t e , t hey range from 3.2 t o 3.8 cm i n t o t a l l e n g t h from t h e t i p o f t h e snout t o t h e f o r k o f t h e t a i l and weigh approx imate ly 0.3 g ( P r i t c h a r d 1944). The f ry feed r e l a t i v e l y l i t t l e , i f a t a l l , d u r i n g t h e i r down- stream m i g r a t i o n and t hus leave t h e streams a t about t h i s same s i z e ( P r i t c h a r d 1944).

Dur ing t h e f i r s t 30-day p e r i o d a t sea (beg inn ing i n August o r Septem- ber ) , t h e f ry inc rease i n we igh t by a f a c t o r o f 6 (LeBrasseur and Parker 1964). Dur ing t h e i r f i r s t 40 days a t sea, p i n k salmon grow t o about 8 cm and by t h e f o l l o w i n g March reach a l e n g t h o f about 33 cm (Parker 1968). Dur ing t h e l a s t s p r i n g and summer i n t h e sea, growth i s r a p i d from about 0 .5 kg up t o an average we igh t o f be- tween 2 and 4 kg (LeBrasseur and Par- ke r 1964).

THE FISHERY

The p i n k salmon i s l e s s d e s i r a b l e t han o t h e r P a c i f i c salmon i n commer- c i a l and s p o r t catches, because o f i t s smal l s i z e , p a l e f l e s h , and f l a c c i d t e x t u r e . However, i t s s t a t u s as a food i t e m and s p o r t f i s h i s inc reas- i ng . The h i s t o r y o f t h e p i n k salm- o n ' s importance i n t h e e a r l y P a c i f i c

Nor thwest f i s h e r y was g iven i n an overv iew by Cobb (1911). E a r l y salmon canner ies, es tab l i shed i n t h e S e a t t l e area i n t h e 18701s, o c c a s i o n a l l y packed p i n k salmon.

Hoar (1951) summarized t h e p i n k salmon f i s h e r y f o r 1917 t o 1947. The p i n k and t h e chum salmon, r e f e r r e d t o as autumn salmon, were considered l e s s d e s i r a b l e species on t h e market than coho, ch inook, and sockeye salmon be- cause t h e i r f l e s h was s o f t and pa le . A t t h e beg inn ing o f World War I, a s u b s t a n t i a l i nc rease occur red i n t h e f i s h e r y because o f war t ime food re - q u i rements and t h e temporary decrease i n t h e Fraser R i ve r c a t c h o f o t h e r salmon due t o t h e r o c k s l i d e a t H e l l ' s Gate i n B r i t i s h Columbia t h a t b locked t h e Fraser R iver . S ince t h a t t ime, t h e p i n k salmon has been i n c r e a s i n g i n importance. The f i s h a re u s u a l l y t ak - en by purse se ines o r g i l l ne ts , b u t t r o l l f i s h i n g f o r p i n k salmon has been i n c r e a s i n g i n importance s i nce 1950 ( D i Donato 1968). The p r imary reason f o r t h e inc reased t r o l l ca t ch i s an expanded market f o r p i nks i n t h e f resh- and f r o z e n - f i s h markets, i n which f i s h caught by t r o l l i n g a re p re - f e r r e d over those taken i n ne t s (Di Donato 1968). From 1935 t o 1963, t h e average commercial ca t ch o f p i n k salm- on i n Washington S t a t e g e n e r a l l y i n - creased (D i Donato 1968). Washington commercial p i n k salmon land ings i n number o f f i s h by gear t ype (Table 1) i n d i c a t e t h e predominance o f t h e s t r o n g odd-year runs i n Washington S ta te .

D i Donato (1968) i n d i c a t e d t h a t t h e Puget Sound r e g i o n was t h e southern geographic l i m i t o f streams suppo r t i ng major p i n k salmon runs i n t h e eas te rn No r th P a c i f i c . S i g n i f i c a n t p i n k salmon runs a re p resen t o n l y i n odd- numbered years i n Puget Sound r i v e r s and i n t h e Fraser R i ve r , which i s t h e major p i n k salmon produc ing r i v e r i n southern B r i t i s h Col umbia (Di Donato 1968). I n c o n t r a s t , t h e more no r t h - e r l y streams o f B r i t i s h Columbia and Alaska have s t r o n g annual runs o f p i n k

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Table 1. Commercial Landings o f number o f p i n k salmon i n Washington S t a t e by gear t y p e (da ta f rom Washington S ta te Department o f F i s h e r i e s 1983).

G i l l Purse Reef Other T o t a l Year T r o l l n e t se ine n e t gears f i s h

salmon i n bo th odd- and even-numbered years (Manzer e t a l . 1965; Donne l l y 1983), w h i l e B r i s t o l Bay, Alaska, has e s s e n t i a l l y j u s t even-numbered yea r runs ( R . Donne l l y , 1986, F ish . Res. I n s t . , S e a t t l e ; pers. comm.). The c y c l i c a l na tu re o f t h e dominant odd- yea r p i n k salmon runs i n Washington S ta te can be observed i n t h e commer- c i a l catches f o r those years (F i gu re 5).

Not u n t i l t he l a t e 1950 's was i t r e a l i z e d t h a t p i n k salmon would take a r t i f i c i a l b a i t . Since then, t h e i r

p o p u l a r i t y as s p o r t f i s h has i n - creased. From 1957 t o 1961, t h e es t imated annual s p o r t ca t ch i n B r i t i s h Columbia v a r i e d f rom 1,000 t o 37,000 f i s h ( S c o t t and Crossman 1973). The s p o r t f i s h e r y f o r p i n k salmon i n Puget Sound, Washington, has been ex t remely p r o d u c t i v e i n r ecen t years , perhaps s t i m u l a t e d by ext remely l i b e r a l ca t ch l i m i t s s e t by t h e Washington Department o f F i she r i es .

P ink salmon f i s h e r i e s management i s made complex by user-group a l l o c a t i o n s

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SOCKEYE CHINOOK CHUM PINK COHO

1981 - TOTAL POUNDS 47,057,650

1982 - TOTAL POUNDS 49,512,433

F igu re 5. C y c l i c a l na tu re o f t h e commercial p i n k salmon ca t ch r e l a t i v e t o o t h e r salmon i n Washington S ta te i s shown f o r odd- and even-numbered years t h a t y i e l d e d 42% and 0% p i n k salmon, r e s p e c t i v e l y (mod i f i ed f rom Washington S t a t e Department o f F i s h e r i e s 1983).

and mixed-stock f i s h e r i e s . Optimum y i e l d i s t h e d e s i r e d goa l . Successful management r e q u i r e s e f f e c t i v e da ta systems, advance p l ann ing , we1 1 - e s t a b l i s h e d spawning escapement ob jec- t i v e s , dependable p r e d i c t i o n s of p o p u l a t i o n s i z e , and r e c o g n i t i o n of p r a c t i c a l d i f f e r e n c e s between rec rea- t i o n a l and commercial f i s h e r i e s (Wr igh t 1981).

Ocean f i s h e r i e s a r e managed by a ca t ch quota system, w h i l e f i s h e r i e s i n t e r m i n a l areas a r e managed by sub- t r a c t i n g escapement goa ls from presea- son r u n f o recas t s and inseason r u n updates. A f t e r s u b t r a c t i o n o f t h e ocean catches and escapement goa ls , t h e es t imated number remain ing g i ves t h e t o t a l a l l owab le harves t , which i s a1 l oca ted among user groups. The Washington Department of F i s h e r i e s and t h e va r i ous t r i b a l e n t i t i e s have

worked much more c l o s e l y toward a u n i f i e d salmon management p l a n due t o t h e B o l d t Dec i s i on i n 1974 (C la r k 1985).

ECOLOGICAL ROLE

When j u v e n i l e p i n k salmon e n t e r t h e e s t u a r i n e environment, t hey feed near t h e sur face , p r i m a r i l y d u r i n g d a y l i g h t hours (Godin 1981a). Stomach analyses o f p i n k salmon i n B r i t i s h Columbia showed t h a t maximum mean p rey biomass occur red near o r a t dusk (Godin 1981a). Apparen t l y , once a f i s h ' s stomach i s f u l l , spontaneous f eed ing resumes a f t e r o n l y 15% o f t h e stomach con ten ts have been evacuated (Godin 1981b). J u v e n i l e p i n k salmon q u i c k l y adapt t o feed ing on p e l a g i c copepods and o t h e r ep iben th i c and p l a n k t o n i c organisms (Gerke and

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Kaczynski 1972; B a i l e y e t a l . 1975; Cooney e t a l . 1981). Food o f j uve- n i l e s i n p r o t e c t e d wate rs such as Chatham Sound, Alaska, and Puget Sound, Washington, i n c l udes h a r p a c t i - c o i d copepods, copepod n a u p l i i , i n v e r - t e b r a t e eggs, t u n i ca tes , and ba rnac l e l a r v a e (Manzer 1969; Gerke and Kaczynski 1972; Kaczynski e t a l . 1973; Simenstad e t a l . 1980; Godin 1981b). P i nk salmon co -occu r r i ng w i t h j u v e n i l e chum salmon o f approx imate ly t h e same s i z e i n Hood Canal had an ove r l ap i n d i e t a r y i tems t h a t approached 75% (Simenstad e t a l . 1980). The estu- a r i n e res idence t i m e o f j u v e n i l e p i n k salmon v a r i e s f rom 4 t o 18 weeks (Simenstad e t a l . 1982). A l l e n and Aron (1958) found t h e p r e f e r r e d food o f p i n k salmon i n t h e ocean t o be amphipods, supplemented by f i s h , euphaus i ids , squ id , and c rus tacean l a r vae . However, d i f f e r e n c e s were no ted i n t h e d i e t s o f f i s h c l ose t o shore versus those o f f i s h f a r f rom shore: f o r f i s h f rom i nsho re wa te rs , amphipods were t h e most impo r t an t and c rus tacean l a r v a e ranked second i n impor tance; f o r f i s h f rom o f f s h o r e wa te rs , copepods and euphaus i ids were t h e dominant food i tems. P i nk salmon appeared t o s e l e c t 1 a rge r food i tems as t hey moved f u r t h e r o f f s h o r e ( A l l e n and Aron 1958). I t has been hypothes- i z e d t h a t app rec i ab l e d i f f e r e n c e s i n average s i z e of i n d i v i d u a l p i n k salmon i n d i f f e r e n t years a re p robab l y due, i n p a r t , t o d i f f e r e n c e s i n f eed ing c o n d i t i o n s (Davidson and Vaughan 1941; Neave 1953).

Compet i t i on and p r e d a t i o n can have s i g n i f i c a n t e f f e c t s on t h e p i n k salmon popu la t i ons . Young p i n k salmon f ry a re preyed upon by a v a r i e t y o f stream f i s h e s , i n c l u d i n g Do1 l y Varden (x- v e l i n u s malma), c u t t h r o a t t r o u t (Salmo c l a r k i ) , ra inbow t r o u t (Salmo g a i r d - n e r i ) , young coho salmon (Oncorhynchus k i s u t c h ) , n o r t h e r n squawf ish (Ptycho- che i 1 us oregonensi s) , and s c u l p i ns (Co t t us spp. ) (Hunter 1959; H a r t 1973; S c o t t and Crossman 1973; Hargreaves and LeBrasseur 1985). K i n g f i s h e r s , mergansers, o t h e r predaceous b i r d s ,

and mammals a1 so p robab l y account f o r some amount o f p r e d a t i o n ( H a r t 1973; S c o t t and Crossman 1973; Simenstad e t a l . 1982). I n f e s t a t i o n s o f leeches on eggs and f r y can a l s o s i g n i f i c a n t l y i n c rease mo r t a l i t y (Earp and Schwab 1954). P reda t i on by and compe t i t i on w i t h j u v e n i l e coho salmon was con- s i d e r e d t h e main c o n t r i b u t o r t o e a r l y sea m o r t a l i t y i n p i n k salmon acco rd i ng t o Parker (1971). Year1 i n g Coho a p p a r e n t l y p rey s e l e c t i v e l y on young p i n k salmon, even i n t h e presence o f chum salmon t h a t a r e b o t h s i g n i f i - c a n t l y sma l l e r and more abundant t han t h e p i n k salmon (Hargreaves and LeBrasseur 1985). E a r l y sea m o r t a l i t y can a l s o be assoc i a t ed w i t h p r e d a t i o n by c e r t a i n spec ies such as h e r r i n g ( T h o r s t e i nson 1962). A d u l t s a t sea a r e preyed upon by humans, mar ine mammals, and, t o a l e s s e r e x t e n t , l a r g e f i s h e s . Upon r e t u r n i n g t o t h e r i v e r s , p i n k salmon a re p reyed on by bears and humans.

ENVIRONMENTAL REQUIREMENTS

Water Temperature

Accord ing t o Rei se r and B j o r n n (1979), t h e optimum wate r temperatures f o r p i n k salmon spawning range f rom 7.2 t o 12.8 O C . P i nk salmon eggs t o l - e r a t e l o n g p e r i o d s o f low temperature, p r o v i d e d t h e i n i t i a l temperature was above 6.0 O C when embryogenesis began and i n i t i a l development o f t h e embryo progressed t o a s tage t h a t was t o l e r - a n t o f c o l d e r wa te r (Combs and Burrows 1957; Combs 1965; Re ise r and B jo rnn 1979). I n c u b a t i o n temperatures f o r success fu l development range f rom 4.4 t o 13.3 O C (Re ise r and B j o r n n 1979). Ext remely c o l d wa te r and low a i r temperatures cause m o r t a l i t y among i n c u b a t i n g eggs and f r y when i c e fo rmat ion reduces wate r in te rchange (Neave 1953; McNeil 1966; Re ise r and B j o r n n 1979).

A d u l t p i n k salmon a r e co ld -wate r f i s h w i t h a p r e f e r r e d temperature range o f 5.6 t o 14.6 O C , an op t ima l

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temperature o f 10 .1 O C , and an upper 1 e t h a l temperature o f 25.8 O C (Rei se r and B jo rnn 1979). The r e l a t i v e l y h i gh stream temperatures (17 O C ) and low d i s s o l v e d oxygen l e v e l s assoc ia ted w i t h d rought c o n d i t i o n s have apparent- l y k i l l e d many mature a d u l t p i n k salm- on i n Alaskan streams (Krueger 1981).

S a l i n i t y

J u v e n i l e p i n k salmon encounter a wide range o f s a l i n i t i e s i n t h e i r m ig ra t i ons . S a l i n i t y g rad ien t s a r e thought t o p l a y a p a r t i n salmon m i - g r a t i o n s (Baggerman 1960; McInerney 1964). Noerenburg (1963) es t imated t h a t over 50% o f t h e p i n k salmon spawning a c t i v i t y i n P r i nce W i l l iam Sound, Alaska, occurs i n i n t e r t i d a l areas. B a i l e y (1969) showed t h a t de- ve l op ing p i n k salmon eggs and a l e v i n s can be adverse ly a f f e c t e d when exposed t o some h i g h i n t e r t i d a l s a l i n i t i e s .

D isso lved Oxygen

D isso lved oxygen (DO) i s supp l i ed t o deve lop ing eggs and a l e v i n s w i t h i n t h e redd by i n t r a g r a v e l f l ow. Dis- so lved oxygen l e v e l w i t h i n t h e redd i s i n f l u e n c e d by t h e d i s s o l v e d oxygen i n t h e stream, t h e r a t e o f i n t r a g r a v e l water f l ow , and t h e b i o l o g i c a l demand f o r oxygen i n t h e immediate area. For i n c u b a t i o n o f salmonid eggs, concen- t r a t i o n s a t o r near s a t u r a t i o n w i t h temporary r educ t i ons t o l e v e l s no low- e r t han 5 .0 mg/l a re op t ima l (Reiser and B jo rnn 1979). However, DO l e v e l s exceeding 6.0 mg/l a re r e q u i r e d f o r successfu l development o f p i n k salmon

, eggs and a l e v i n s ( B a i l e y e t a l . 1980). Oxygen consumption g e n e r a l l y increases as embryo development progresses, and DO g e n e r a l l y decreases i n an area sup- p o r t i n g l a r g e numbers o f eggs o r a1 e v i ns (Bai l e y e t a1 . 1980). Low DO concen t ra t i ons may r e s u l t i n a v a r i e t y o f f r y abno rma l i t i e s ( S i l v e r e t a l . 1963; Shumway e t a l . 1964; Re iser and B jo rnn 1979). Dur ing t h e l a t t e r p a r t o f i n cuba t i on , reduced DO can cause premature ha t ch ing (A lde rd i ce e t a l . 1958; B a i l e y e t a l . 1980). Growth

r a t e , food consumption, and e f f i c i e n c y o f food u t i l i z a t i o n by j u v e n i l e salmon a l l dec l i ned a t lowered DO (Reiser and B jo rnn 1979). Reduced DO concentra- t i o n s can s i g n i f i c a n t l y hamper t h e swimming performance o f m i g r a t i n g a d u l t salmonids (Jones 1971; Re iser and B jo rnn 1979). I n a d d i t i o n , mor- t a l i t i e s may occur i n a d u l t s when low DO i s combined s y n e r g i s t i c a l l y w i t h h i g h temperatures (Krueger 1981).

Subs t ra te

The d iameter o f g rave l t h a t com- poses t h e subs t ra te f o r spawning p i n k salmon ranges from 1.3 t o 10.2 cm (Lucas 1959; Col l i n q s 1974). P i nk salmon spawn over a v a r i e t y of sub- s t r a t e m a t e r i a l s. The s i z e , shape, dens i t y , and embeddedness o f t h e mate- r i a l , c u r r e n t v e l o c i t y , water depth, and d e n s i t i e s o f f i s h can i n f l u e n c e s u b s t r a t e se lec t i on . Successful f ry emergence i s phys i ca l l y h indered by excess ive amounts o f sand and s i l t i n t h e g r a v e l , which a l s o l i m i t t h e p e r c o l a t i o n o f water w i t h i t s DO con ten t (Reiser and B jo rnn 1979). Hourston and MacKinnon (1957) suggest- ed t h a t ve r y smal l g rave l s made i d e a l p i n k salmon spawning areas. No d i f - ferences i n s i z e o r t ime o f r e t u r n o f a d u l t s cou ld be t r a c e d t o t h e na tu re o f t h e g rave l environment from which t hey came (Bai 1 ey e t a1 . 1976).

Eggs and deve lop ing a l e v i n s a re i n f 1 uenced by t h e subs t ra te . Produc- t i v e p i n k salmon streams g e n e r a l l y con ta ined l e s s t han 5.0% by volume o f f i n e sediments ( t 0 . 8 mm), whereas l e s s p roduc t i ve streams had 15.0% o r more f i n e s i n t h e subs t ra te (Krueger 1981).

Water Depth

Water depth se lec ted by p i n k salmon i s determined by c u r r e n t ve loc - i t y and subs t ra te type. P re fe r red water depths f o r spawning p i n k salmon i n se l ec ted Washington S t a t e streams ranged from 0.2 t o 0.5 m ( C o l l i n g s 1974). Hourston and MacKi nnon (1957) s t u d i e d water depth preferences i n a

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spawning channel and found t h a t t h e f i r s t f i s h t h a t en te red t h e channel chose s i t e s w i t h mean depths o f 0 .4 m and subsequent f i s h occupied p rogres - s i v e l y sha l l owe r mean depths. It ap- pears t h a t i n na tu re these sha l lower depths a r e used even though t hey a re l e s s t han op t ima l s i t u a t i o n s (Krueger 1981; Wi l son e t a l . 1981).

Water V e l o c i t y

A1 though severa l c u r r e n t v e l o c i - t i e s a r e s u i t a b l e f o r p i n k salmon spawning, t h e p r e f e r r e d v e l o c i t i e s range f rom 2 1 t o 1 0 1 cm/s ( C o l l i ngs 1974). P i nk salmon i n t h e T e r r o r R i ve r , A laska, spawned i n areas w i t h c u r r e n t v e l o c i t i e s r ang ing f rom 0.19 t o 0.66 m/s, b u t t h e p r e f e r r e d spawn- i n g v e l o c i t i e s ranged f rom 0.35 t o 0.47 m/s (Wi lson e t a l . 1981). Flow regimes can a f f e c t deve lop ing p i n k salmon eggs and a l e v i n s th rough e i t h e r mechanical damage, temperature changes, o r reduced i n t r a g r a v e l d i s - so l ved oxygen l e v e l s (McNeil 1966; Re ise r and B j o r n n 1979; Krueger 1981).

The h i g h c u r r e n t v e l o c i t i e s asso- c i a t e d w i t h h i g h s t ream d ischarges

sometimes exceed t h e swimming c a p a b i l - i t i e s o f p i n k salmon and p reven t up- stream m ig ra t i on . However, p i n k salmon can n e g o t i a t e c u r r e n t v e l o c i - t i e s up t o about 2 . 1 m/s (Krueger 1981). Upstream m i g r a t i o n s o f a d u l t s can be hampered by e i t h e r t o o l i t t l e o r t oo much stream f l o w (W icke t t 1958; Re ise r and B jo rnn 1979; Krueger 1981).

T u r b i d i t y

Accord ing t o Rei s e r and B j o r n n (1979), m i g r a t i n g salmon a v o i d o r cease t o m ig ra te i n wa te rs w i t h ve r y h i g h s i l t loads (4,000 mg/l) . T u r b i d wa te r absorbs more r a d i a t i o n t han c l e a r wa te r , thus p o s s i b l y r e s u l t i n g i n temperature b a r r i e r s t o upstream m i g r a t i o n . Inc reased t u r b i d i t y a1 so may cause osmoregul a t o r y problems i n young f i s h by i n t e r f e r i n g w i t h g i l l f u n c t i o n . T h i s r e s u l t s i n de layed e n t r y i n t o seawater and inc reased p re - d a t i o n on t h e e s t u a r i n e p o p u l a t i o n (Kehoe 1982). Eggs may be s u f f o c a t e d by t h e inc reased d e p o s i t i o n o f f i n e sediment and a d u l t s may s u f f e r f rom impa i r ed oxygen t r a n s p o r t , dec l i ne i n n u t r i t i o n a l i n t a k e , and p h y s i c a l dam- age t o t h e g i l l f i l a m e n t s by ab ras i on ( R i v i e r and Seguier 1985).

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LITERATURE CITED

A lderd ice , D.W., W. P. Wicke t t , and t h e area. U.S. N a t l . Mar. F ish. J.R. B r e t t . 1958. Some e f f e c t s o f Serv. F ish . B u l l . 73(4):846-861. temporary exposure t o 1 ow d i sso l ved oxygen l e v e l s on P a c i f i c salmon B a i l e y , J.E., J .J . P e l l a , and S.G. eggs. J. F ish. Res. Board Can. Tay lo r . 1976. Produc t ion o f f ry 15(2) : 229-250. and a d u l t s o f t h e 1972 brood o f p i n k

A lexande rsdo t t i r , M. , and 0. A. Math i - sen. 1983. L i f e h i s t o r y o f p i n k salmon (Oncorhynchus gorb;scha)' and i m ~ l i c a t i o n s f o r manaaement. Univ. ~ a ' s h i n ~ t o n , F ish. ~es ' : I n s t . , Rep. No. FRI-UW-8313. 72 pp.

A1 1 en, G. H. , and W. Aron. 1958. Food o f sa lmonid f i s h e s o f t h e western n o r t h P a c i f i c Ocean. U.S. F i s h W i l d l . Serv. Spec. Sc i . Rep. F ish. No. 237. 11 pp.

Anas, R. E. 1959. Three-year-01 d p i n k salmon. J. F ish. Res. Board Can. 16(1) : 91-92.

Baggerman, B. 1960. S a l i n i t y p re - ference, t h y r o i d a c t i v i t y , and t h e seaward m i g r a t i o n o f f o u r species o f P a c i f i c s a l mon (Oncorhynchus). J. F ish. Res. Board Can. 17(2): 295-322.

B a i l e y , J.E. 1969. E f f e c t s o f s a l i - n i t y on i n t e r t i d a l p i n k salmon sur - v i v a l . Alaska Dep. F i s h Game I n f . L e a f l . No. 87:12-15.

B a i l e y , J.E. B.L. Wing, and C.R. Mat t - son. 1975. Zooplankton abundance and f eed ing h a b i t s o f f r y o f p i n k salmon, oncorhynchus gorbuscha, and chum salmon, Oncorhynchus ke ta , i n T r a i t o r s Cove, A1 aska, w i t h spec- u l a t i o n on t h e c a r r y i n g c a p a c i t y o f

salmon, Oncorhynchus gorbuscha, from grave l i ncuba to r s and n a t u r a l spawn- i n g a t Auke Creek, Alaska. U.S. N a t l . Mar. F ish . Serv. F ish. B u l l . 74(4): 961-971.

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C a r l , G . C . , W.A. Clemens, and C.C . L indsey. 1959. The f reshwate r f i s h e s o f B r i t i s h Columbia. B r i t i s h Columbia Prov. Mus. Handb. No. 5. 192 pp.

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Gerke, R. J. , and V. W. Kaczynski . 1972. Food o f j u v e n i l e p i n k and chum salmon i n Puget Sound, Washington. Wash. Dep. F ish . Tech. Rep. No. 10. 27 PP-

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Oncorhynchus gorbuscha Wal baum. J. F i s h B i o l . 19(1): 63-71.

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Hargreaves, N. B. , and R.J. LeBrasseur. Kaczynski , V.W. , R . J. F e l l e r , J. 1985. Species s e l e c t i v e p r e d a t i o n C lay ton , and R.G. Gerke. 1973. on j u v e n i l e p i n k (Oncorhynchus Troph ic a n a l y s i s o f j u v e n i l e p i n k

orbuscha) and chum salmon (0. ke ta ) and chum salmon i n Puget Sound. J. :y coho salmon (0. k i s u t c h r 5. Fish. Res. Board Can. 30(7):1003- J. F ish . Aquat. Sc i . 42(4): 659-668. 1008.

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Kehoe, D.M. 1982. The e f f e c t s o f Grays Harbor Es tuary sediment on t h e osmoregul a t o r y a b i 1 i t y o f coho salm- on smol t s , Oncorhynchus k i sutch. Army Corps o f Engineers Repor t (Se- a t t l e D i s t r i c t ) . 27 pp.

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McDonald, J . 1960. The behaviour o f P a c i f i c salmon f r y d u r i n g t h e i r downstream m i g r a t i o n t o f r eshwa te r and s a l t w a t e r nu r se r y areas. J. F ish. Res. Board Can. 17(5): 655-676.

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P r i t c h a r d , A. L. 1939. A study o f t h e n a t u r a l propagat ion o f t he p i n k salmon, dncorhynchus gorbuscha,' i n B r i t i s h Columbia. Trans. Am. F ish.

P r i t c h a r d , A. L. 1944. Phys ica l char- a c t e r i s t i c s and behavior o f p i n k salmon f r y a t McCl in ton Creek, B.C. J. F ish. Res. Board Can. 6(3):217- 227.

Reiser , D. W. , and T.C. Bjornn. 1979. H a b i t a t requirements o f anadromous salmonids. Pages 1-54 jJ W.R. Meehan, ed. I n f l u e n c e o f f o r e s t and rangeland management on anadromous f i s h h a b i t a t i n t he Western Un i t ed

oxygen concen t ra t i on and water move- ment on t he growth o f s teelhead t r o u t and coho salmon embryos. Trans. Am. F ish. Soc. 93(4): 342-356.

S i l v e r , S.T., C.E. Warren, and P. Doudoroff . 1963. D isso lved oxygen requirements o f developing steelhead t r o u t and chinook salmon embryos a t d i f f e r e n t water v e l o c i t i e s . Trans. Am. Fish. Soc. 92(4): 327-343.

Simenstad, C.A. , W. J. Kinney, S.S. Parker, E.O. Salo, J. R. Co rde l l , and H. Buechner. 1980. Prey commu- n i t y s t r u c t u r e and t r o p h i c ecology o f ou tm ig ra t i ng j u v e n i l e chum and p i n k salmon i n Hood Canal, Washing- ton. Univ. Wash. F ish . Res. I n s t . , Rep. No. FRI-UW-80 26. 113 pp.

S ta tes and Canada. U. S. For. Serv. Gen. Tech. Rep. PNW-96. Por t land , Simenstad, C.A. , K. L. Fresh, and E.O.

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Phys ica l and b i o l o g i c a l e f f e c t s o f salmon: an unappreciated f unc t i on .

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Washington S t a t e Department o f F isher - i e s . 1983. 1982 f i s h e r i e s s t a t i s - t i c a l r e p o r t . Olympia, Wash. 77 pp.

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W icke t t , W.P. 1958. Review o f c e r t a i n env i ronmenta l f a c t o r s a f f e c t i n g t h e p r o d u c t i o n o f p i n k and chum salmon. J. F ish . Res. Board Can. 15(5):1103- 1126.

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Wi lson, W. , W. T r i hey , J. Ba ld r i dge , C. Evans, J. T h i e l e , and D. Trudgen. 1981. An assessment o f environmen- t a l e f f e c t s o f c o n s t r u c t i o n and ope ra t i on o f t h e proposed T e r r o r Lake H y d r o e l e c t r i c F a c i l i t y , Kodiak, Alaska. A r c t i c Env i ronmenta l I n f o r - mat ion and Data Center , Anchorage, Alaska. 429 pp.

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50172 -101

~ o r t h w e s t ) - - p i n k Salmon 1 -

REPORT DOCUMENTATION I PAGE I B i o l o a i c a l r i e ~ o r t 82( 11.88) *

4. Title and Subtitle 1 5. Repor( Date

Species P r o f i l e s : L i f e H i s t o r i e s and Environmental January 1989

?. Authods) 8. Performing Organization Rept. Nc

S c o t t A. Bonar, G i l b e r t B. Pauley, and G. L. Thomas 9. Performing Orsanixation Name and Address

2. I 3. R n p ~ e n t ' s Accession NO,

Reauirements o f Coastal F ishes and I n v e r t e b r a t e s ( P a c i f i c

Washington Cooperat ive F i she ry 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

.S

12. Sponsoring Organization Name and Address

U.S. Dep. o f t h e I n t e r i o r U.S. Army Corps o f Engineers F i s h and W i l d l i f e Serv ice Waterways Experiment S t a t i o n Na t i ona l Wetlands Research Center P. 0. Box 631 Washington, DC 20240 Vicksburg, MS 39180

1'; Contract'c) or Grant'") NO.

(GI

13. Type of Repor( L Period Coverec

-- IS. Supplementary Notes

*U.S. Army Corps o f Engineers Report No. TR EL-82-4

16. Abstract (Limit: 200 word.)

Species p r o f i l e s a re l i t e r a t u r e summaries o f t h e taxonomy, morphology, range, l i f e h i s t o r y , and environmental requirements o f coas ta l aqua t i c species. They a re designed t o a s s i s t i n environmental impact assessment. The p i n k salmon, o f t e n c a l l e d humpback salmon o r humpy, i s e a s i l y i d e n t i f i e d by i t s ex t remely smal l sca les (150 t o 205) on t h e l a t e r a l l i n e . They a re t h e most abundant o f t h e P a c i f i c salmon species and spawn i n No r th American and Asian streams bo rde r i ng t h e P a c i f i c and A r c t i c Oceans. They have a ve ry s imple two-year l i f e c y c l e , which i s so i n v a r i a b l e t h a t f i s h runn ing i n odd- numbered yea rs a re i s o l a t e d f rom f i s h runn ing i n even-numbered years so t h a t no gene f l o w occurs between them. Adu l t s spawn i n t h e f a l l and t h e young f r y emerge i n t h e sp r i ng . The p i n k salmon i s l e s s d e s i r a b l e i n commercial and s p o r t catches than most o t h e r salmon because o f i t s smal l s i z e and i t s s o f t p a l e f l e s h . The Puget Sound r e g i o n o f Washington S t a t e i s t h e southern geographic l i m i t o f streams suppor t i ng major p i n k salmon runs i n t h e eas te rn Nor th P a c i f i c . P ink salmon runs a r e p r e s e n t l y o n l y i n odd- numbered yea rs i n t h i s reg ion . Optimum water temperatures f o r spawning range f rom 7.2 t o 12.8 OC. P roduc t i ve p i n k salmon streams have l e s s than 5.0% by volume o f f i n e sediments (50.8 mm).

17. Document Analysis a. Descrlpton

Es tua r i es Temperature L i f e c y c l e F i s h e r i e s Sediments Growth Salmon Suspended sediments Oxygen c o m p e t i t i o n S a l i n i t y Feeding h a b i t s Animal m i g r a t i o n s b. 1dentlfienlOpen.Ended Terms

Pink salmon L i f e h i s t o r y Predators Humpback salmon Spawning Compet i tors Oncorhynchus gorbuscha Temperature requirements

Sal i n i t y requirements

.a Availability Statement / 19. Secur~ty Class (This Repom 1 21. No. of Pages

Oepertment of Commerce

U n c l a s s i f i e d U n l i m i t e d

U n c l a s s i f i e d

18 -- 22. Price

ee ANSI-239.18) OPTIONAL FORM 272 (4-77 (Fornwrly NTIS35)

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TAKE PRIDE in Amerzcd

DEPARTMENT OF THE IWTERIOR U.S. FISH AND Wl lDL lA SERVICE

As the Nation's principal conservation agency. the Department of the Interior has respon- sibility for most of our .nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resourcm, protecting our fish and wildlife, preserving thsenvironmental and cultural values of our national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department as- sesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for paople who live in island territories under U.S. administration.