Species Profiles: Life Histories and EnvironmentalRequirements of Coastal Fishes and Invertebrates

(Pacific Southwest): Northern anchovy

Item Type monograph

Authors Kucas, Stephen T.

Publisher United States Fish and Wildlife Service

Download date 09/09/2021 22:41:31

Link to Item http://hdl.handle.net/1834/21626

REFERENCECOPY Do Not Remove from the Library

U. S. Fish and Wildlife Service - - - 1 \ A G' r-

Biological Apr i l 1 9 8 6 Report 8 2 ( 1 1-60) "=Gvard TR EL-82-4

Lofoyette, Louisiana 70506

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

NORTHERN ANCHOVY

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station

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

B i o l o g i c a l Repor t 82(11.50) TR EL-82-4 A p r i l 1986

Spec ies P r o f i l e s : L i f e H i s t o r i e s and Env i ronmen ta l Requi rements o f C o a s t a 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 Southwest )

NORTHERN ANCHOVY

Stephen T. Kucas, J r . Ca l i f o r n i a C o o p e r a t i v e F i s h e r y Research U n i t

Humboldt S t a t e U n i v e r s i t y A rca ta , CA 95521

P r o j e c t O f f i c e r John Parsons

N a t i o n a l C o a s t a l Ecosystems Team U.S. F i s h and W i l d l i f e S e r v i c e

1010 Gause Bou leva rd S l i d e l l , LA 70458

Per formed f o r

C o a s t a l Eco logy Group Waterways Exper imen t S t a t i o n U .S. Army Corps o f E n g i n e e r s

V icksburg , MS 39180

and

N a t i o n a l Coas ta l Ecosystems Team D i v i s i o n o f B i o l o g i c a l S e r v i c e s

Research and Development F i s h and W i l d l i f e S e r v i c e

U.S. Department o f t h e I n t e r i o r Washington, DC 20240

T h i s se r i es should be referenced as f o l l ows :

U.S. F i sh and W i l d l i f e Service. 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 environmental requirements o f coas ta l f i s h e s and inver tebra tes . U.S. F i sh Wi ld l . Serv. B io l . Rep. 82(11). U.S. Army Corps o f Engineers, TR EL-82-4.

Th i s p r o f i l e should be c i t e d as f o l l ows :

Kucas, S.T., Jr . 1986. Species p r o f i l e s : 1 i f e h i s t o r i e s and environmental r equ i 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 Southwest)--northern anchovy. U.S. F i sh Wi ld l . Serv. B io l . Rep. 82(11.50). U.S. Army Corps o f Engineers, TR EL-82-4. 11 pp.

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f a s e r i e s on c o a s t a l a q u a t i c o rgan isms, p r i n c i p a l l y f i s h , o f s p o r t , commerc ia l , o r e c o l o g i c a l i m p o r t a n c e . The p r o f i l e s a r e d e s i g n e d t o p r o v i d e c o a s t a l p l a n n e r s , managers, e n g i n e e r s , and b i o l o g i s t s w i t h a b r i e f s k e t c h o f t h e 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 e n v i r o n m e n t a l r e q u i r e m e n t s o f t h e s p e c i e s and t o d e s c r i b e how p o p u l a t i o n s may be expec ted t o r e a c t t o e n v i r o n m e n t a l changes caused b y c o a s t a l deve lopment . Each p r o f i l e has s e c t i o n s on taxonomy, l i f e h i s t o r y , e c o l o g i c a l r o l e , e n v i r o n m e n t a l r e q u i r e m e n t s , and economic impor tance , if a p p l i c a b l e . A t h r e e - r i n g b i n d e r i s used f o r t h i s s e r i e s so t h a t new p r o f i l e s can be added as t h e y a r e p repa red . T h i s p r o j e c t i s j o i n t l y p lanned and f i n a n c e d b y t h e U.S . Army Corps o f E n g i n e e r s and t h e U.S. F i s h and W i l d l i f e S e r v i c e .

S u g g e s t i o n s o r q u e s t i o n s r e g a r d i n g t h i s r e p o r t s h o u l d be d i r e c t e d t o :

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l C o a s t a l Ecosystems Team U.S. F i s h and W i l d l i f e S e r v i c e NASA-S 1 i d e l l Computer Complex 1010 Gause B o u l e v a r d S l i d e l l , LA 70458

U.S. Army Eng inee r Waterways Expe r imen t S t a t i o n A t t e n t i o n : WESER-C P o s t O f f i c e Box 631 V i cksbu rg , MS 39180

CONVERSION TABLE

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

m i l l i m e t e r s (mn) cen t imete rs (an) meters (m) k i l omete rs ( h )

2 square meters (m ) 10.76 square k i l o m e t e r s ( h 2 ) 0.3861 hec ta res (ha) 2.471

l i t e r s ( 1 ) c u b i c mete rs (m3) cub ic meters

To O b t a i n

inches inches f e e t mi 1 es

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

gal 1 ons c u b i c f e e t a c r e - f e e t

m i l 1 igrams (mg) 0.00003527 ounces grams ( g ) 0.03527 ounces k i log rams ( k ) 2.205 pounds m e t r i c tons q t ) 2205.0 pounds m e t r i c tons 1.102 s h o r t tons k i 1 ocal o r i e s ( kca l ) 3.968 B r i t i s h thermal u n i t s

Cel s i u s degrees 1.8("C) + 32 Fahrenhei t degrees

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

inches 25.40 inches 2.54 f e e t ( f t ) 0.3048 f a thoms 1.829 m i i e s ( m i ) 1.609 nacl;ical m i l e s ( m i ) 1.852

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

ga l 1 ons ( g a l ) cub ic f e e t ( f t 3 ) a c r e - f e e t

ounces (02) 28.35 pounds ( l b ) 0.4536 s h o r t tons ( t o n ) 0.9072 B r i t i s h thermal u n i t s ( B t u ) 0.2520

m i l 1 i m e t e r s cen t imete rs meters meters k i l o m e t e r s k i 1 ometers

square meters hectares square k i l ometers

1 i t e r s c u b i c meters cub ic meters

grams k i 1 og rams m e t r i c tons k i 1 ocal o r i e s

Fahrenhei t degrees 0.5556("F - 32) C e l s i u s degrees

CONTENTS

PREFACE ................................................................... iii CONVERSION T A B L E .......................................................... i v ACKNOWLEDGMENTS ........................................................... v i

NOMENCLATURE/TAXONOMY/RANGE ........................................ 1 MORPHOLOGY/IDENTIFICATION A I D S ............................................ 1 REASON FOR I N C L U S I O N :IN S E R I E S ........................................ 3 L I F E H I S T O R Y .............................................................. 3

S p a w n i n g ................................................................ 3 L a r v a l S t a g e ............................................................ 3 J u v e n i l e s ............................................................... 4 M a t u r i t y a n d L i f e S p a n .................................................. 4

GROWTH C H A R A C T E R I S T I C S .................................................... 4 COMMERCIAL AND B A I T F I S H E R I E S ........................................ 4 ECOLOGICAL ROLE ........................................................... 6 ENVIRONMENTAL REQUIREMENTS ................................................ 6

T e m p e r a t u r e ............................................................. 6 ................................................................... D e p t h 7

O t h e r E n v i r o n m e n t a l F a c t o r s ............................................. 7

L I T E R A T U R E C I T E D .......................................................... 9

ACKNOWLEDGMENTS

Thanks t o Rick K1 i n g b e i l , Joseph Lesh, and J e r r y S p r a t t (Cal i f o r n i a Department of F i s h and Game) f o r rev iewing t h e manuscr ipt and t o Thomas Hass le r (Gal i f o r n i a Cooperat ive F ishery Research U n i t ) f o r a s s i s t i n g i n i t s p repara t ion .

F i g u r e 1. N o r t h e r n anchovy.

S c i e n t i f i c name . . . . . . E n p r a u l i s mordax ( G i r a r d

P r e f e r r e d common name . . . . N o r t h e r n anchovy ( F i g u r e 1 )

O the r c o n o n names . . . . . Pinhead ( a p p l i e d t o s m a l l f i s h )

C lass . . . . . . . . . . O s t e i c h t h y e s Order . . . . . . . . . . C l u p e i f o r m e s F a m i l y . . . . . . . . . . E n g r a u l i d a e

Geograph ic range: Queen C h a r l o t t e I s l a n d s , B r i t i s h Columbia, sou th t o Cape San Lucas, B a j a C a l i f o r n i a ; c e n t e r o f abundance ex tends f r o m Magdalena Bay, Mexico, t o San F ranc i sco , Cal i f o r n i a (Ah1 s t r o m 1966). Three p o p u l a t i o n s a r e recogn ized : one f r o m B r i t i s h Columbia t o n o r t h e r n C a l i f o r n i a , a second o f f s o u t h e r n Cal i f o r n i a and t h e n o r t h e r n B a j a C a l i f o r n i a p e n i n s u l a i n Mexico, and a t h i r d

o f f c e n t r a l and s o u t h e r n B a j a C a l i f o r n i a ( F i g u r e 2). A1 though t h e ranges o f t h e s e p o p u l a t i o n s over1 ap somewhat, each i s g e n e t i c a l l y d i s t i n c t (Vrooman and Smi th 1971).

F i n r a y s : d o r s a l 14-19, ana l 19-26, p e c t o r a l 13-20; m i d l a t e r a l s c a l e s 41-50; g i 11 r a k e r s 28-41 and 37-45 on l o w e r a r c h (number i n c r e a s e s w i t h s i z e ) ; g i 11 r a k e r s much l o n g e r t h a n eye; p e l v i c f i n s abdomina l ; v e r t e b r a e 43-47. Body i s l o n g and s l i g h t l y compressed. Head i s a n t e r i o r l y compressed and n e a r l y t w i c e as l o n g as i t i s deep; s n o u t p r o t r u d e s and i s p o i n t e d w i t h l a r g e eye near t h e

- -

' ~ a r g e l y e x t r a c t e d f rom Jo rdan and Evermann (1908) and M i l l e r and Lea (1972).

WASHINGTON SUBPOPULATIO

SUBPOPULATI

SUBWULATIO

F igu re 2. D i s t r i b u t i o n o f no r the rn anchovy ( f rom P a c i f i c F ishery Management Counci 1 1978).

t i p ; g i l l open ings ex tend under jaw i n t o t h r o a t ; mouth i s s u b t e r m i n a l w i t h sma l l t e e t h ; maxi 1 l a r y ex tends beyond r o o t of mandib le ; o p e r c l e i s p l a c e d o b l i q u e l y and deeper t h a n i t i s l ong . C o l o r i n l i f e : b l u i s h above, s i l v e r on s i d e and below; i t i s n o t t r a n s l u c e n t and has no s i l v e r y l a t e r a l band.

REASON FOR INCLUSION IN SERIES

The n o r t h e r n anchovy i s abundant i n t h e C a l i f o r n i a C u r r e n t and i s e c o l o g i c a l l y and e c o n o m i c a l l y i m p o r t a n t i n t h e c o a s t a l w a t e r s of s o u t h e r n C a l i f o r n i a ( S o u l e and Ogur i 1972-1976; Mais 1974) . Though i t s u p p o r t s a sometimes t h r i v i n g i n d u s t r i a l f i s h e r y and a l u c r a t i v e l i v e - b a i t f i s h e r y , b o t h y i e l d much l e s s p r o f i t t h a n anchovy f i s h e r i e s i n o t h e r p a r t s o f t h e w o r l d ( B a x t e r 1967). A sha rp i n c r e a s e i n t h e biomass o f t h e n o r t h e r n anchovy o f f C a l i f o r n i a i n r e c e n t decades and a l a t e r d e c l i n e i n l a n d i n g s i n t h e l a t e 1 9 7 0 ' s have l e d t o an i n t e n s i v e a t t e m p t b y f i s h e r y agenc ies i n C a l i f o r n i a and Mex ico t o u n d e r s t a n d t h e b i o l o g y and p o p u l a t i o n dynamics o f t h i s s p e c i e s .

LIFE HISTORY

Spawning

B i o l o g i c a l i n f o r m a t i o n abou t t h e n o r t h e r n anchovy was s u n a r i z e d b y B a x t e r (1967) . He r e p o r t e d t h a t anchovy spawning, a1 though r e c o r d e d f rom B r i t i s h Columbia t o a p o i n t be low Magdalena Bay, B a j a C a l i f o r n i a , i s h e a v i e s t between P o i n t Concept ion, Cal i f o r n i a , and P o i n t San Juan ico, B a j a C a l i f o r n i a . I n w a t e r s n o r t h o f P o i n t Concept ion, spawning success has v a r i e d w i d e l y . Two ma jo r spawning a reas a r e s o u t h o f P o i n t Concep t ion . One i s o f f s o u t h e r n C a l i f o r n i a and n o r t h e r n B a j a C a l i f o r n i a , and t h e o t h e r i s o f f c e n t r a l and s o u t h e r n B a j a C a l i f o r n i a ( A h l s t r o m 1956) .

I n t h e w i n t e r , anchov ies u s u a l l y move t o deeper wa te r o f f sho re , and i n t h e s p r i n g t h e y r e t u r n t o i n s h o r e sha l low w a t e r s . Spawning i s m o s t l y w i t h i n 6 0 mi of t h e c o a s t , a1 though i t has been reco rded up t o 300 m i o f f - shore. Anchovies s t a y n e a r t h e bo t tom i n t h e day t ime and come t o t h e s u r f a c e a t n i g h t . They spawn m o s t l y a t dep ths l e s s t h a n 10 m, a t w a t e r t empera tu res o f 12 t o 15 " C ( A h l s t r o m 1959).

A1 though anchov ies spawn t h r o u g h o u t t h e y e a r , p a r t i c u l a r l y i n t h e s o u t h e r n p a r t o f t h e i r range ( B a x t e r 19671, most spawn i n w i n t e r and s p r i n g ( S t a u f f e r and Parke r 1980) ; however, spawning n o r t h o f P o i n t Concept ion peaks d u r i n g t h e p e r i o d f r o m mid-June t o mid-August ( H u n t e r and Macewicz 1980 1.

Anchovies spawn s e v e r a l t i m e s a yea r ; i n d i v i d u a l f ema les may l a y as many as 20,000 t o 30,000 eggs a y e a r ( H a r t 1973) . D u r i n g t h e peak spawning season, t h e f i s h appear t o spawn about once a week ( H u n t e r and Goldberg 1980) . The eggs a r e p e l a g i c and f l o a t p a s s i v e l y i n t h e upper l a y e r s o f t h e ocean. Eggs have been c o l l e c t e d ( a t 10 m d e p t h ) a t w a t e r t e m p e r a t u r e s o f 9.9 t o 23.3 "C ; more t h a n 90% were t a k e n a t 13.0 t o 17.5 "C.

The eggs of anchov ies a r e o v o i d (1.23 t o 1.55 m a l o n g t h e ma jo r a x i s and 0.65 t o 0.82 mn a l o n g t h e m ino r a x i s ) , c l e a r , and t r a n s 1 u c e n t . They h a t c h i n 2 t o 4 days, depending on wa te r t e m p e r a t u r e ( B o l i n 1936) .

L a r v a l Staqe

Newly ha tched l a r v a e a r e 2.5 t o 3 mm l o n g and weigh 0.0246 + 0.0014 mg d r y we igh t , of wh ich 53% i s y o l k (Hun te r 1977) . The l a r g e and e l o n - ga ted yo1 k sac i s absorbed i n about 36 h. The l a r v a e a r e e l o n g a t e d , t r a n s - p a r e n t , and t h r e a d 1 i ke; 01 f a c t o r y and l a t e r a l l i n e o rgans a r e w e l l deve loped. A f t e r h a t c h i n g , l a r v a e a r e i n a c t i v e and f l o a t m o t i o n l e s s i n t h e wa te r e x c e p t d u r i n g s h o r t b u r s t s of

i n t e n s e swimming a t about 1-mi n i n t e r v a l s . The mouth i s t e r m i n a l i n t h e e a r l y s tages. La rvae about 10 mm l o n g come t o t h e s u r f a c e a t n i g h t t o g u l p a i r t o i n f l a t e t h e i r swim b ladder and t h u s conserve energy t h a t wou ld o t h e r w i s e be r e q u i r e d t o m a i n t a i n t h e i r p o s i t i o n i n t h e water column (Hun te r and Sanchez 1976 1. R ichardson (1981 1 hypo thes i zed t h a t t h e s e n i g h t l y v e r t i c a l m i g r a t i o n s cause southward and o f f s h o r e t r a n s p o r t o f t h e l a r v a e o f f C a l i f o r n i a . L a b o r a t o r y measurements i n d i c a t e d t h a t s c h o o l i n g b e g i n s i n l a r v a l anchov ies when t h e y a r e 11 t o 12 mn s tandard l e n g t h (SL ) . The onse t o f s c h o o l i n g i s c o n c u r r e n t w i t h an i n c r e a s e i n p a t c h i n e s s of l a r v a e i n t h e sea (Hun te r and Coyne 1982 1. Rap id s t r u c t u r a l and b e h a v i o r a l changes occu r when t h e f i s h a r e 12-15 mm l o n g . The l e n s r e t r a c t o r musc le becomes f u n c t i o n a l and t h e number of r o d s i n t h e r e t i n a i n c r e a s e s ( 0 ' Connel l 1981) . The young about 25 mn l o n g resemble t h e a d u l t s .

I n one s tudy, su rvey c r u i s e s were made f r o m t h e C a l i f o r n i a - O r e g o n b o r d e r t o t h e s o u t h e r n t i p o f Ba ja C a l i f o r n i a f r o m 1949 t o 1964 t o d e t e r m i n e t h e d i s t r i b u t i o n and abundance of anchovy eggs and l a r v a e ( A h l s t r o m I967 1. C r u i s e s were made about month:_v f r o m 1949 t o 1960, and q u a r t e r l y f r o m i 9 6 1 t o 1964. O f a l l l a r v a e c a p t u r ~ d d u r i n g su rveys f r o m 1951 t o 1960, 96% were t a k e n between P o i n t Concept i o n , Cal i f o r n i a , and Magdalena Bay, Ba ja C a l i f o r n i a . Most were c o l l e c t e d f r o m January t o May and t h e fewest f r o m August t o October . Water t empera tu res ( a t 10 m d e p t h ) were 1' t o 18 "c .

Juven i 1 es

L i t t l e i s known about t h e movement and h a b i t a t p r e f e r e n c e o f j u v e n i l e anchov ies . Tag r e t u r n s have shown an i n t e r c h a n g e o f f i s h between t h e c e n t r a l Cal i f o r n i a , s o u t h e r n Cal i f o r n i a, and n o r t h e r n B a j a C a l i f o r n i a f i s h i n g grounds, o r between t h e c e n t r a l and s o u t h e r n subpopu-

l a t i o n s (Chavez e t a l . 1977). Most i n v e s t i g a t o r s m a i n t a i n t h a t b o t h j u v e n i l e s and a d u l t s move o f f s h o r e i n w i n t e r and r e t u r n toward shore i n s p r i n g . Changing w ind p a t t e r n s i n f a l l , f r o m n o r t h e r l y t o s o u t h e r l y , c o u l d cause a s h i f t i n s u r f a c e c u r r e n t s f r o m southward t o no r thward , a dampening o f u p w e l l i n g , and an onshore d r i f t o f s u r f a c e wa te rs (Wya t t e t a l . 1972) . These f a c t o r s may c o n t r i b u t e t o a n o r t h e r l y onshore movement o f j u v e n i l e s a l o n g t h e c o a s t o f C a l i f o r n i a .

M a t u r i t y and L i f e Span

A t l e a s t h a l f o f a l l female anchov ies r e a c h sexua l m a t u r i t y when about 96 mm l o n g a t t h e end of t h e i r f i r s t yea r o f l i f e (Hun te r and Macewicz 1980); a l l anchov ies a r e mature i n t h e i r second y e a r o f l i f e (age group I ) , a c c o r d i n g t o t h e P a c i f i c F i s h e r y Management Counci 1 (PFMC 1978 1.

Anchovies a r e g e n e r a l l y s h o r t - l i v e d ; f i s h more t h a n 158 mn l o n g and 4 y e a r s o l d a r e r a r e , b u t anchov ies 229 mn l o n g and 7 yea rs o l d have been r e p o r t e d ( B a x t e r 1967 1.

GROWTH CHARACTERISTICS

The age and g rowth r a t e s of anchov ies i n c e n t r a l and s o u t h e r n C a l i f o r n i a w a t e r s were f i r s t r e p o r t e d by C l a r k and P h i l l i p s ( 1 9 5 2 ) . The f i s h grew 92 mm SL i n t h e f i r s t y e a r o f 1 i f e ; t h e r e a f t e r , s u c c e s s i v e annual i nc remen ts (mn) i n t h e second t h r o u g h f i f t h g row ing seasons were 28, 29, 13, and 4. Anchov ies i n s o u t h e r n C a l i f o r n i a w a t e r s i n t h e f i r s t yea r o f l i f e were g e n e r a l l y s m a l l e r t h a n t h o s e i n c e n t r a l C a l i f o r n i a w a t e r s ( S p r a t t 1975 1.

COMMERCIAL AND BAIT FISHERIES

The n o r t h e r n anchovy i n Cal i f o r n i a s u p p o r t s a commercial

f i s h e r y and a l i v e - b a i t f i s h e r y . I t has a w ide range of uses: human food, b a i t ( l i v e o r dead), f e e d f o r f i s h h a t c h e r i e s and mink farms, and i n d u s t r i a l f i s h meal and o i 1.

Records o f t h e Cal i f o r n i a commercial anchovy c a t c h f r o m 1916 t o 1967 were sumnari zed b y Ta l b o t (1973 1. The annual c a t c h averaged about 325 m e t r i c t o n s ( t ) f r o m 1916 t o 1921; most of t h e f i s h were reduced t o o i l and meal . Enactment o f r e s t r i c t i v e laws i n 1919 and 1921 made i t i m p r a c t i c a l t o c o n t i n u e t o c a t c h f i s h f o r r e d u c t i o n , and annual l a n d i n g s f rom 1922 t o 1938 averaged o n l y 145 t . I n 1939 t o 1946, annual average c a t c h e s i n c r e a s e d t o 1,319 t, and then, as a r e s u l t o f d e c l i n i n g s a r d i n e ca tches , t h e demand f o r anchov ies f o r c a n n i n g i n c r e a s e d and t h e c a t c h r o s e t o 8,591 t b y 1947. A f t e r 1947, more r e s t r i c t i o n s were p l a c e d on t h e anchovy f i s h e r y , and t h e l a n d i n g s dropped f o r t h e n e x t 3 yea rs ; however, t h e boom was s t i l l t o come. The c o l l a p s e o f t h e s a r d i n e f i s h e r y i n C a l i f o r n i a i n 1952 r e s u l t e d i n a sharp i n c r e a s e i n t h e anchovy c a t c h t o 39,000 t i n 1953. Annual ca tches o f ove r 15,000 t c o n t i n u e d u n t i l 1958; consumer acceptance o f canned anchov ies t h e n presumably dw ind led , and t h e annual c a t c h a g a i n d e c l i n e d t o o n l y 1,200 t a n n u a l l y f r o m 1959 t o 1965. P e r m i t s f o r t h e r e d u c t i o n o f anchov ies were a g a i n i s s u e d i n 1966, and a r e c o r d h i g h c a t c h o f 143,000 t was r e p o r t e d i n 1975.

The c e n t r a l s u b p o p u l a t i o n o f anchov ies o f f t h e c o a s t o f C a l i f o r n i a and n o r t h e r n B a j a C a l i f o r n i a s u p p o r t s f i s h e r i e s o f b o t h t h e U n i t e d S t a t e s and Mex ico. The I n s t i t u t i o Nac iona l de Pesca and t h e C a l i f o r n i a Department of F i s h and Game have e n t e r e d i n t o an i n f o r m a l c o o p e r a t i v e program t o m o n i t o r t h e anchovy f i s h e r y (Chavez e t a l . 1977) . A n a l y s i s o f f i s h sampled f r o m t h e Mexico-Cal i f o r n i a l a n d i n g s has i n d i c a t e d m a j o r changes i n age c o m p o s i t i o n f r o m 1977 t o 1980 (Ma is 1981) . B e f o r e 1977, t h e c a t c h was

h e a v i l y dominated b y f i s h o f age groups I and 11; some were even o l d e r . S i n c e 1977, f i s h o f age groups 0 and I have dominated catches, and o l d e r age groups have d i m i n i shed. The o p t imum y i e l d f o r t h e c e n t r a l C a l i f o r n i a p o p u l a t i o n f o r t h e 1981-82 f i s h i n g season was e s t i m a t e d t o be 545,000 t ( S t a u f f e r and Char te r 19821, as s p e c i f i e d b y t h e f o r m u l a g i v e n i n t h e P a c i f i c Management P lan. A c c o r d i n g t o Mais (19811, under t h e p r e s e n t h i g h f i s h i n g p r e s s u r e (and w i t h p r o d u c t i o n n e a r i n g 182,000 t pe r y e a r ) , f a i l u r e s of two success i ve y e a r c l a s s e s c o u l d depress anchovy s t o c k s t o a b n o r m a l l y low l e v e l s .

The s i z e of t h e anchovy p o p u l a t i o n o f f Cal i f o r n i a has changed ove r t h e l a s t t h r e e decades. A marked i n c r e a s e i n abundance c o i n c i d e d w i t h a s t e a d y decrease i n s a r d i n e s i n t h e same area. I t has been e s t i m a t e d t h a t anchovy spawning biomass i n c r e a s e d from 640,000 t i n 1951 t o 5 t o 8 m i l l i o n t i n t h e m id -1960 ' s (Smi th 1972 1. The c e n t r a l s u b p o p u l a t i o n c o n t r i b u t e d about 78% o f t h e t o t a l C a l i f o r n i a biomass (Vrooman and Smi th 1971). As judged b y r e c e n t su rveys of l a r v a l abundance, t h e biomass of a d u l t f i s h i n t h e c e n t r a l s u b p o p u l a t i o n may be as h i g h as 2.7 m i l l i o n t ( S t a u f f e r and C h a r t e r 1982). These f i n d i n g s a r e d i f f i c u l t t o i n t e r p r e t because of t h e c o n f l i c t i n g d a t a on m o r t a l i t y i n t h e c e n t r a l s u b p o p u l a t i o n . M o r t a l i t i e s were e s t i m a t e d b y Hanan (1981) t o be 62% f o r annual m o r t a l i t y and 0.97% f o r i n s t a n t a n e o u s t o t a l m o r t a l i t y . He a1 so i n d i c a t e d t h a t i n s t a n t a n e o u s t o t a l m o r t a l i ty i n c r e a s e d a f t e r 1976-- an i n c r e a s e t h a t c o i n c i d e d w i t h a sha rp decrease of o l d e r anchov ies i n t h e commercial c a t c h and a d e c l i n e i n t h e t o t a l U.S. c a t c h .

The commercial l a n d i n g s o f anchov ies , m o n i t o r e d by t h e C a l i f o r n i a Department o f F i s h and Game, had an e s t i m a t e d ex-vesse l v a l u e o f $3.2 m i l - l i o n i n 1981 (PFMC 1983). More r e c e n t e s t i m a t e s have n o t been pub l i shed .

Much o f t h e v a l u e of t h e c a t c h i s due t o l a n d i n g s f o r r e d u c t i o n . The ex- vesse l p r i c e o f anchov ies v a r i e s c o n s i d e r a b l y i n response t o changes i n domest ic and w o r l d marke ts f o r f i s h meal and o t h e r p r o t e i n meals. Because of t h e g r e a t v a r i a b i l i t y i n f i s h meal demand, t h e anchovy ex-vesse l p r i c e i s expected t o c o n t i n u e t o f 1 u c t u a t e .

The l i v e - b a i . t f i s h e r y f o r anchov ies has c o n t q i b u t e d 98% o f t h e t o t a l l i v e b a i t c a t c h i n C a l i f o r n i a ( B a x t e r 1967). The f i s h e r y i s a c t i v e a t most c o a s t a l p o r t s between San F r a n c i s c o and San Diego, and expans ion of t h i s f i s h e r y t o t h e n o r t h has been a t tempted (Wal dvoge l 1977) . San Diego Harbor i s t h e c e n t e r o f t h e l i v e b a i t i n d u s t r y . L i v e - b a i t ca tches have f 1 u c t u a t e d between 1,500 and 7,000 t a n n u a l l y , and s a l e s were e s t i m a t e d t o have been about $2.3 m i l l i o n pe r y e a r (PFMC 1978) .

ECOLOGICAL ROLE

The n o r t h e r n anchovy cannot be ass igned t o a s i n g l e t r o p h i c l e v e l , l a r g e l y because i t s d i e t c o n s i s t s o f zoop lankton, p h y t o p l a n k t o n , an3 f i s h (Loukashk in 1970) . Nonethe less , t h e anchovy i n a l l l i f e s tages i s p lank tophagous and i t s h o u l d be ass igned t o a low t r o p h i c l e v e l .

On t h e b a s i s o f o b s e r v a t i o n s i n t h e f i e l d and i n t h e l a b o r a t o r y , anchov ies o f a1 1 s i z e s a r e b o t h f i l t e r feeders and p a r t i c u l a t e feeders , depending on t h e f o o d a v a i l a b l e ( M i l l e r 1968) . Anchovies p r o b a b l y feed c h i e f l y d u r i n g t h e day ( B a x t e r 1967). A l though t h e y o l k sac i s absorbed about 1.5 days a f t e r t h e f i s h ha tch , l a b o r a t o r y o b s e r v a t i o n has r e v e a l e d t h a t t h e l a r v a e do n o t feed u n t i l about 2.5 days a f t e r h a t c h i n g (Scu ra and Je rde 1977). Anchovy l a r v a e l o n g e r t h a n 7 rml " a c t u a l l y a t e t h e g r e a t e s t v a r i e t y o f food," a c c o r d i n g t o Be rne r (1959 ) . Stomach c o n t e n t s of f i s h 43 t o 215 mn SL were 72% c r u s t a c e a n s i n v a r i o u s deve lop -

menta l s tages . Zoop lank ton o t h e r t h a n c rus taceans were second i n abundance 1 1 5 % , and i n d e t e r m i n a t e zoop lank- t o n i c rema ins and f l e s h y p a r t s were t h i r d . The r e s t o f t h e d i e t c o n s i s t e d o f ; phy top lank ton (6 .6%) and f o r e i g n m a t t e r (0 .6%) (Loukashk in 1970). Anchovies a l s o sometimes e a t t h e i r own eggs and l a r v a e .

The c h i e f c o m p e t i t o r o f t h e n o r t h e r n anchovy a t a l l l i f e s tages i s t h e P a c i f i c sa rd ine , Sard inops saqax ( B a x t e r 1967) . C o m p e t i t i o n b e g i n s i n t h e l a r v a l s tages and c o n t i n u e s t h r o u g h 1 i f e . Anchovies and s a r d i n e s e a t s i m i l a r foods, and b o t h s p e c i e s a r e most abundant between P o i n t Concept i o n , Cal i f o r n i a , and Magdalena Bay, Ba ja C a l i f o r n i a ( B a x t e r 1967) .

Essent i a1 l y e v e r y p r e d a t o r y f i s h , b i r d , and mammal i n t h e Cal i f o r n i a C u r r e n t e a t s anchov ies . The PFMC (19781, i n a sumnary o f r e p o r t s on anchovy p r e d a t o r s , no ted t h a t anchovy eggs and l a r v a e a r e t h e p r e y o f an asso r tmen t o f i n v e r t e b r a t e and v e r t e b r a t e p l a n k t i v o r e s , i n c l u d i n g a d u l t anchov ies . D u r a t i o n o f t h e p l a n k t o n i c l i f e s t a g e i s o n l y about 2 t o 4 months, and m o r t a l i t y i s h i g h . J u v e n i l e anchov ies near sho re a r e e x t r e m e l y v u l n e r a b l e t o p i s c i v o r e s - - p r i m a r i l y b l u e f i n tuna and a1 bacore. O the r p r e d a t o r s a r e sha rks , p o r p o i s e s , sea ls , and b i r d s . I t has been e s t i m a t e d t h a t a d u l t anchov ies t a k e n a n n u a l l y b y p r e d a t o r s wou ld compose about 73% o f t h e spawning biomass i f no f i s h i n g were done (PFMC 1978).

ENVIRONMENTAL REQUIREMENTS

Temperat u r e

Anchovy l a r v a e , j u v e n i 1 es, and a d u l t s have been observed a t w a t e r t empera tu res r a n g i n g f r o m about 8 t o 25 "C . Eggs have been sampled a t wa te r t empera tu res o f f rom 9.9 t o 23.3 " C ( A h l s t r o m 1956) . Water tem- p e r a t u r e s a t a d e p t h of 10 m were

r e p o r t e d t o be r e p r e s e n t a t i v e of t h e upper mixed l a y e r where eggs t h r i v e d . Most eggs were taken when water temperatures were 13.0 t o 17.5 " C . Anchovy l a r v a e have been taken a t wa te r temperatures o f 10.0 t o 19.7 "C; b u t 95% were taken a t 14.0 t o 17:4 "C (Ah ls t rom 1959). Most l a r v a e 1 i v e above t he thermoc l ine . A d u l t anchovies have been r e g u l a r l y observed a t water temperatures o f 12 t o 20 "C (PFMC 1978); some anchovies appa ren t l y avo i d h i g h su r f ace temperatures because they 1 i v e i n deeper water (Mais 1974). Anchovies u s u a l l y spawn a t water temperatures o f 12 t o 15 "C, which a r e t y p i c a l d u r i n g l a t e w i n t e r (PFMC 1978).

A l though da ta on water temperature and f i s h d i s t r i b u t i o n a re d i f f i c u l t t o i n t e r p r e t , changes i n water temperatures appa ren t l y a f f ec t t h e d i s t r i b u t i o n o f j u v e n i l e and a d u l t anchovies. For example, when average water temperatures a re lower than usual , a d u l t anchovies a re l e s s abundant near shore, and j u v e n i 1 es dominate t h e catches t h e r e (Bax te r 1967 1.

Depth

Adu l t s avo i d su r f ace water d u r i n g t h e day b u t move near t h e s u r f a c e a t

n i g h t (Bax te r 1967). Anchovy l a r v a e t end t o avo id water depths exceeding 48 m (Ah ls t rom 1959). I n c o n t r a s t , Mais (1974) r e p o r t e d t h a t a d u l t anchovies a r e common a t depths o f 183 m o r more d u r i n g t h e day, b u t f r equen t t h e upper 73 m a t n i g h t . C l e a r l y , f u r t h e r i n v e s t i g a t i o n o f t h e depth d i s t r i b u t i o n of t h e n o r t h e r n anchovy and r e l a t e d env i ronmenta l v a r i a t i o n s i s needed.

Other Environmental Fac to r s

I n f o rma t i on about water qual i t y requi rements and p re fe rences f o r t h e anchovy i s scarce. Anchovies of t e n congregate i n areas of sewage o u t f a l l s , and p e r i o d i c d i e - o f f s have been caused by oxygen d e f i c i e n c i e s (PFMC 1978). Anchovies tend t o move away f rom water d e f i c i e n t i n oxygen and avo id h i gh oxygen concen t ra t i ons d u r i n g p l ank ton blooms. Weather may a l s o e x e r t an i n f l u e n c e on water qual i t y and anchovy d i s t r i b u t i o n . Anchovies sometimes leave harbor waters j u s t be fo re heavy w i n t e r storms and h i g h f reshwate r i n f l o w . At tempts t o i n t e r p r e t t h e e f f e c t s o f env i ron- mental v a r i a t i o n s (e.g., temperature, depth, and oxygen) on t h e d i s t r i b u t i o n o f anchovies have been i n c o n c l u s i v e (Lasker and Smith 1977; Brewer and Smith 1982).

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Scura, E.D., and C.W. Jerde. 1977. Va r ious spec ies of p h y t o p l a n k t o n as f o o d f o r l a r v a l n o r t h e r n anchovy, E n q r a u l i s mordax, and r e l a t i v e n u t r i t i o n a l v a l u e o f t h e d i n o - f l a g e l l a t e s G m n o d i n i urn so ledens and Gonyaulax p o l y e d r a . U.S. N a t l . Mar. F i s h . Serv . F i s h . B u l l . 75:577-584.

Smith, P.E. 1972. The i n c r e a s e i n spawning biomass o f n o r t h e r n

anchovy, E n g r a u l i s mordax. U.S. N a t l . Mar. F i s h . S e r v x s h . B u l l .

Soule, D.F., and M. Ogur i , eds. 1972- 1976. Mar ine s t u d i e s o f San Pedro Bay, C a l i f o r n i a , p a r t s 1-12. A l l a n Hancock Foundat ion, U n i v e r s i t y o f Southern Cal i f o r n i a , Los Angeles.

S p r a t t , J.D. 1975. Growth r a t e s o f t h e n o r t h e r n anchovy E n q r a u l i s mordax i n s o u t h e r n Cal i f o r n i a wa te rs , c a l c u l a t e d f r o m o t o l i t h s . C a l i f . F i s h Game 61:116-126.

S t a u f f e r , G.D., and R.L. Char te r . 1982. The n o r t h e r n anchovy spawning biomass f o r t h e 1981-82 C a l i f o r n i a f i s h i n g season. C a l i f . Coop. Oceanic F i s h . I n v e s t . Rep. 23:15-19.

S t a u f f e r , G.D., and K.R. Pa rke r . 1980. E s t i m a t e o f t h e spawning biomass o f t h e n o r t h e r n a w h c v y c e n t r a l s u b p o p u l a t i o n f o r t h e 1978-

79 f i s h i n g season. C a l i f . Coop. Oceanic F i s h . I n v e s t . Rep. 21:12-16.

T a l b o t , G.B. 1973. The C a l i f o r n i a sard ine-anchovy f i s h e r i e s . T rans . Am. F i s h . Soc. 102:178-187.

Vrooman, A.M., and P.E. Smith. 1971. Biomass o f t h e s u b p o p u l a t i o n s o f t h e n o r t h e r n anchovy Enqrau l i s mordax G i r a r d . C a l i f . Coop. Oceanic F i s h . I n v e s t . Rep. 15:49-51.

Waldvogel, J.B. 1977. Age, m a t u r i t y and d i s t r i b u t i o n o f n o r t h e r n anchovy, En r a u l i s rnordax, i n Hum- + -- b o l d t Bav. Ca i f o r n i a . M.S. Thesis. ~ u m b o l d t " - s t a t e U n i v e r s i t y , A rca ta , C a l i f . 3 6 p p .

Wyatt, B., W.V. B u r t , and J.B. P a t t u l l o . 1972. S u r f a c e c u r r e n t s o f f Oregon as de te rm ined f r o m d r i f t b o t t l e r e t u r n s . J. Phys. Oceanogr. 2: 286-293.

PAGE 1 B i o l . R ~ D . 82(11.501* 1 a. r.ti. .W S-OI~II.

Species P r o f i l e s : L i f e H i s t o r i e s and Env i ronmenta l Requirements of Coas ta l F i shes and I n v e r t e b r a t e s ( P a c i f i c Sou thwes t ) - -Uor thern

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U.S. Department o f t h e I n t e r i o r P.O. BO-x 631. Washington, DC 20240 V icksburg , MS 39180

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Species p r o f i l e s a r e l i t e r a t u r e summaries of t h o taxonomy, morphology, d i s t r i b u t i o n , 1 i f e h i s t o r y , and env i r onmen ta l r equ i r emen ts of c o a s t a l a q u a t i c species. They a r e p repa red t o a s s i s t i n env i r onmen ta l impac t assessment. The n o r t h e r n anchovy (Enarau l i s mordax) g e n e r a l l y i s f ound i n c o a s t a l waters. It moves o f f s h o r e i n w i n t e r and r e t u r n s i n s h o r e i n sp r i ng . A l t hough anchov ies spawn t h roughou t t h e year , most spawn i n w i n t e r and sp r i ng . Few n o r t h e r n anchov ies l i v e l o n g e r t h a n 7 yea rs . Commercial ca t ches (1916-52) f o r C a l i f o r n i a ranged f r o m 27 t o 143,000 t. Anchovies a r e p r i m a r i l y p lanktophagous i n a l l 1 i i e s tages ; t h e i r d i e t c o n s i s t s o f zoop lank ton , phy top lank ton , and f i s h . Anchovy have been c o l l e c t e d a t wa te r tempera tu res o f 8 t o 25 "C.

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REGION 1 Regional Director U.S. Fish and Wildlife Service Lloyd Five Hundred Building, Suite 1692 500 N.E. Multnornah Street Portland, Oregon 97232

REGION 4 Regional Director U.S. Fish and Wildlife Service Richard B. Russell Building 75 Spring Street, S.W. Atlanta, Georgia 30303

REGION 2 REGION 3 Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service P.O. Box 1306 Federal Building, Fort Snelling Albuquerque, New Mexico 87 103 Twin Cities, Minnesota 55 1 1 1

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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 resources, protecting our fish and wildlife, preserving theenvironmental 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 people who live in island territories under U.S. administration.