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U. S. Fish and Wildlife Service Biological Report 82 (1 1.58) Nationa 1 Wetlands Research Center TR EL-82-4

June 1986 700 Cajun Dome Boulevard Lafayette, Louisiana 70506

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Florida)

KING AND SPANISH MACKEREL

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 Report 82(11.58) TR EL-82-4 June 1986

Species P ro f i les : L i f e H i s t o r y and Environmental Requirements o f Coastal Fishes and Inve r teb ra tes (South F l o r i d a )

KING MACKEREL AND SPANISH MACKEREL

Mark F. Godcharles and

Michael D. Murphy D i v i s i o n o f Marine Resources

F l o r i d a Department o f Natural Resources 100 8 t h Ave. S.E.

St. Petersburg, FL 33701

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

Nat iona l Coastal Ecosystems Team U.S. F i sh and W i l d l i f e Serv ice

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. Army Corps o f Engineers

Vicksburg, MS 39180

and

Nat iona l Coastal Ecosystems Team Research and Development F i s h and W i l d l i f e Serv ice

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

This s e r i e s should be referenced as fo l lows:

U. S. F i s h and W i l d l i f e Service. 1983-19 Species p r o f i l e s : 1 i f e h i s t o r i e s and environmental requirements o f c o a s t a r f i shes and inver tebra tes . U. S. F i s h Wi ld l . Serv. B i o 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 fo l lows:

Godcharles, M. F. , and M. D. Murphy. 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 requirements o f coas ta l f i s h e s and i nve r teb ra tes (south F l o r i d a ) -- k ing mackerel and Spanish mackerel. U. S. F i s h W i l d l . Serv. B i o l . Rep. 82(11.58). U. S. Army Corps o f Engineers, TR EL-82-4. 18 pp.

PREFACE

This 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 spo r t , commercial , o r eco log i ca l importance. The p r o f i l e s 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 the 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 he 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 sec t ions on taxonomy, 1 i f e h i s t o r y , eco log i ca l r o l e , environmental requirements, and economic importance, i f app l icab le . A t h r e e - r i ng b inde r i s used f o r t h i s se r i es so t h a t new p r o f i l e s can be added as they a re prepared. This 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 quest ions regard ing t h i s r e p o r t should be d i r e c t e d t o one of the 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 iona l 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 1 , 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

CONVERSION T A B L E

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

Mu1 t i p l y !?L To O b t a i n

m i l l i r ne te rs (mm) c e n t i m e t e r s (n) mete rs (m) k i 1 ometers ( km)

2 square m e t e r s (m ) 10.76 square k i 1 m e t e r s ( km2) 0.3861 h e c t a r e s ( h a ) 2.471

l i t e r s ( 1 ) c u b i c m e t e r s (m3) c u b i c me te rs

i nches inches f e e t m i l es

square f e e t square ( n i l es acres

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

m i l 1 i g rams (mg) 0.00003527 ounces grams ( g ) 0.03527 ounces k i l o j ra lns ( k g ) 2.205 pounds m e t r i c tons ( t ) 2205.0 pounds m e t r i c t o n s 1.102 s h o r t t o n s k i 1 oca l o r i e s ( k c a l ) 3.968 B r i t i s h thermal u n i t s

C e l s i u s degrees 1.8("C) + 32 F a h r e n h e i t degrees

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

i nches 25.40 i nches 2.54 f e e t ( f t ) 0.3048 fathoms 1.829 m i l e s ( m i ) 1.609 n a u t i c a l m i l e s ( m i ) 1.852

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

g a l 1 ons ( g a l ) c u b i c f e e t ( f t 3 ) a c r e - f e e t

ounces ( 0 2 ) 28.35 pounds ( l b ) 0.4536 s h o r t t o n s ( t o n ) 0.9072 B r i t i s h thermal un i t s ( B t u ) 0.2520

m i l 1 i m e t e r s c e n t i m e t e r s me te rs me te rs k i l o m e t e r s k i 1 o n e t e r s

squa re me te rs h e c t a r e s square k i 1 ometers

1 i t e r s c u b i c me te rs c u b i c me te rs

grams k i 1 og rams m e t r i c t o n s k i 1 oca l o r i e s

F a h r e n h e i t degrees 0.5556("F - 32) C e l s i u s degrees

CONTENTS

Page

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . PREFACE CONVERSIONTABLE . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . TABLES . . . . . . . . . . . . . . . . . . . . . . . . . . . . ACKNOWLEDGMENTS

NOMENCLATURE/TAXONOMY/RANGE . . . . . . . . . . . . . . . . . . . . . . MORPHOLOGY/IDENTIFICATION AIDS . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . REASON FOR INCLUSION I N SERIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . LIFE HISTORY

Seasonal D i s t r i b u t i o n and Populat ions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spawning

M a t u r i t y and Fecundity . . . . . . . . . . . . . . . . . . . . . . . Eggs. Larvae. and Juven i les . . . . . . . . . . . . . . . . . . . . . Longevity and Growth . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . COMMERCIAL AND SPORT FISHERIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . ECOLOGICAL ROLE

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Food H a b i t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Predators

. . . . . . . . . . . . . . . . . . . . . . ENVIRONMENTALREQUIREMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temperature

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S a l i n i t y

. . . . . . . . . . . . . . . . . . . . . . . . . . . . LITERATURE CITED

iii i v v i

v i i

TABLES

Number Page

1 M e r i s t i c c h a r a c t e r i s t i c s o f mackerels i n F l o r i d a . . . . . . . . . 4

2 Regressions o f f ecund i t y on t o t a l weight and f o r k l eng th f o r k i n g mackerel and Spanish mackerel. . . . . . . . . . . . . . . 7

3 Mean back-calculated f o r k lengths o f k i n g mackerel and Spanish mackerel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

4 Von B e r t a l a n f f y growth parameters f o r k i ng mackerel and Spanish mackerel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

5 Length-weight r e l a t i o n s f o r k i n g mackerel and Spanish mackerel. . . 10

6 F l o r i d a commercial landings o f k i n g and Spanish mackerel i n t he Gu l f o f Mexico and along the A t l a n t i c coast, 1960-83. . . . . . . . 12

ACKNOWLEDGMENTS

We a re g r a t e f u l f o r rev iews by J. Connor Davis, F l o r i d a Marine F i s h e r i e s Commission; Roy W i 11 iams, F l o r i d a Department of Natura l Resources; Dr. B i l l Seaman, U n i v e r s i t y o f F l o r i d a ; and s t a f f o f t h e Na t i ona l Mar ine F i s h e r i e s Se rv i ce Laboratory i n Panama City, FL.

King mackerel

Spanish mackerel

Figure 1. King and Spanish mackerel ( f r a n Goode 1884).

KING MACKEREL AND SPANISH MACKEREL

S c i e n t i f i c name. . Scomberomorus caval l a (Cuvier)

p re fe r red - common name.. . k ing mackerel (F igure 1)

Other common names. ......... k i ng f i sh, k ings, serucho

S c i e n t i f i c name........ . Scomberomorus maculatus (M i t ch i 11 )

Pre fer red common name. ........ Spanish mackerel (F igure 1)

Class .................... Osteichthyes Order ..................... Perciformes Family ..................... Scombridae

Geographic range: K ing mackerel i n h a b i t A t l a n t i c coas ta l waters from the G u l f o f Maine t o Rio de Janei ro , Braz i 1 , i n c l u d i n g t h e Gulf o f Mexico and t h e Caribbean Sea ( B r i ggs 1958; Beaumari age 1973). They a re concentrated o f f t h e coast o f t he Carol inas i n t h e spr ing , summer, and f a l l ; i n the no r the rn Gulf o f Mexico from Texas t o nor th - west F l o r i d a i n summer; and o f f southern F l o r i d a and Louis iana i n w in te r . The d i s t r i b u t i o n s and concentrat ions i n south F l o r i d a are shown i n F igure 2.

T. PETERSBURG FT. PIERCE

Coasta l d istr ibut ion

Areaa of high concentrat ion

Figure 2. Distribution of mackerel along the coast of South Florida.

2

Spanish mackerel i n h a b i t coas ta l waters o f t h e western A t l a n t i c Ocean from t h e Gu l f o f Maine t o t he Yucatan Peninsula (Col l e t t e e t a1 . 1978). They a re common as f a r n o r t h as Chesapeake Bay i n summer (Bigelow and Schroeder 1953). Large schools a re common i n south F l o r i d a waters i n l a t e f a l l and w i n t e r (F igure 2).

MORPHOLOGY/IDENTIFICATION AIDS

King Mackerel

The elongated, fus i fo rm, compressed body o f k i n g mackerel i s 4.25 t o 5.00 t imes as long as t h e head, and 5.50 t o 6.25 t imes as l ong as i t i s deep (Be r r i en and Finan 1977a). They have two dorsa l f i n s , t h e second fo l lowed by e i g h t o r n ine f i n l e t s . The l a t e r a l l i n e , which a b r u p t l y curves downward below t h e second dorsa l f i n , d i s t i ngu i shes t h e k i n g mackerel from the Spanish mackerel and t h e cero mackerel (S. rega l i s ) . The caudal peduncle h a s a l a r g e f l eshy keel. The body i s e n t i r e l y scaled w i t h rudimentary scales, except f o r most o f t h e pec to ra l f i n . The mouth i s l a r g e and ob l i que w i t h t h e maxi1 1ary reaching p o s t e r i o r l y t o s l i g h t l y beyond t h e eye o r b i t . Each s ide o f t h e jaw has about 30 s t rong l y compressed t r i a n g u l a r tee th . Th is species gets considerably 1 arger than the Spanish mackerel.

'The dorsa l sur face o f t h e k i n g mackerel i s b lack w i t h i r i d e s c e n t hues o f green and blue. The l a t e r a l and ven t ra l s ides a re s i l v e r y whi te. The l a t e r a l ye1 low s p o t t i n g on t h e young i s almost always absent on mature specimens (Be r r i en and F i nan 1977a).

second f i n i s concave and o r i g i n a t e s a s h o r t d is tance i n f r o n t o f the anal f i n , which i s s i m i l a r i n form and s ize. These are fo l lowed by e i g h t o r n ine dorsa l and anal f i n l e t s . The l a t e r a l 1 i n e i s wavy, and t h e caudal peduncle i s keeled. The caudal f i n i s lunate, and t h e pec to ra l s a re no t covered w i t h scales. See Table 1 f o r o the r key d iagnos t i c characters.

Spanish mackerel a re dark b l u e o r blue-green above, p a l e and s i l v e r y below. The i r s ides a re marked w i t h many smal l , oblong, d u l l orange o r y e l l o w i s h spots t h a t a re prominent bo th above and below t h e l a t e r a l l i n e . The membrane i s b lack on t h e a n t e r i o r one - th i rd o f t h e f i r s t dorsa l f i n and t h e p o s t e r i o r i s greenish-whi t e . The second dorsa l and the pec to ra l f i n s a re p a l e y e l l o w w i t h dusky edges. The anal and v e n t r a l f i n s a re wh i te (Be r r i en and Finan 1977b).

REASON FOR INCLUSION I N SERIES

King and Spanish mackerel support commercial and spo r t f i s h e r i e s o f major importance. Both species i n - h a b i t coas ta l waters b u t Spanish mackerel u s u a l l y come c l o s e r t o shore, e.g., along t h e beaches and i n t h e ou te r waters o f es tuar ies . I n t h e summer, l a rge so l i t a r y k i n g mackerel a re sometimes captured o f f p i e r s and near deepwater i n l e t s . Juveni 1 e k i n g mackerel sometimes mix w i t h schools o f Spanish mackerel. Both species feed p r i n c i p a l l y on estuary-dependent spe- c i e s such as menhaden (B revoo r t i a spp. ) and anchovies (Anchoa spp. ). Excessive a l t e r a t i o n s o f es tua r ies cou ld reduce t h e product ion o f coas ta l forage f i s h on which k i n g and Spanish mackerel feed.

Spanish Mackerel LIFE HISTORY

Spanish mackerel have compressed, Seasonal D i s t r i b u t i o n and Populat ions elonaated bodies about 4.5 t o 5.0 time; as long as they a re deep. The K ing mackerel. Extensive tagg ing f i r s t dorsa l f i n i s t r i a n g u l a r ; t h e s tud ies i n d i c a t e t h a t t h e r e a re a t

Table 1. M e r i s t i c c h a r a c t e r i s t i c s o f mackerels i n F l o r i d a (Mago-Leccia 1958).

K i ng C h a r a c t e r i s t i c mackerel

Spanish Cero mackerel mackerel

To ta l number o f ver tebrae

k i r s t c losed haemal arch a t ve r teb ra number

F i r s t haemal spine a t ve r teb ra number

Dorsal spines

Dorsal rays

Dorsal f i n l e t s

Anal spines

Anal rays

Anal f i n l e t s

+- Angle G i l l rakers

Lower 1 imb

l e a s t two m ig ra to ry groups o f k i n g mackerel. Those tagged i n t h e w i n t e r south o f Cape Canaveral , F lo r i da , usual l y moved southward through the F l o r i d a Keys, entered the Gu l f o f Mexico i n l a t e w i n t e r and spr ing , and then cont inued northward a long the west F l o r i d a Cont inenta l Shel f . Some were recaptured as f a r west as Texas and a few as f a r south as Veracruz and Yucatan, Mexico. These western migrants, some tagged o f f t he coasts o f Texas and northwest F l o r i d a i n t he summer, re turned t o south F l o r i d a i n t h e w i n t e r ( W i l l iams and Tay lor 1978;

Wi l l iams and Suther land 1979; Suther l and and Fable 1980).

A second group o f k i n g mackerel, tagged i n the s p r i n g o f f southeast F l o r i d a , moved northward i n the summer a long t h e A t l a n t i c coas t t o t he Carol inas. One was recaptured as f a r n o r t h as Chincoteague, V i r g i n i a . King mackerel tagged i n South Caro l ina i n the sp r i ng genera l l y moved south i n May through August and were recap- t u r e d i n F l o r i d a t h a t summer. I n the f a l l they migrated t o t he nor thern 1 i m i t o f t h e i r d i s t r i b u t i o n , and many

were recaptured i n Nor th Caro l ina ( W i 11 iams and Suther land 1979; W i 11 iams and Godcharl es 1983).

Pre l im inary e l e c t r o p h o r e t i c anal- yses o f 48 enzymes ex t rac ted from k i n g mackerel t i ssues (hear t , 1 i v e r , eye, and muscle) g i ve evidence t h a t t h e r e a re two popu la t ions o f k i n g mackerel. V a r i a t i o n o f two a l l e l e s f o r t h e pep- t i d a s e w i t h g l y c y l leuc ine-2 locus gave the g rea tes t d i f f e rences between South Carol i na and Texas specimens ; however, o ther l o c i were min imal ly polymorphic and o f l i m i t e d value f o r separa t ing popu la t ions (May, unpubl . MS.). Homing tendencies are a d d i t i o n a l evidence o f two populat ions. A f t e r a year o r more o f freedom, k i n g mackerel u s u a l l y a re recaptured a t o r near t h e i r re lease s i t e a t about t he same t ime o f year they were f i r s t tagged ( W i 1 1 i ams and Godcharl es 1983).

Spanish mackerel. Spanish mack- e r e l are a l so migra tory , general l y moving northward each sp r i ng , spending summer i n t h e nor thern art o f t h e i r range, and m ig ra t i ng south i n f a l l . I n spr ing , summer, and f a l l , they are most abundant i n t h e no r the rn Gu l f o f Mexico and along t h e eas t coast o f t h e Un i ted States up t o V i r g i n i a (Wollam 1970; Dwinel l and Futch 1973; Powell 1975). The i r major w i n t e r i n g grounds a re o f f south F l o r i d a , b u t some over- w i n t e r o f f t h e eas t coast o f Mexico (Mendoza 1968; Suther l and and Fable 1 980).

Spanish mackerel from d i f f e r e n t geographic areas may mix i n south F l o r i d a du r ing the w in ter . I n summer, t h e south A t l a n t i c and eas tern G u l f o f Mexico popu la t ions spawn i n i s o l a t i o n i n the no r the rn p a r t s o f t h e i r ranges (Wol lam 1970). E lec t ropho re t i c pa t - t e rns o f two hemoglobin phenotypes demonstrated t h a t nor thwestern g u l f f i s h a re d i s t i n c t from f i s h captured a long t h e U.S. eas t coast (Skow and Chi t tenden 1981).

Spawni ng

King mackerel. This species spawns i n t he coasta l waters o f t h e no r the rn G u l f o f Mexico, and o f f t h e south A t l a n t i c coast (Dwinel l and Futch 1973; McEachran e t a l . 1980; Burns 1981 ; Powles, unpubl. MS. ). Because of a p r o t r a c t e d spawning season, l a r vae have been c o l l e c t e d by sampling gear from May through October. Catches were h ighes t i n September. The p a u c i t y o f l a r vae i n t h e eastern G u l f o f Mexico south o f Cape San Blas, o f f t h e Yucatan Channel and southeast F l o r i d a , i n d i c a t e s m in i - mal spawning i n these waters (Wol lam 1970; Houde e t a l . 1979; Burns 1981).

King mackerel l a r vae were d iscov- ered o f f nor thwest F l o r i d a and Texas between t h e Middle and Outer (35-183 m) Cont inenta l She l f (Dwinel l and Futch 1973; McEachran e t a l . 1980). Larvae captured near Palm Beach, F l o r i d a , were c l o s e r t o shore than those captured f a r t h e r n o r t h o f f Cape Canaveral, F l o r i d a , Savannah, Georgia, and Cape Fear, Nor th Carol ina. Nor th o f Cape Canaveral , 1 arvae were found near o r o f f t h e s h e l f a long t h e 200 m depth contour and near t h e Gu l f Stream (Wollam 1970; Burns 1981; Powles, unpubl. MS.).

Fu r the r evidence o f p r o t r a c t e d spawning f o r k i n g mackerel i s t h e ho ld ing o f v i t e l l ogenic eggs (eggs w i t h forming y o l k ) from May through October (Beaumariage 1973). Mu1 t i p l e spawning i s suggested by bimodal d i s - t r i b u t i o n s o f mean oocyte diameter o f yo1 kbear ing v i t e 1 logen ic (s tage 4) eggs. The f i r s t mode was i n l a t e May through e a r l y J u l y and t h e second was i n l a t e J u l y and e a r l y August. The appearance o f spent males from September throqgh December co inc ided w i t h a commensurate absence o f v i t e l - logen ic eggs beginning i n August, suggesting a f i n a l c l i m a c t i c spawn.

Spanish mackerel . Spanish mackerel a l so spawn over a p r o t r a c t e d season (Powel 1 1975). ~ i p e females

have been c o l l e c t e d from A p r i l through September i n F l o r i d a waters (Kl ima 1959; Powel 1 1975; Finucane and Col 1 i ns , unpubl . MS. ) . Larvae have been c o l l e c t e d from s p r i n g u n t i l l a t e summer i n t h e eas tern G u l f o f Mexico (Dwinel l and Futch 1973; Houde e t a l . 1979), and from May through mid- September from Cape Fear, Nor th Carol i nay t o Cape Canaveral , F l o r i d a (Pow1 es , unpubl . MS. ).

The onset o f spawning progresses from south t o nor th . Spawning begins i n A p r i l i n t h e Carol tnas, i n mid-June i n Chesapeake Bay, and from l a t e August t o l a t e September o f f Sandy Hook, New Jersey, and Long I s l a n d , New York (Ear l 1 1883). Beaumariage (1970) repo r ted t h a t few mackerel spawned a t water temperatures below 26 OC. I n Texas, mackerel spawned when water temperatures exceeded 25 O C and a t s a l i n i t i e s between 30 and 36 p p t (Hoese 1965). Spanish mackerel apparent ly spawn a t n i g h t ( E a r l l 1883; Smith 1907).

Col l e c t i o n s o f small 1 arvae i n d i c a t e t h a t Spanish mackerel spawn over t he Inne r Cont inenta l She l f i n waters 12-34 m deep (McEachran e t a1 . 1980). I n t h e nor theastern G u l f o f Mexico o f f nor thwest F lo r i da , most l a r vae were c o l l e c t e d i n water shal lower than 13 m y a l though some were i n water as deep as 91.5 m (Dwinel l and Futch 1973). A l l l a r vae c o l l e c t e d a long t h e west coast o f F l o r i d a by Houde e t a l . (1979) were taken i n s i d e t h e 50 m depth contour; most were w i t h i n t h e 20 m contour.

M a t u r i t y and Fecundi ty

I n south F l o r i d a , most male k i n g mackerel probably spawn i n t h e i r f o u r t h year o f l i f e (Age 111) when about 718 mm f o r k l e n g t h (FL); most females spawn a t Age I V when about 857 mm FL (Beaumariage 1973). Finucane e t a l . (unpubl. MS.) repor ted mature fe - males. about 600 mm FL d i s t r i b u t e d from Texas t o t he Carol inas, w i t h egg count est imates rang ing from 69,000 (446 mm

FLY Age I, 0.68 kg) t o 12,207,000 (1,489 mm FLY Age X I I I , 25.6 kg). The bes t i n d i c a t o r o f fecund i ty i s t he t o t a l weight o f t he f i s h (Finucane e t a l . unpubl . MS). Fecundi ty -we ight equations are l i s t e d i n Table 2.

Spanish mackerel i n south F l o r i d a become sexua l ly mature i n t h e i r second and t h i r d year o f 1 i f e (Ages I and 11) when about 250 t o 350 mm FL. I n southeast F lo r i da , t h e l eng th o f most mature males and females ranges between 325 and 349 mm FL; a l l f i s h l a r g e r than 375 mm FL were mature. Kl ima (1959) found mature females as small as 250 mm FL and mature males between 280 and 340 mm FL. He e s t i - mated t h a t most o f these f i s h were 1 o r 2 years o ld. On t h e o the r hand, Powell (1975) repor ted t h a t Kl ima overest imated a l l ages by one year because he misread the f i r s t annul us ; the re fo re , f i s h l ess than 1 year o l d may have been mature. Many Age I f i s h had r i p e oocytes, b u t observat ions i n t h e Apr i 1 -September spawning season suggested t h a t t h e eggs o f Age I f i s h were n o t advanced enough t o be spawned t h a t season (Powel 1 1975). Spanish mackerel Age I11 and o l d e r cons t i - t u t e t h e b u l k o f t he spawning stock.

The fecund i t y o f Spanish mackerel (Table 2) i n southeast F l o r i d a i n - creases w i t h i nc reas ing l e n g t h and weight (F i nucane and Col 1 i ns , unpubl . MS.). Egg number est imates ranged from 194,000 t o 1,491,000 f o r females 354 t o 664 mm FL. E a r l l (1883) re - por ted 1.5 m i l l i o n eggs from a 6 - l b (2.7- kg) female c o l 1 ec ted from Chesapeake Bay.

Eggs, Larvae, and Juveni 1 es

Examination o f k i n g and Spanish mackerel l a r vae 2.0 t o 2.9 mm standard l e n g t h (SL) o f f e r e d d iagnos t i c a ids f o r separa t ing t h e l a rvae based on pigment (me1 anophore) d i f f e rences i n t he jaw, head, and nape areas (Richardson and McEachran 1981). The myomere counts, mouth, t ee th , preoper- c u l a r spines, f i n elements, and p ig -

Table 2. Regressions o f f e c u n d i t y (F) on t o t a l weight (TW) and f o r k l e n g t h (FL) f o r k i n g mackerel (Finucane e t a1 . , unpubl . MS. ) and Spanish mackerel ; r i s t h e c o r r e l a t i o n c o e f f i c i e n t (Finucane and C o l l i n s , unpubl. MS.).

Equation Number o f

100r2 f i s h

KING MACKEREL (SOUTHEAST U. S. )

F = 1.854 x 10 (TW) 1.361 85.6 6 5

F = 4.391 x (FL) 3.974 82.0 64

SPANISH MACKEREL (SOUTHEAST FLORIDA)

F = 9.076 x l o 2 (TW) 0.919 94.1 11

F = 1.027 x (FL) 2.863 92.5 11

mentat ion o f l a r g e r k i ng mackerel l a r vae (3.3 t o 17.0 mm SL) were desc r i bed by Wol lam (1970). King mack- e r e l eggs have n o t y e t been descr ibed.

King mackerel. Because most k i n g mackerel l a r vae are c o l l e c t e d near t he sur face , t he ref inement o f quan t i t a - t i v e sampl i n g techniques t o c o l l e c t l a r vae 3 mm SL (about 3 days o l d ) would a i d i n t h e d e l i n e a t i o n o f spawning grounds (McEachran e t a l . 1980). I n t he g u l f , mackerel l a r vae have been taken a t surface s a l i n i t i e s and temperatures from 27 t o 36 p p t and 26 t o 3 1 OC, and i n t he south A t l a n t i c between 30 and 37 p p t and 22 t o 28 OC

(Dwinel l and Futch 1973; Powles, unpubl . MS. ).

Spanish mackerel. The eggs and newly hatched l a r vae o f Spanish mack- e r e l have been descr ibed by Ryder (1882). The eggs a re pe lag i c , smooth, and t ransparent , w i t h a s i n g l e o i l d rop le t . Eggs are round and about 1 mm (0.9 - 1.3 mm) i n diameter; t h e p e r i v i t e l l i n e space ( t h e c l e a r space

i n s i d e t h e egg between the cho r i on and t h e o i l g lobu le ) i s about 0 .1 mm across, and the o i l g l obu le i s 0.25 mm i n diameter. Hatching takes p lace a f t e r about 25 h a t a temperature o f 26 O C (Smith 1907). Larvae longer than 3 mm SL were descr ibed by Wollam (1970). Most l a r vae have been c o l - l e c t e d i n coas ta l waters o f t he G u l f o f Mexico (Wollam 1970; McEachran e t e l . 1980) and the eas t coas t o f t he Un i ted Sta tes (Powles, unpubl . MS. ).

Juveni 1 e Spanish mackerel have been c o l l e c t e d from low s a l i n i t y es tua r i es and h igh s a l i n i t y beach waters. A Spanish mackerel 58 mm l ong was c o l l e c t e d from Sabot Pond, Louis iana, a t a s a l i n i t y o f 0.2 p p t (Kel l e y 1965). Juven i l es (133 t o 158 mm SL) were c o l l e c t e d a long h igh s a l i - n i t y (33.8 p p t ) beaches and low s a l i - i n i t y (12.8 t o 19.7 pp t ) bayous i n Tampa Bay, F l o r i d a (Spr inger and Wood- burn 1960). Apparent ly , some j uven i 1 e Spanish mackerel use es tua r i es as nursery grounds, b u t most s tay near- shore i n open beach waters.

Longevity and Growth The von B e r t a l a n f f y growth equation i s as fo l lows:

K in mackerel. A1 though Beau- mari age 4-i- 1973 estimated age, growth, and m o r t a l i t y o f k ing mackerel, recent est imates (Johnson e t a l . 1983) are probably more accurate because the data base inc luded more o f t h e l a r g e r and o lde r k i n g mackerel c o l l e c t e d from Nor th Caro l ina t o Texas. Most o f Beaumariagels (1973) specimens came from w in te r c o l l e c t i o n s i n South F lo r i da . The mean back-calculated f o r k lengths a re l i s t e d i n Table 3.

where 1 i s l e n g t h a t age t, L, i s asymptot.c length , K i s the growth c o e f f i c i e n t , and t i s the age when t h e t h e o r e t i c a l l eng th i s zero (Table 4)

Both Beaumariage (1973) and Johnson e t a l . (1983) aged k i n g mack- e r e l by using o t o l i t h s . Although

Tab1 e 3. Mean back-cal cu la ted f o r k 1 engths (mm) o f k ing mackerel (Beaumariage 1973; Johnson e t a l . 1983) and Spanish mackerel (Kilma 1959; Powell 1975). Beaumariage's lengths were converted t o f o r k lengths using h i s r e l a t i o n FL = 1.096(SL) - 17.143. Powel 1 I s lengths were converted by us ing h i s r e l a t i o n FL = 1.073(SL) + 2.427. Lengths f o r female k i n g mackerel reported by Johnson e t a l . (1983) excluded Louis iana specimens.

Ma1 es Females Beaumariane Johnson e t a l . Beaumariage Johnson e t a l .

K i ng mackerel

I11 I v v VI VI I VIII I X X

Femal es Kilma Powel 1

Spanish mackerel

t h e r e was good c o r r e l a t i o n between the growth o f o t o l i t h s and l e n g t h o f f i s h , n e i t h e r study c l e a r l y demonstrated t h a t opaque o t o l i t h growth r i n g s were v a l i d i n d i c a t o r s o f age (Powers and E ldr idge, unpubl. MS. a).

Females l i v e longer than males and u s u a l l y grow f a s t e r a f t e r Age I 1 (Table 3). The o l d e s t females c o l - l e c t e d were 14 years o f age o r o lder : a 1.4 m FL f i s h from Louis iana (Johnson e t a l . 1983) and a 90 l b (40.8 kg) f i s h from Key West (Beaumariage, unpubl . data). The o l d e s t male (979 mm SL) was 12 years o l d (Johnson e t a l . 1983).

King mackerel growth i s h i g h l y va r i ab le . For example, females 850 t o 899 mm FL cou ld be 1 t o 8 years o l d ; males about t he same s i z e cou ld be 3 t o 8 years o ld . Johnson e t a l . (1983) suggested t h a t compensatory growth occurs i n "s low growing" f i s h du r ing t h e i r second year when t h e i r growth increment surpasses t h a t o f " f a s t growers. I' Nevertheless , "s low growing" mackerel remain small e r than " f a s t growers" throughout t h e i r l i v e s .

Equations r e l a t i n g weight t o l eng th o f k i n g and Spanish mackerel a re l i s t e d i n Table 5. The t o t a l annual m o r t a l i t y est imate by Johnson e t a l . (1983) was 0.37; Beaumariage (1973) repor ted 0.54.

Spanish mackerel. The growth o f l a r v a l and l u v e n i 1 e S ~ a n i s h mackerel has no t b&n measured i n t h e labor - a tory . The p r o t r a c t e d spawning season makes i t d i f f i c u l t t o est imate growth from leng th d i s t r i b u t i o n s . Hi ldebrand and Cable (1938) c o l l e c t e d l a rvae 4 mm long as e a r l y as June o f f Nor th Carol ina; some j u v e n i l e s were 80 mm 1 ong by October.

By determining age and growth from o t o l i t h s i n south F l o r i d a , Powell (1975) and Kl ima (1959) repor ted t h a t females grow f a s t e r than males (Table 3). The average l eng th o f Age I 1 f i s h reported by' Klima (1959) , however, i s

about equal t o t he average l eng th o f Age I f i s h reported by Powell (1975). One author probably misread the f i r s t annulus. Growth parameters f o r each sex a re i n Table 4.

Spanish mackerel o f t he same age a re smal le r i n t he southwestern G u l f o f Mexico than i n south F l o r i d a (Powel 1 1975). Mendoza (1968) and Doi and Mendizabal (1979) a l s o used o t o l i t h s t o determine age, b u t t he data f o r bo th sexes were combined. Mendoza (1968) repor ted average f o r k lengths (FL) o f 333, 408, 471, 543, and 593 mm f o r Ages 1 1 - V I , respec- t i v e l y , and an est imated asymptot ic l e n g t h o f 860 mm FL. Est imated average t o t a l lengths (TL) repor ted by Doi and Mendizabal (1979) f o r Ages I - V I were 262, 426, 475, 512, 575, and 638 mm, r espec t i ve l y .

The l i f e span o f Spanish mackerel i s about 5 t o 8 years (Kl ima 1959; Powell 1975; Doi and Mendizabal 1979). The t o t a l annual m o r t a l i t y r a t e based on Powe l l ' s (1975) data was est imated a t 0.62 (GM&SAFMC 1982), which approx- imates t h e r a t e ca l cu la ted by Doi and Mendizabal (1979) f o r Spanish mackerel taken o f f t h e Mexican coast (0.59). Weight- length r e l a t i o n s f o r Spanish mackerel a re g iven i n Table 5.

COMMERCIAL AND SPORT FISHERIES

K i ng Mackerel

F l o r i d a has h i s t o r i c a l l y produced about 90% o f t he k i n g mackerel commer- c i a l land ings i n t h e Un i ted States (Table 6). The species i s a l s o h i g h l y regarded as a s p o r t f i sh . Sport f i s h - ermen i n t h e south A t l a n t i c reg ion a re est imated t o have caught 598,000 k i n g mackerel i n 1979 and 1,370,000 i n 1980. I n t h e g u l f , about 600,000 were taken i n 1979 and 1 m i l l i o n i n 1980 (Trent e t a l . 1983). A t t imes, keen compet i t ion f o r t h i s resource has l e d t o ser ious user c o n f l i c t s .

Table 4. Von B e r t a l a n f f y growth parameters f o r k i n g mackerel (Beaumariage 1973; Johnson e t a l . 1983) and Spanish mackerel (Powell 1975). Johnson e t a l . d i d not i nc lude la rge, most ly female f i s h captured o f f Louisiana i n data used t o est imate growth parameters. Beaurnariage' s and Powel 1 ' s est imates o f asymptotic length (Lm) were converted t o f o r k lengths by us ing t h e i r length- length regressions (Table 3). K i s t he growth c o e f f i c i e n t .

Species and sex

Growth parameters K Lm(mm FL) to(years) Source

King mackerel Ma1 es 0.35 903 -2.50 Beaumariage 1973

0.28 965 -1.17 Johnson e t a l . 1983 Females 0.21 1,243 -2.40 Beaumari age 1973

0.29 1,067 -0.97 Johnson e t a l . 1983

Spani sh mackerel Ma1 es 0.48 555 -1.12 Powel 1 1975 Females 0.45 694 -0.78 Powell 1975

Tab1 e 5. Length-weight r e l a t i o n s f o r k ing mackerel and Spanish mackerel. Weights (W) a re i n grams and lengths (L) i n m i l l ime te rs .

Species Length Number o f W = aL b

and sex measure* f i s h a b Source

King mackerel Ma1 es S L 237 1.330 x 101: 2.9372 Beaumari age 1973

F L 701 0.8064 x 2.9928 Johnson e t a l . 1983 Females S L 292 3.907 x 3.1256 Beaumariage 1973

FL 2,023 0.8801 x 10 2.9827 Johnson e t a l . 1983

Spanish mackerel Ma1 es S L 135 1.1519 x 1012 2.9822 Powel 1 1975 Females S L 217 4.7491 x 10 3.1373 Powel 1 1975

%L = Standard 1 ength FL = Fork length.

Major commerci a1 catches a1 ong the Florida ea s t coast a re centered between Cape Canaveral and Palm Beach, and on the west coast from Key West t o Naples (Beaumariage 1973). A1 though king mackerel support a year-round f ishery, most a re caught i n winter and ea r ly spring. The percentages of the to ta l catch taken from 1950 t o 1974 along the ea s t coast of Florida were 67% by t r o l l i ng , 29% by runaround gi 11 nets , and 4% by handlines. The percentages f o r the west coast were 56% f o r runaround g i l l ne t s , 34% fo r t r o l l ing, 6% fo r hand1 ines , and 4% fo r other methods (Trent e t a l . 1983). Purse seining has recently been per- mitted i n Federal waters (GWSAFMC 1982).

Major changes i n the locations and the in tens i ty of f ishing in Flori- da have taken place since about 1960. The ea s t coast center of production has gradually sh i f t ed northward from Dade County toward Volusia County and a spring f ishery of major importance has developed off Palm Beach County near Jup i te r and Juno (Williams and Godcharles 1983). As a r e su l t of higher dockside pr ices , f ishing e f f o r t s increased sharply: from 1969 t o 1977 there was a three-fold increase i n hook-and-1 i ne vessels , 100 t o 300, and g i l l net vessels , 12 t o 33 (GM&SAFMC 1982). By 1983, the number of gi 11 net vessels had r isen t o about 80, and the larger net boat f l e e t had moved onto f i sh ing grounds t ha t had formerly been used pr inci- pal ly by t r o l l e r s and handliners. In addit ion, the net f l e e t has increased i t s eff ic iency through the use of spot ter a i rplanes , monof i lament net construction, 1 arger and deeper nets mechanical ly re t r ieved by power rol l e r s , and e lec t ron ic equipment.

Spanish mackerel. This species i s of major commercial im~ortance in south ~ i o r i d a (Klima 1959; Powell 1975). The main f i sh ing areas a re the Florida Keys and the Atlant ic coast between Palm Beach and Cape Canaveral.

Small numbers a re caught as an inci- dental o r supplemental commercial species off the coasts of Alabama, Mississippi, Louisiana, North Carolina, and, t o a smaller extent , Georgia and South Carol ina.

Spanish mackerel are primarily captured w i t h g i l l nets deployed from small boats 20 t o 22 f t long, and with power r o l l e r s used on large boats 30 t o 60 f t long. Since 1976, commercial production on the gulf coast has f luc- tuated between 1 .5 and 3.5 mill ion Ib, and production on the Atlant ic coast has fluctuated between 3.4 and 11 mil l ion lb (Table 6). The causes of f luctuat ion in catch cannot be ident i - f i ed because there are no catch-per- u n i t -o f -e f for t data.

Spanish mackerel a l so a re an important species f o r the pr ivate boat and char ter boat spor t f ishery a1 ong the gulf and south Atlant ic coasts. Most anglers f i s h from pr ivate boats, although good catches a re made from char ter boats, f i shing p i e r s , and beach f ishing (Deuel 1973).

The limited spor t s t a t i s t i c s suggest t ha t 1979 commercial landings on the Atlant ic coast were double the spor t catch. On the gulf coast , however , the spor t catch was probably 50% higher than the commercial catch (GWSAFMC 1982).

Fisheries Management Plan. The Fishery Management Plan (FMP) fo r coastal pelagic f i sh species , includ- ing mackerel, was implemented in March 1983 by the Gulf of Mexico and South A t 1 an t i c Fishery Management Counci 1 s (GM&SAFMC). Quotas based on theore t i - cal y ie lds par t i t ioned according t o h i s to r ica l landings were es tabl ished f o r the commercial hook-and-1 ine (3,877,200 l b ) , net (5,122,800 l b ) , and spor t (28,000,000 1b) f i she r i e s , and were in e f f ec t from 1 July 1982 t o 30 June 1983. In May 1983, the commercial hook-and- 1 ine f ishery was o f f i c i a l l y closed when t h a t quota was a t ta ined. This ea r ly closure was

Table 6. F l o r i d a commercial landings (thousands o f pounds) o f k i n g and Spanish mackerel i n the Gu l f o f Mexico and along the A t l a n t i c coast, 1960-83 (NMFS Annual Landings).

G u l f o f Mexico King Spanish

Year mac ke r e 1 mackerel

A t l a n t i c Kina Spanish

mackerel mackerel

a t t r i b u t e d t o increased catches i n Nor th Caro l i na (0.7 m i l l i o n I b ) and Louis iana (1.2 m i l l i o n Ib ) . These new developments i n con junct ion w i t h decreasing catches i n south F l o r i d a are c u r r e n t l y under review by the GM&SAFMC. Recent s tud ies suggest t h a t increased catches are r e l a t e d t o s t rong year c lasses, t h a t more than one migra tory group e x i s t s , and t h a t a maximum sustainable y i e l d o f 37.7 m i 11 i o n 1 b may have been overestimated (Powers and E ldr idge, unpubl. MS. a, b; W i 11 iams and Godcharl es 1983).

A maximum sustainable y i e l d o f 27 m i l l i o n 1b f o r Spanish mackerel was

es tab l ished by the f i s h e r i e s management p l a n (GM&SAFMC 1982). Sport and commercial catch s t a t i s t i c s from the Nat ional Marine F i she r ies Serv ice reveal t h a t on l y about h a l f o f t he maximum sustainable y i e l d was landed i n 1979.

ECO LOG I CA L ROLE

Food Habi ts

King and Spanish mackerel juve- n i l e s and adu l ts are p r imar i l y pe lag i c carnivores. Analys is o f t he stomach

contents o f 84 j u v e n i l e k i n g mackerel (103-309 mm FL) from Cape Canaveral, 130 Spanish mackerel (117-432 mm FL) from Cape Canaveral, and 214 Spanish mackerel from Gal veston Bay revealed t h a t j u v e n i l e s o f bo th species were p r i n c i p a l l y p isc ivorous , b u t k i n g mackerel showed a g rea te r preference f o r inver tebra tes . Anchovies (Anchoa spp. ) , menhaden (Brevoor t ia spp.), and A t l a n t i c th read h e r r i na - (Opi sthonema ogl i num) were t h e dominant forage o f t h e mackerels. Much l ess common were mugi 1 i ds , sc iaenids, carangids, and e l e o t r i d s . Squid was the major i n v e r t e b r a t e prey f o r bo th species. Juven i les f e d h e a v i l y on anchovies, as shown by t h e i r frequency o f occurrence (19%-39%) and volume (30%-54%) i n mackerel stomachs. 'The body shape o f anchovies appears t o make them h i g h l y s u i t a b l e prey f o r juven i l e mackerel (Naughton and Sal oman 1981).

King mackerel feed most ly on school ing f i s h , secondar i l y on c rus ta- ceans, and m in ima l l y on mollusks. The dominant p rey by number (59%) were c l upeids ( A t l a n t i c th read he r r i ng ) and scaled sardines, Haren u l a jagua?a. The minor f i s h p r e y d r e specles o f Carangidae, Lut janidae, Pomadasyi- dae (Haemul idae) , Sparidae, and T r i g1 idae. Inver tebra tes , p a r t i c u - l a r l y squ id and shrimps, made up 33% o f t h e d i e t (Beaumariage 1973).

I n south F l o r i d a , t he k ina mack- e r e l fed p r i m a r i l y on t h e bai lyhoo, ~emiramphus b r a s i l i e n s i s , f o l l owed by l u t j a n i d s ( f i v e species), c lupe ids , scombrids, mugi l i d s - (two spec ies j , and serranids (Saloman and Naughton 1983a). The i nve r teb ra tes eaten were most ly penaeid shrimp and some squ id and nematodes. I n eas t c e n t r a l F lo r i da , c lupe ids ( p r i n c i p a l l y t h e Spanish sardine, (Sardi ne l 1 a a u r i t a ) were the dominant f i s h prey. -Other prey were anchovies, m u l l e t , f l y i n g f i s h , drum, and jacks. Squid was t h e major i nve r teb ra te food; o thers were nematodes, penaeid shrimp, and i s o - pods. Most f i s h eaten by a d u l t k i n g

mackerel were about t h e same s ize : 100 t o 150 mm FL (Saloman and Naughton 1983a).

The food o f a d u l t k i n g and Spanish mackerel i s s i m i l a r . Kl ima (1959) exami ned 190 Spanish mackerel stomachs and repo r ted t h a t 76% con- t a i n e d h e r r i n g l i ke f i shes , p r i n c i p a l l y t he scaled sard ine and A t l a n t i c th read her r ing . Shrimp (Penaeus spp. ), mu1 l e t (Mugi l spp.),eedlefish (S t rongy lura spp.), and anchovies were l e s s abundant. I n Texas, M i l es and Simmons (1951) examined 2,274 Spanish mackerel stomachs con ta in ing food, and found t h a t 30% contained menhaden. Kemp (1950), a l so working i n Texas, repor ted t h e contents o f 611 Spanish mackerel stomachs: 13% contained shrimp; 5%, squid; 9%, r i b b o n f i s h ; I%, menhaden; I%, o the r species; and t h e remainder, u n i d e n t i f i a b l e . The round scad (Decapterus punctatus) was a l s o 1 i s t e d as a food o f t h e Spanish mack- e r e l (Anderson and Gehringer 1957).

Stomachs o f 6,933 Spanish mack- e r e l (64% were empty) were examined from t h e no r the rn G u l f o f Mexico, from east c e n t r a l F l o r i d a , and from the Carol inas. I n vo l ume, anchovies made up 96% o f t h e food i n Texas and 99% i n eas t c e n t r a l F l o r i d a (Saloman and Naughton 1983b). Anchovies a1 so made up 94% o f t h e d i e t by number i n Louis iana and 98% i n Nor th and South Carol ina. The predominance o f anchovies, he r r i ng , and small jacks i n t he Spanish mackerel d i e t demonstrates major p reda t i on on small , school i n g pe lag i c f i shes .

Predators

Larvae and j u v e n i l e s o f k i n g and, p r i n c i p a l ly, Spanish mackerel have been i d e n t i f i e d as prey f o r t h e l i t t l e tunny, Euth nnus a l l e t t e r a t u s , and do lph in , Coryphaena h i ppuru; (Carl son 1952: Klawe r a a o v ~ c h 969: Rose and^ ' ~ a s s l e r 1974). - k e l a t i v e l y , l a rge k i n g and Spanish mackerel a re eaten by

pelagic sharks, l i t t l e tunny, and dolphins. The bottlenose dolphin ( ~ u r s i o ~ s truncatus) in te r fe res w i t h commercial f ishing bv ~ i r a t i n a k i n a mackerel hooked on t i 1 l'ing 1 i t e s ani in nets (Cato and Prochaska 1976). Sharks sometimes in te r fe re w i t h g i l 1 net s e t s by eating mackerel caught i n the mesh. The most common shark spe- c ies a re the t i g e r shark, Galeocerdo cuvieri ; bull shark, Carcharhin~ leucase dusky shark, C. obsc

-9 - smooth hammerhead, Sphyyna zva shortf in mako, Isurus ox

- , ,aena; - y r i nchus ;

lemon shark, Negaprion brevi ros t r i s ; and porbeagl e , Lamna nasus (Bigelow and Schroeder 1948: CTark and von Schmidt 1965; GMASAFMC 1982).

ENVIRONMENTAL REQUIREMENTS

Temperature

Temperature and sa l i n i t y a re be- lieved t o be the most important fac- t o r s governing the dis t r ibut ion of the two mackerels. Their northern range extends only t o the 20 O C isotherm within the 18 m depth contour (Munro 1943; Berrien and Finan 1977a). Their northern range l imi t i s in the vici- ni ty of Block Island, Rhode Island (Beaumari age 1970). Duri ng years of warm water temperatures, Spanish mack-

erel have been reported as f a r north as North Bay, Massachusetts (Arnold 1951). According t o Earll (1883), water temperatures of 21 t o 27 O C are preferred by the Spanish mackerel ; rarely a re they observed i n waters cooler than 18 O C .

The a r r iva l of king mackerel off west central Florida i n the spring depends on changes in water tempera- tu re and on the preceding winter ' s a i r temperature (Wi 11 iams and Taylor 1980). Furthermore, sport catch data from northwest Florida indicate t h a t catch-per-hour i s usually higher fol lowing warm winters and lower following cold winters (Fable e t a l . 1981).

Sa l in i ty

All l i f e stages of king and Spanish mackerel usually inhabit waters within s a l i n i t i e s of 32 t o 36 ppt. Spanish mackerel usually avoid freshwater or low s a l i n i t i e s near the mouths of r ivers (Earll 1883). Ex- ceptions were reported by Tagatz and Dudley (1961), who collected young Spanish mackerel in a s a l i n i t y of 4.7 ppt in the Neuse River, North Carol i na. Other investigators have reported juveni 1 es i n low s a l i n i t i e s (17.8 ppt, Springer and Woodburn 1960; 0.2 ppt, Kelley 1965).

LITERATURE CITED

Anderson, W. W. , and J. W. Gehringer. sharks). Sears Foundation f o r Marine 1957. Physical oceanography, Research, New Haven, Conn. 546 pp. b i o l o g i c a l , and chemical data, South A t l a n t i c coast o f t h e Uni ted States, Bigelow, H.B., and W.C. Schroeder. Theodore N. G i l l c r u i s e 3. U.S. 1953. Fishes o f the Gu l f o f Maine. F i sh Wi ld l . Serv. Spec. Sci. Rep. U.S. F i sh Wi ld l . Serv. Fish. B u l l . Fish. 210. 208 pp. 53. 577 pp.

Arnold, E.L., J r . 1951. Northward Burns, K.M. 1981. Seasonal and area l d ispersa l o f warm-water f i shes i n d i s t r i b u t i o n o f scombrid la rvae i n southern New England dur ing the the v i c i n i t y o f Palm Beach, F lo r i da . s umme r o f 1949. Copei a M.A. Thesis. U n i v e r s i t y o f South 1951(1): 87-88. F lo r i da , Tampa. 66 pp.

Beaumariage, D. S. 1970. Current sta- t us of b i o l o g i c a l i nves t i ga t i ons o f F l o r i d a ' s mackerel f i s h e r i e s . Proc. Gulf Caribb. Fish. I n s t . , 22nd Annual Meeting, 1969: 79-86.

Beaumariage, D. S. 1973. Age, growth, and reproduct ion o f k i n s mackerel . comber om or us caval l a , i n ~ l o r i d a : Fla. Mar. Res. Publ. 1. 45 pp.

Berr ien, P., and D. Finan. 1977a. B io log i ca l and f i s h e r i e s data on k ing mackerel , Scomberomorus caval 1 a (Cuvier). U.S. Na t l . Mar.- ~ e r v . , -sandy Hook Lab. , Highlands , N.J. Tech. Ser. Rep. 8. 40 pp.

Ber r ien P . , and D. Finan. 1977b. B io log i ca l and f i s h e r i e s data on Spanish mackerel , Scomberomorus maculatus (M i t ch i 11 ). U. S. Na t l . Mar. Fish. Serv., Sandy Hook Lab., Highlands, N. J. Tech. Ser. Rep. 9. 52 PP.

Bigelow, H.B., and W.C. Schroeder. 1948. Fishes o f t he western Nor th A t l a n t i c ( l ance le ts , cyclostomes,

Br iggs, J.C. 1958. A l i s t of F l o r i d a f i shes and t h e i r d i s t r i b u t i o n . B u l l . Fla. S ta te Mus. B i o l . Sci. 2(8). 318 pp.

Carlson, C. B. 1952. Exp lora tory f i s h i n g f o r t he l i t t i e tuna (Euthynnus a l l e t t e r a t u s ) o f f the A t l a n t i c coast o f t he Uni ted States. Proc. Gu l f Caribb. Fish. I n s t . , 4 t h Annu. Session 1951:89-94.

Cato, J.C. , and F. J. Prochaska. 1976. Porpoise a t t a c k i n g hooked f i s h i r k and i n j u r e F l o r i d a fisherman. Nat l . Fisherman 56(9): 3b, 16b.

Clark, E., and K. von Schmidt. 1965. Sharks o f t he c e n t r a l Gu l f coast o f F lo r i da . B u l l . Mar. Sci. 15:13-83.

Col l e t t e , 0.0. , J. L. Russo, and L.A. Zavala-Cami n. 1978. Scomberomorus b r a s i l i e n s i s , a new species o f Spanish mackerel from t h e western A t l a n t i c . U.S. Nat l . Mar. Fish. Serv. Fish. B u l l . 76(1): 273-280.

Deuel, D.G. 1973. The 1970 s a l t - water ang l i ng survey. U.S. Nat l .

Mar. F ish. Serv. Curr. F ish. S ta t . 6200. 54 pp.

Doi , T., and D. Mendizabal. 1979. Evaluacion ~ r e l i m i n a r de l a ~ o b l a - c i o n de ' s i e r r a , comber om or us rnaculatus ( M i t c h i l l ) , f r e n t e a l a s costas de Veracruz. Proceedings: col loquium on the Spanish and kTng mackerel resources o f the Gu l f o f Mexico. Gu l f States Mar. F ish. Comm. No. 4:43-55.

Dragovich, A. 1969. Review o f s tud ies o f tuna food i n the A t l a n t i c Ocean. U.S. F i sh W i l d l . Serv. Spec. Sci . Rep. F ish. 593. 2 1 pp.

Dwine l l , S.E. , and C. R. Futch. 1973. Spanish and k ing mackerel l a r vae and juven i 1 es i n t h e nor theastern G u l f o f Mexico, June through October, 1969. Fla. Dep. Nat. Resour. Mar. Res. Lab. Leaf l . Ser. 4(24). 14 pp.

E a r l l , R.E. 1883. The Spanish mack- e r e l , Cybium maculatum ( M i t ch . ), Ag. ; i t s na tu ra l h i s t o r y and a r t i f i - c i a l propagation, w i t h an account o f i t s o r i g i n and development o f t he f i s h e r y . Rep. U.S. Comm. F i sh Fish. (1880) p t . 8: 395-424.

Fable, W.A., J r . , H.A. Brusher, L. Trent , and J. Finnegan, J r . 1981. Possib le temperature e f f e c t s on c h a r t e r boat catches o f k i n g mackerel and o the r coasta l pe lag i c species i n northwest F lo r i da . U. S. Na t l . Mar. Fish. Serv. Mar. Fish. Rev. 43(8): 21-26.

Finucane, J.H., and L.A. C o l l i n s . R e ~ r o d u c t i o n o f S ~ a n i sh mackerel . comber om or us macul a tus , from the G u l f o f Mexico and South A t l a n t i c Ocean o f t h e Un i ted States. U.S. Nat l . Mar. Fish. Serv., Panama C i t y , F la. 23 pp. (unpubl. MS.)

Finucane, J.H., L.A. C o l l i n s , H.A. Brusher, and C.H. Saloman. Repro- duc t i ve b io logy o f k i n g mackerel ,

Scomberomorus caval 1 a, from the southeastern Un i ted States. U. S. Nat l . Mar. Fish. Serv.. panama ~ i t v . - - Fla. 28 pp. (unpubl. MS.)

Goode, G. B. 1884. The f i s h e r i e s and f i s h e r i e s i n d u s t r i e s o f the Un i ted States. Sect ion 1. Natural h i s t o r y o f usefu l aquat ic animals. U.S. Government P r i n t i n g O f f i c e , Washington, D.C.

Gu l f o f Mexico and South A t l a n t i c F ishery Management Counci 1 s (GM&SAFMC). 1982. F ishery management p l an/ f i nal environmental impact statement/regul a t o r y impact review/ d r a f t r egu la t i ons f o r the coasta l pe lag i c resources (mackerels) i n the Gu l f o f Mexico and South A t l a n t i c region. Tampa, F la . (GMFMC), and Charleston, S. C. (SAFMC).

Hi ldebrand, S. F. , and L. E. Cable. 1938. Fur ther notes on the development and l i f e h i s t o r y o f some t e l e o s t s a t Beaufor t , N.C. B u l l . U.S. Bur. Fish. 48(24):505-642.

Hoese, H. D. 1965. Spawning o f marine f i s h e s i n t he Po r t Aransas, Texas, area as determined by the d i s t r i - b u t i o n o f young and larvae. Ph.D Diss. Univ. Texas, Aust in. 144 pp.

Houde, E.D., J.C. Leak, C.E.Dowd, S.A. Berkeley, and W.A. Richards. 1979. Ichthyoplankton abundance and d i ve r - s i t y i n t he eastern G u l f o f Mexico. F i n a l r e p o r t t o t he Bureau o f Land Management, Contract No. AA550-CT7- 28. 546 pp.

Johnson, A.G., W.A. Fable, J r . , M. L. Wi l l iams, and L.E. Barger. 1983. Age, growth, and m o r t a l i t y o f k i ng mackerel , Scornberomorus caval 1 a, from the Southeastern Uni-ted-. U.S. Nat l . Mar. Fish. Serv. Fish. B u l l . 81(1): 97-106.

Ke l ley , J.K. 1965. A taxonomic survey o f t he f i shes o f D e l t a Nat iona l W i l d l i f e Refuge w i t h emphasis on

d i s t r i b u t i o n and abundance. M. S. Munro, I. S. R. 1943. Revis ions o f Thesis. Lou is iana S ta te U n i v e r s i t y , A u s t r a l i a n species o f Scomberomorus. Baton Rouge. 126 pp. Mem. Queens1 . Mus. 12(21): 65-69.

Kemp, R. J. 1950. Report on stomach ana l ys i s from June 1, 1949 through August 31, 1949. Pages 101-127 i n Texas Game and F i s h Commiss i f i Laboratory Annual Report f o r f i s c a l yea r 1948- 1949.

Klawe, W. L. 1961. Young scombroids from t h e waters between Cape Hat te ras and Bahama I s l ands . B u l l . Mar. Sc i . G u l f Car ib . ll(1): 150-157.

Kl ima, E.F. 1959. Aspects o f t h e b i o l o g y and f i s h e r y f o r Spanish mackerel , ~comberombrus macul a tus ( M i t c h i 11 ) , o f southern F l o r i d a . la. ~ o a r d - Conserv. Mar. Res. Lab. Tech. Ser. 27. 39 pp.

Mago-Leccia, F. 1958. The comparat ive os teo losy o f t h e scombroid f i s h e s o f -- t h e genus Scomberomorus from F l o r i d a . B u l l . Mar. Sci. G u l f Car ib .

May, B. Genet ic v a r i a t i o n i n t h e - . k i n g mackerel (Scomberomorus caval 1 a). Cornel 1 U n i v e r s i t y , I t h a c a , N.Y. F i n a l r e p o r t on Con- t r a c t - No. C1434 t o t h e F l o r i d a Department o f Na tu ra l Resouces. (unpubl . MS. )

McEachran, J.D. , J.H. Finucane, and L.S. H a l l . 1980. D i s t r i b u t i o n , seasonal i t y and abundance o f k i n g and Spanish mackerel l a r v a e i n t h e nor thwestern G u l f o f Mexico (Pisces: Scombridae). Nor theas t G u l f Sc i . 4(1): 1-16.

Mendoza, N. A. 1968. Consideraciones sobre l a b i o l o g i a pesquera de l a s i e r r a Scomberomorus maculatus ( M i t c h i l l ) , en e l estado de

Naughton, S. P. , and C. H. Saloman. 1981. Stomach con ten ts o f j u v e n i l e s o f k i n g mackerel (Scomberomorus c a v a l l a and Spanish mackerel (S. - maculatus). Nor theas t G u l f Sc i . 5 m.

Powel 1, D. 1975. Age, growth and rep roduc t i on i n F l o r i d a stocks o f Spanish mackerel , Scomberomorus maculatus. F la . Mar. Res. Publ. 5. 2 1 PP.

Powers, J.E., and P. E ld r idge . A p r e l im ina ry assessment o f k i n g mackerel resources o f t h e southeast Un i t ed States. U.S. N a t l . Mar. F ish. Ser., Miami, F la . (unpubl. MS. a)

Powers, J.E., and P. E ld r i dge . Assessment o f G u l f o f Mexico and South A t l a n t i c k i n g mackerel. U. S. N a t l . Mar. F ish. Serv., Miami, F la. (unpubl. MS. b)

Powles. H. Abundance and d i s t r i b u t i o n o f k i n g mackerel (Scomberomorus c a v a l l a ) and S ~ a n i s h mackerel (S. macul a t i s ) 1 arvae o f f t h e southeaJt Un i t ed States. South Carol i na Marine Resources Center. Char1 eston. 16 pp. (unpubl. MS.)

Richardson, S. L. , and J. D. McEachran. 1981. I d e n t i f i c a t i o n o f smal l ( l e s s than 3 mm) l a r v a e o f k i n g and Spanish mackerel , Scomberomorus caval l a and S. maculatus. Nor theas t G u l f Sc i . 5 ( i ) : 75-79.

Rose, C. D. , and W. W. Hassler . 1974. Food h a b i t s and sex r a t i o s o f do lph in , Coryphaena c a ~ t u r e d i n t h e western ocean o f f Hat te ras , No r th Carol i na. Trans. Am. F ish. Soc. 103:94-100.

Veracruz. B ios l ( 2 ) : 11-22. Ryder, J.A. 1882. Development o f t h e

Mi les , D.W. , and E.G. Simmons. 1951. The menhaden f i s h e r y . Tex. Game F i s h Oyster Comm. B u l l . 30. 28 pp.

- spani sh mackerel ( C b i um' maculatum). B u l l . U. S. %h Comm.

Saloman, C.H., and S.P. Naughton. 1983a. Food o f k ina mackerel. Scomberomorus caval l a ," from the Southeastern Uni ted States i nc lud ina the Gu l f o f Mexico. U.S. l3ep: Commer. NOAA Tech. Memo. NMFS-SEFC- 126. 2 5 p p .

Saloman, C. H. , and S. P. Naughton. 1983b. Food o f Spanish mackerel. Scomberomorus macul atus, from the G u l f o f Mexico and southeastern seaboard o f the Un i ted States. U.S. Dep. Commer. NOAA Tech. Memo. NMFS- SEFC-128. 22 pp.

Skow, L.C. , and M.E. Chittenden, J r . 1981. D i f fe rence i n hemoglobin phenotypes among Spanish mackerel, Scomberomorus maculatus. Northeast G u l f Sci. 5(1):67-7

Smith, H.M. 1907. The f i shes o f North Carol ina. N.C. Geol. Econ. Surv. 2. 433 pp.

Springer, V.G. , and K. D. Woodburn. 1960. An eco log ica l study o f the f i shes o f t h e Tampa Bay area. Fla. Board Conserv. Mar. Lab. Pro f . Pap. Ser. 1. 104 pp.

Sutherland, D. F., and W.A. Fable, J r . 1980. Results o f a k ing mackerel (Scomberomorus caval l a ) and A t l a n t i c S ~ a n i sh m a c k e ~ ( ~ c o m b e r o m o r u s maculatus) m ig ra t i on study, 1975- 1979. U.S. Dep. Commer. NOAA Tech. Memo. NMFS-SEFC-12. 18 pp

Tagatz, M. E. , and D. L. Dud1 ey. 1961. Seasonal occurrence o f marine f i shes i n f o u r shore hab i ta t s near Beau- f o r t , N.C. 1957-60. U.S. F i s h Wi ld l . Serv. Spec. Sc i . Rep. Fish. 390. 19 pp.

Trent , L., P. E ldr idge, and E. Anthony. 1983. Commercial and rec- r e a t i o n a l f i s h e r i e s s t a t i s t i c s f o r k ing mackerel i n the southeast Uni ted States. Pages 1-2 i n Abstracts o f papers presented a t t& k ing mackerel symposi um November 4- 5, 1983, Orlando, Fla. F la . Dep. Nat. Resour. and U.S. Nat l . Mar. F ish. Serv. (Abstr . )

W i 11 iams, R. 0. , and M. F. Godcharl es. 1983. King mackerel tagging and stock assessment. Report f o r 1981-1982 t o U.S. Na t l . Mar. Fish. Serv. (Unpubl.) F la. Dep. o f Nat. Resour. PL 88-309: P r o j e c t No. 341-a. 30 pp. (16 pp. t e x t . )

W i l l iams, R.O. , and D. F. Sutherland. 1979. King mackerel migra t ions . Pro- ceedings: co l loquium on the Spanish and k ing mackerel resources o f the G u l f o f Mexico. G u l f States Mar. F ish. Comm. No. 4:57. (Abst r . )

Wi l l iams, R . O . , and R.G. Taylor . 1978. King mackerel tagg ing and stock assessment study (unp l . ) Com- p l e t i o n r e p o r t t o Nat l . Mar. Fish. Serv. F la. Dep. Nat. Resour. PL 88-309: P ro jec t No. 2-254-r. 70 pp. (19 pp. t e x t . )

W i l l iams, R.O. , and R.G. Taylor . 1980. The e f f e c t o f water temperature and w in te r a i r temperature on spr ingt ime migra t ions o f k ing mackerel i n the v i c i n i t y o f Tampa Bay, Fla. F la. Sci. 43 (suppl.):26. (Abstr . )

Wollam, M.B. 1970. Desc r ip t i on and d i s t r i b u t i o n o f l a rvae and e a r l y juven i 1 es of k i ng mackerel , Scomber- omorus c a v a l l a (Cuvier), and Spanish mackerel, S. maculatus ( M i t c h i l l ) ; (Pisces: ~combridae); i n the western North A t l a n t i c . F la. Dep. Nat. Resour. Mar. Res. Lab. Tech. Serv. 61. 35 pp.

60172 -101

15. Supplementaty Nates

*U.S. Army Corps o f Engineers Report No. TR EL-82-4 16. Abstrmsl (Umit: 200 words) This Species P r o f i l e on k i ng and Spanish mackerel summarizes the tax-

onomy, morphology, d i s t r i b u t i o n , 1 i f e h i s t o r y , f i s h e r y descr ip t ions , eco log ica l r o l e , and environmental requirements o f these coasta l pe lag ic f i s h t o a s s i s t environmental impact assessment. King and Spanish mackerel support major commercial and spor t f i she r i es i n south F lo r ida . I n 1974-83, Gu l f o f Mexico and A l t a n t i c commerical landings o f k ing mackerel dec l ined from 10.4 t o 4.3 m i l l i o n I b. ; Spanish mackerel have f luc tua ted between 4.9-17.4 m i l 1 i o n 1 b. Both i n h a b i t coasta l waters, bu t Spanish mackerel are genera l l y found c loser t o beaches and i n ou te r estuar ine waters. Both species feed p r i n c i p a l l y on estuarine-dependent species. They are h i g h l y migratory , e x h i b i t i n g seasonal migrat ions t o w in te r feeding grounds o f f south F l o r i da and summer spawning/feeding grounds i n the nor thern Gu l f o f Mexico and o f f t h e A t l a n t i c coast o f the Southeastern U.S. Spawning occurs from March/Apri l through September/October between the middle and Outer Cont inental Shel f (35-183 m i ) f o r k ing mackerel and t he inner s h e l f (12-34 m i ) f o r Spanish mackerel. King mackerel reach sexual ma tu r i t y i n ' t h e i r 3 rd and 4 t h years and Spanish, between t h e i r 2nd and 3rd. Female k ing mackerel l i v e longer and grow l a rge r and f a s t e r than males. Spanish mackerel l i v e t o 8 years; females a lso grow f a s t e r than males. King and Spanish mackerel feed p r i n c i p a l l y on school ing f ishes. Larvae and juven i les o f both species are prey t o l i t t l e tunny and dolph in ; adu l t s are prey f o r sharks and bot t lenose dolphin. Temperature and s a l i n i t y are impor tant f ac to r s r egu la t i ng mackerel d i s t r i b u t i o n .

17. Document Analysus a. Deacripton

REPORT DOCUMENTATION 1. REPORT NO. 2. PAGE B i o l o a i c a l Report 82(11.58)* 1 .

I - I

4. Title and Subtitle Species P r o f i 1 es: L i f e H i s t o r y and Environmental Requirements o f Coastal Fishes and Inver tebrates (South F lor ida)- - King Mackerel and Spanish Mackerel.

7. Author(s)

Mark F. Godcharles and Michael D. Murphy 9. perform in^ Organization Name end Address

F l o r i d a Department o f Natural Resources D i v i s i o n o f Marine Resources, Bureau o f Marine Research 100 Eighth Avenue, SE St. Petersburg, FL 33701 12. Sponsorinn Organlzatlon Name and Address

National Coastal Ecosystems Team U. S. corps of ~~~i neers F ish and W i l d l i f e Service Waterways Experiment S ta t i on U.S. Department o f t he I n t e r i o r P. 0. Box 631 Washington, DC 20240 Vicksburg, MS 39180

Growth Feeding hab i t s

3. Reclplent's Accass~on No.

L RepoR Date

J~~~ 1986

8. Performin. Oraan~zatlon Re@. NO.

10. PmiectlTaskIWod~ Unit No.

11. bntract(C) or Crant(C) NO.

(c)

(C)

13. TYP* of Report 6 Penad Covemd

14.

1 b. Identlfier8lOoan.Ended Terms

King mackerel Scomberomorus cava l l a Spani sh mac k e r r

I c. COSATI !GeldlGroup

Temperature Sal i n i t y

L i f e cyc les Marine f i shes Animal migrat ions

Scomberomorus maculatus kco l og i ca l r o l e s Environmental requirements

18. Avallmbllity Statement 19. Sccurkty Class (This ReporO 21. No. of Pages

Unclass i f ied Re1 ease Unl i m i ted m. Security C I ~ S S (This Page)

Unclass i f ied (See ANSI-Z39.lB) OPTIONAL FORM 272 (4-77)

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