REFERENIP': CO?Y Do Not Remove fr9m the Library

U. S. Fish and Wildlife Service National Wetlands Research Center

FWSIOBS-82/11 .27 700 Cajun Dome Bo~~levard September 1984

TR EL-82-4 La fayette, Lou isiano 70506

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

WHITE SHRIMP

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of ~ n ~ g e r s

FWSjOBS-82/11.27 TR EL-82-4 September 1984

Species Pro f i 1 es : L i f e H i s t o r i e s and Environmental Requirements o f Coastal Fishes and Inver tebra tes (South At1 a n t i c )

WHITE SHRIMP

Robert J. Muncy M iss i ss ipp i Cooperative F ish and Wild1 i f e Research U n i t

P.O. Drawer BX M iss i ss ipp i S ta te U n i v e r s i t y M i s s i s s i p p i State, MS 39762

P ro jec t Manager La r r y Shanks

P ro jec t O f f i c e r John Parsons

Nat ional Coastal Ecosystems Team U.S. F i s h 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

U.S. Army Corps o f Engineers Waterways Experiment S t a t i o n

Vicksburg, MS 39180

and

Nat iona l Coastal Ecosystems Team D i v i s i o n of B i o l o g i c a l Services

Research and Devel oprnent F i sh and W i l d l i f e Serv ice

U.S. Department o f t he I n t e r i o r Was h i ng ton, DC 20240

Th i s s e r i e s should be re fe renced as f o l 1 ows :

U.S. F i s h and W i l d l i f e Serv ice. 1983-19-. Species p r o f i l e s : l i f e h i s t o r i e s and env i ronmenta l requ i rements o f coas ta l f i s h e s and i nve r t eb ra tes . U.S. F i s h W i l d l . Serv. FWS/OBS-82/11. U.S. Army Corps o f Engineers, TR EL-82-4.

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

Muncy, R. J. 1984. Species p r o f i l e s : 1 i f e h i s t o r i e s and env i ronmenta l r e q u i r e - ments 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 (South A t l a n t i c ) - - w h i t e shrimp. U.S. F i sh W i l d l . Serv. FWS/OBS-82/11.27. U.S. Army Corps o f Engineers, TR EL-82-4. 19 pp.

PREFACE

This species p r o f i l e i s one o f a ser ies on coastal aquat ic organisms, p r i n c i p a l l y f ish, o f sport , commercial , o r eco log ica l importance. The p r o f i l e s a re designed to provide coastal 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 of the b i 01 og i c a l c h a r a c t e r i s t i c s and environmental requ i re- ments o f the species and t o descr ibe how populat ions o f the species may be expected to react t o e n v i r o m e n t a l changes caused by coastal development. Each p r o f i l e has sect ions on taxonany, 1 i f e h i s t o r y , eco log ica l ro le , envi ronmental requirements, and economic importance, i f appl i cab l e. A t h ree - r i ng b inde r i s used f o r t h i s ser ies so t h a t new p r o f i l e s can be added as they are prepared. This p r o j e c t i s j o i n t l y planned and f inanced by the U.S. Army Corps o f Engineers and the U.S. Fish 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 o f t he f o l l o w i n g addresses.

In format ion Transfer Special i s t Nat ional Coastal Ecosys tems Team U.S. Fish and Wild1 i f e Service NASA-Sl i d e l 1 Computer Compl ex 1010 Gause Boul evard Sl i d e l 1 , LA 70458

U.S. Army Engineer Waterways Experiment S ta t i on At tent ion : WESER-C Post O f f i c e Box 631 Vicksburg, MS 39180

CONTENTS

Page

PREFACE ........................................................... iii ...................................................... CONVERSION TA9LE v ACKNOWLEDGMENTS ..................................................... v i

........................................... NOMENCLATURE/TAXONOMY/RANGE 1 ............................................ MORPHOLOGY/IDENTIFICATION 1 ....................................... REASONS FOR INCLUSION IN SERIES 3 LIFE HISTORY .......................................................... 4 ........................................................... Spawning 4

Eggs .............................................................. 4 Larvae ............................................................. 4 ........................................... Post la rvae and Juven i les 5 ............................................................. Adu l t s 6 M i g r a t i o n .......................................................... 6

GROWTH ................................................................ 7 .......................................................... Mor ta l i t y 8 ............................................. D i seases and Parasi t e s 8 ........................................................... THE FISHERY 9 ....................................................... ECOLOGICAL ROLE 11 ............................................ ENVIRONMENTAL REQUIREMENTS 12 Temperature ........................................................ 12 ........................................................... Sal i n i ty 12 .................................. Temperature-Sal i n i t y I n t e r a c t i o n s 13 .......................................................... Subst ra te 13 ................................. Other Environmental Considerat ions 13

LITERATURE CITED ...................................................... 15

CONVERSION TABLE

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

m i l 1 imeters (nun) cent imeters (an) meters (m) k i 1 ometers ( km)

2 square meters (m ) square k i 1 ometers ( km2) hectares (ha)

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

m i l 1 igrams (mg) grams ( g ) k i 1 ograms ( kg) m e t r i c tons ( t ) m e t r i c tons k i 1 ocal o r i e s ( kcal )

Cel s i u s degrees

To Obta in

inches inches fee t m i l es

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

0.00003527 ounces 0.03527 ounces 2.205 pounds

2205.0 pounds 1.102 s h o r t tons 3.968 B r i t i s h thermal u n i t s

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

gal 1 ons cub ic f e e t acre- fee t

1.8(c0) + 32 Fahrenhei t degrees

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

inches 25.40 inches 2.54 fee t ( f t ) 0.3048 f a thoms 1.829 m i l e s ( m i ) 1.609 n a u t i c a l m i les ( m i ) 1.852

ga l l ons ( g a l ) cub ic f e e t ( f t 3 ) acre- f e e t

ounces (oz ) 28.35 pounds ( I b ) 0.4536 s h o r t tons ( t o n ) 0.9072 B r i t i s h thennal u n i t s ( B t u ) 0.2520

m i l 1 imeters cent imeters meters meters k i l ometers k i 1 ometers

square meters hectares square k i 1 ometers

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

grams k i 1 ograms m e t r i c tons k i 1 ocal o r i es

Fahrenhei t degrees 0.5556(F0 - 32) Cel s i u s degrees

ACKNOWLEDGMENTS

For t h e i r he1 p f u l rev iews and suggest ions, I thank Susan Shipman, Coasta l F i s h e r i e s Sect ion, Georgia Department o f Na tu ra l Resources; Erlwin A. Joyce, Jr., Di v i s i o n o f Mar ine Resources, F l o r i d a Department o f Na tu ra l Resources ; and David Whitaker, D i v i s i o n o f Mar ine Resources, South Ca ro l i na W i l d l i f e and l l a r i n e Resources Department.

Figure 1. White shrimp.

WHITE SHRIMP

S c i e n t i f i c nave.. ..... .Penaeus s e t i f e r u s ( L i nnaeus)

Common name.. ... . M i t e shrimp (F igu re 1) Other names.. .............. .Gray shrimp,

l a k e shrimp, green shrimp, green- t a i l e d shrimp, b l u e - t a i l e d shrimp, rainbow shrimp, Daytona shrimp, com- mon shrimp, southern shrimp; i n Mexico: camaron b l anco (Pe'rez- Far fan te 1969).

Class......... ................. Crustacea Order.... ....................... Decapoda Fami ly ......................... Penaeidae

Geographic range: White shrimp a re d i s - t r i b u t e d a long the A t l a n t i c coast

from F i r e Is land, New Yqrk, t o Sa in t Luc ie In1 e t , F l o r i d a (Perez-Farfante 1969), u s u a l l y i n water l e s s than 2 7 m deep (McKenzi e 1981). White shrimp i n h a b i t coasta l waters o f t h e Gul f o f Mexico from Ochl ockonee R ive r of Apalachee Bay, F lo r i da , t o Ciudad Campeche, Mexico. Centers o f abun- dance i n North and South Carol ina, Georgia, and nor theas t F l o r i d a are shown i n F igure 2 (Whi taker 1981).

Freshly caught whi te shrimp o f t e n have w ide l y spaced body chromatophores; consequently they a re 1 i g h t e r co lo red than p ink o r brown shrimp (Perez- Far fan te 1969).

MILES 0

KILOMETERS

Coasta l distribution

Centers of abundance

ATLANTIC OCEAN

F i g u r e 2. D i s t r i b u t i o n o f w h i t e shr imp a long t he coas t s o f Nor th and South Carol i n a , Georgia, and no r t heas t F l o r i d a .

2

The wh i t e shrimp i s sometimes c a l l e d t h e non-grooved shrimp (Whitaker 1981) because i t s a d r o s t r a l c a r i n a does n o t extend behind t h e midd le o f t h e carapace i n a d u l t s (L indner and Cook 1970) nor t o t h e p o s t e r i o r margin o f t h e carapace i n j u v e n i l e s (Wi 11 i ams 1965). I n con t ras t , t h e brown shrimp (P. aztecus) and -p ink shrimp (P. duorarumT, c o - e x i s t i n a alona t h e south7ast and a u l f coas ts o f t h e ~ n q t e d States, each has a deep groove ( a d r o s t r a l su lcus) ex tend ing almost t o t h e p o s t e r i o r margin o f t h e carapace. I n t h e wh i t e shrimp, t h e a d r o s t r a l su lcus i s shor t , extending t o t h e e p i g a s t r i c back t o o t h o f t h e rost rum. Gas t ro f ron ta l c a r i n a i s absent. Thelycum i s open, w i t h a n t e r o l a t e r a l r idges ; mesi a1 l y tu rned p a i r o f , f leshy protuberances on s t e r n i t e X I V (Perez-Farf ante 1969). Antenna1 f l a q e l l a 2.5 t o 3 t imes t h e body l enq th i n penaeus s e t i f e r u s as i n P. schmitt ; a species found i n Cuba, - t m q i n ~ s l a n d s , and a long eas te rn Cen t ra l and South American At1 a n t i c coast south t o Laguna, Braz i 1 ( ~ g r e z - F a r f ante 1969). Zamora and T ren t (1968) noted t h a t t h e kee l was smooth on t h e s i x t h abdominal somi t e o f post1 arvae w h i t e shrimp b u t bore sp ines on brown shrimp and p ink shrimp.

Sexes a r e e a s i l y d i s t i n g u i s h a b l e by t h e mod i f ied endopod o f t he f i r s t p a i r o f pleopods on t he males and t he open- type thelycum between t h i r d , f o u r t h , and f i f t h pereopods on t h e females (L indner and Cook 1970). At leng ths o f 28 mm, males can be d i s t i n g u i s h e d f rom females by t h e s h o r t e r and narrower endopods o f t h e f i r s t pleopods and by two prqtuberances on s t e r n i t e X I V ( Perez-Farf ante 1969) .

REASONS FOR INCLUSION IN SERIES

The wh i t e shrimp was t h e f i r s t American shrimp t o be e x t e n s i v e l y marketed f o r food. Commercial shr imping i n t h e Un i ted S ta tes began as e a r l y as 1709 (McKenzie 1981); catches reached 8,181 m e t r i c t ons ( t ) i n 1917 (when

t r a w l s rep1 aced haul se ines) , and peaked i n t h e l a t e 19201s. By t h e 19301s, t he re was concern t h a t t h e i n t e n s i v e f i s h e r y a long t h e southeastern A t l a n t i c coast (Nor th Carol ina, South Carol ina, Georgia, and nor theas t F l o r i d a ) was d e p l e t i n g t h e resource (Wi 11 i ams 1965). White shrimp c o n t r i b u t e d an est imated 95% o f shrimp catches i n 1931 (McKenzie 1981). The annual w h i t e shrimp 1 andings f o r 1976-80 averaged 3,480 t (13% below t h e 1957-80 average). The decrease was a t t r i b u t e d t o recent severe w in te r s and c o l d waters du r i ng t h a t per iod (McKenzie 1981). The harvest o f 2,685 t o f wh i te shrimp from the Southeastern States i n 1982 was valued a t $29.4 m i l l i o n (com- p u t e r i zed data from Nat iona l Marine F i she r i es Serv ice, Miami, F l o r i d a ) .

I n 1957-80, t h e mean y e a r l y ca tch o f 3,991 t o f whi te shrimp c o n t r i b u t e d 58% o f the t o t a l ca tch o f penaeid shrimp i n t h e Uni ted States (McKenzie 1981). Shrimp (80% wh i te ) accounted f o r 82% o f t h e ex-vessel 1971-77 values o f marine products landed i n Georgia (Music 1979).

Catches by spo r t shr impers are r a r e l y est imated b u t t hey must be considerable. Despi te d i f f i c u l t i e s i n e s t i m a t i n g r e c r e a t i o n a l shrimp f i s h i n g , McKenzie (1981) repo r ted t h a t i n 1973, 43% o f coas ta l boat owners i n South Ca ro l i na f i s h e d f o r shrimp w i t h non- 1 icensed gear and i n 1978, 46% o f shrimp c r a f t owners i n Nor th Ca ro l i na repo r ted " spo r t shr imping" (Thei 1 i n g 1981). The wh i t e shrimp i s an impor tan t food o f many mar ine and e s t u a r i n e f i s h e s and i n v e r t e b r a t e s and i s heavi l y used f o r b a i t i n South Carol ina, Georgia, and F l o r i d a . I n Georgia, 1,479 noncom- merc ia l r ec rea t i ona l b a i t shr impers were 1 icensed i n 1982-83 (Susan Shipman, Ga . Dep. Nat. Resour. ; pers. comm.) . I n 1980, t he est imated commercial wholesale b a i t shrimp ca tch i n nor theas t F l o r i d a was 19.6 m i l l i o n l i v e shrimp valued a t $0.77 m i l l i o n and 413.6 t o f dead shrimp valued a t $1.32 m i l 1 i o n (PlcKenzie 1981).

LIFE HISTORY

Spawni ng

Along t he south A t l a n t i c coas t o f t h e Uni ted S ta tes , wh i t e shrimp spawn from March t o November, b u t mos t l y from A p r i l (May i n South Caro l ina) t o October (Joyce 1965; L indner and Anderson 1956; Music 1979; McKenzie 1981; Shipman 1983a) . According t o McKenzi e (1981) spawning condi t i o n s extended i n t o September i n South Carol ina, Georgia, and nor theas t F l o r i d a , and i n t o October i n c e n t r a l F l o r i da . Shrimp spawn as l a t e as November i n Georgia, a c t i v i t y decreasing from south t o n o r t h (Shipman 1983a). Spawning peaks i n May and June a long the o f f s h o r e waters o f no r theas t F l o r i d a (Joyce 1965). I n general , t h e inc rease o f bottom water temperatures i n sp r i ng t r i g g e r s spawning, and r a p i d decreases i n water temperature i n t h e f a l l co inc ide w i t h t h e end o f spawning ( L i ndner and Anderson 1956; Whi t a k e r 1981). As judged by t he low percentages o f spent females i n June t o August, wh i t e shrimp may spawn as many as f o u r t imes d u r i n g t h e i r l i f e span (L indner and Anderson 1956) ; however, t h e r e i s some evidence t h a t they spawn on l y once i n Carol i na waters (Wi 11 i ams 1965).

White shrimp spawn a long t h e South A t l a n t i c coast o f t h e Un i ted S ta tes i n water more than 9 m deep (Whi taker 1983a), and w i t h i n 9 km from the shore (L indner and Cook 1970; Whi t a k e r 1983b). Spawni ng shrimp seemingly p r e f e r s a l i n i t i e s o f 27 p p t o r more (Cook and Murphy 1969). I n t he Gu l f o f Mexico most wh i t e shrimp spawn a t depths o f 8 t o 31 m ( ~ g r e z - Fa r fan te 1969) . Sexual l y mature and spent female wh i te shrimp were captured a long t he nor theas t F l o r i d a coast o n l y i n o f f s h o r e waters a t depths over 11 m (Joyce 1965). L i t t l e i s known about t he spawning l o c a t i o n o f f s h o r e from North and South Caro l ina (W i l l i ams 1965), b u t a d u l t shrimp tagged i n No r th Ed is to R iver es tuary i n South Caro l ina i n May 1983 were recaptured w i t h i n 9 km from the coas t (Whi t a k e r 1983b).

White shrimp were f i r s t spawned i n c a p t i v i t y i n 1980. The general r e q u i r e - ments f o r matura t ion and rep roduc t i on u s u a l l y f a l l i n t he range o f 20% t o 60% 1 i g h t i n t e n s i t y , 10 t o 14 h photoper iod, 20" t o 28OC water temperature, and 26 t o 34 p p t s a l i n i t i e s . I n some l a b o r a t o r y experiments, wh i te shrimp spawned o n l y a t n i g h t (L indner and Cook 1970) b u t i n o thers some spawned d u r i n g day1 i g h t (Lawrence e t a1 . 1980). The food source requ i red was f r esh marine i n v e r t e b r a t e s and f i s h supp l ied a t 3% t o 5% d r y weight o f t he weight o f t he shrimp (Lawrence e t a1 . 1983).

I n copu la t i on ( l i m i t e d t o hard- s h e l l e d i n d i v i d u a l s ) , t he male a t t aches a spermatophore onto the thelycum o f the femal e. Spermatozoa are be1 i eved t o be re leased from the spermatophore simul taneously w i t h expu ls ion o f t h e ova. About 0.5 t o 1 m i l l i o n eggs a r e d ischarged per spawn from each female (Pckez- Far fan te 1969).

The eggs o f wh i t e shrimp a re d ischarged d i r e c t l y i n t o the water and s i n k t o the bottom (Anderson 1966; L indner and Cook 1970). The sphe r i ca l and opaque r i p e eggs, which a r e 0.192 t o 0.3 mm i n diameter, have a pu rp l i sh -b lue chor ion.

Larvae

White shrimp a re i n the l a r v a l form f o r about 10 days o r more, depending on food and h a b i t a t cond i t i ons (Johnson and F i e l d i n g 1956). Eggs hatch i n t o 0.3 mm long p lank ton i c naupl i i w i t h i n 10 t o 12 h a f t e r f e r t i l i z a t i o n (Kl ima e t a1 . 1982). The nonfeeding naupl ii are c a r r i e d by p r e v a i l i n g cu r ren t s w h i l e t hey undergo f i v e mo l t s over a 24- t o 36-h pe r i od t o become f ree - feed i ng protozoea, 1 mm t o t a l l eng th (TL) (Anderson 1966). Protozoea grow t o a l e n g t h o f 2.5 nnn through t h r e e p ro tozoea l stages be fo re a t t a i n i n g t h e f i r s t stage mysis (Dobkin 1961). Postembryonic stages o f wh i t e shrimp

w y e f i r s t descr ibed by Pearson (1939). Perez-Farf ante (1969) repo r ted f i v e naupl i a1 , t h r e e pro tozoea l , and t h r e e mysis stages, fo l lowed by t h e f i r s t mastigopus o r f i r s t p o s t l a r va l stage.

Post 1 arvae and Juveni l e s

P lank ton i c post1 arvae 1 i v e o f fshore , and then move i nsho re w i t h t i d a l cu r ren t s toward e s t u a r i e s (Whi taker 1983a). At t h e end of two p o s t l a r v a l stages, about 15 t o 20 days a f t e r hatching, t h e shrimp are s t i l l p l anktonic (Anderson 1966). They en te r e s t u a r i e s du r i ng t h e second p o s t l a r va l s tage (7 mm long; 2 t o 3 weeks a f t e r ha tch ing) and then become ben th i c ( W i 1 1 i ams 1965) .

Favorable c u r r e n t s t r a n s p o r t 1 arvae and e a r l y pos t l a r vae shoreward (Perez- Farf ante 1969). Duronsl e t e t a1 . (1972) sampled p o s t l a r v a l wh i t e shrimp i n g r e a t e r numbers a t n i g h t a t t h e sur face t han a t t h e bottom o f a t i d a l pass, b u t found no depth d i f f e r e n c e s du r i ng day- 1 i g h t sampling, a l though abundance was lowest near t h e surface (0.8 m). No s i g n i f i c a n t d i f f e r e n c e s were de tec ted i n p l ank ton ne t sampling o f p o s t l a r v a l shr imp i n Georgia waters a t d i f f e r e n t depths, t imes of day, t i d a l stages, o r l una r pulses. High t u r b i d i t i e s may have in f luenced d i s t r i b u t i o n (Baisden 1983). Water temperature had l i t t l e e f f e c t on t h e movement of pos t l a r vae i n t o t h e e s t u a r i e s (McKenzie 1981).

Post1 a r va l wh i t e shrimp en te r e s t u a r i e s i n South Ca ro l i na and Nor th Carol i na from June through September (Anderson 1965). I n Georgia, nearshore and n o r t h e r l y bot tom c u r r e n t s c a r r y wh i t e shrimp post 1 arvae i n t o e s t u a r i e s and sounds (McKenzie 1981). The l a r vae en te r e s t u a r i e s i n A p r i l and e a r l y May i n t h e south A t l a n t i c , and i n June and J u l y i n Nor th Ca ro l i na (McKenzie 1981). I n nor theas te rn F l o r i d a e s t u a r i e s j u v e n i l e s were f i r s t taken i n June (Joyce 1965). White shrimp 25 t o 75 mm l ong were c l a s s i f i e d as j u v e n i l e s by Christmas e t a l . (1976), whereas

P & - e z - ~ a r f ante (1969) considered wh i t e shrimp t o be j u v e n i l e s a f t e r t hey a t t a i n e d an u l t i m a t e of r o s t r a 1 tee th : 4 t o 10 on t h e upper ros t rum and 0 t o 3 on t h e lower ros t rum w i t h modes o f 8 upper and 2. lower.

The abundance o f wh i t e shrimp peaks i n June through August i n Georgia e s t u a r i e s (Shipman 1983a). Whi le i n es tuar ies , j u v e n i l e wh i t e shrimp tend t o move f a r t h e r upstream than do j u v e n i l e p i n k o r brown shrimp -- as f a r as 160 km i n Louis iana and 210 km i n no r theas t F l o r i d a (Pgrez-Far fante 1969).

Juven i l e white, brown, and p ink shrimp tend t o i n h a b i t d i f f e r e n t subs t ra tes (Wi 11 i ams 1958). White shrimp p re fe r muddy subs t ra tes w i t h loose peat and sandy mud. They l a y t h e i r long antennae above t h e surface o f t h e subs t ra te when burrowing -- i n con t ras t t o brown and p ink shrimp, which o f t en bury t h e i r s h o r t e r antennae. Resp i ra to r y requi rements du r i ng burrowing o r cover-seekiqg, as w e l l as food, i n f l uence wh i t e shrimp pre fe rence f o r a muddy o r pea ty subs t ra te . Wi l l i ams (1958) r e p o r t e d t h a t j u v e n i l e wh i t e shrimp and brown shrimp avoid coarse subs t ra te and i n h a b i t s o f t e r bottoms because food ( r a t h e r than cover) i s more r e a d i l y avai 1 ab le there. Shallow, muddy bottoms i n waters o f low t o moderate s a l i n i t y (Anderson 1966) serve as optimum nursery grounds f o r j u v e n i l e wh i t e shrimp (Whi taker 1983a). I n t h e south A t l a n t i c s ta tes , es tua r i ne nursery areas of wh i t e shrimp are predominate ly associated w i t h Soa r t i na a l t e r n i f l o r a wetlands (McKenzie ' 1981). Juven i l e wh i t e shrimo conareaate i n sandy-muddy subs t ra te bu t juvGni7 e brown shrimp sometimes f o r c e f u l l y d i sp lace w h i t e shrimp f rom t h i s h a b i t a t (Rul i f s o n 1981). Juveni l e brown shrimp d i s p l aced j u v e n i l e wh i t e shrimp f rom grass cover i n aquar ia ( G i l e s and Zamora 1973). Staggered rec ru i tmen t of wh i t e and brown shrimp probab ly reduces compet i t ion f o r h a b i t a t (Shi pman 1983a). Inc reas ing water temperatures reduced t h e preference o f wh i t e shrimp a f f i n i t y f o r

Adu l ts sandy-mud subs t ra tes r a t h e r than s h e l l subs t ra tes .

I n 1971-81, wh i te shrimp were 1 argest i n South Ca ro l i na land ings when d e n s i t i e s were lowest, suggest ing t h a t i n t r a s p e c i f i c compet i t ion was reduced o r t ime spent i n t h e nursery areas was 1 onger (McKenz i e 1981). I n Georgi a, 1 arge shrimp predomi nated t h e harves t a f t e r w in te r freezes, l end ing suppor t t o t h e suggest ion t h a t t h e abundance of wh i t e shrimp i n south A t l a n t i c e s t u a r i e s determines t h e s i z e o f t h e shrimp i n t h e f i s h e r y o r du r i ng emigra t ion i n t h e f a l l ( Sh i pman 1983a) . As t h e season progresses t o June o r J u l y and j u v e n i l e s reach leng ths o f about 51 mm, t hey move f rom shal low marshes i n t o deeper creeks, r i v e r s , and bays (Anderson 1966). Along t h e nor theas t F l o r i d a coast, wh i te shrimp were 70 t o 80 mm l ong i n June and J u l y (Joyce 1965). A sharp decrease i n t h e mean l eng th along nor theas t F l o r i d a ' s o f f s h o r e waters i n August corresponded c l o s e l y w i t h t h e a r r i v a l of sma l l e r shrimp 120 t o 140 mm l ong ) from e s t u a r i e s (Joyce 1965). Simi 1 a r l y , an increased count (an inc rease i n numbers per pound) o f shrimp t r aw led i n

. Georgia 's o f f s h o r e waters d u r i n g August s i g n a l l e d t h e emigra t ion o f t h e summer's f i r s t r e c r u i t s (Shipman 1983a).

Among j uven i l es , wh i t e shrimp are u s u a l l y more a c t i v e than are brown o r p ink shrimp du r i ng day1 i g h t . I n Galveston Bay, Texas, t r a w l catches o f j u v e n i l e shrimp 35 t o 97 mm l ong were s i g n i f i c a n t l y l a r g e r i n dayt ime than a t n i g h t (C la rk and C a i l l o u e t 1975). I n F l o r i d a , t he percentage o f wh i te shr imp taken du r i ng dayt ime was 83% o f a l l shrimp i n inshore waters bu t o n l y 57% i n deep water (Joyce 1965). Small wh i t e shrimp may be more a c t i v e du r i ng t h e day than Targer ones (Joyce 1965). I n t h e l abo ra to ry , wh i te shrimp d i d n o t burrow, bu t they were quiescent on the bottom o r i n shal low depressions f o r several hours du r i ng t he day (Wickham and M i n k l e r 19 75).

White shrimp u s u a l l y mature sexua l l y a t age I d u r i n g t h e calendar year a f t e r t hey hatch. Mature males have j o i n e d petosmal endopods a t 105 t o 127 mm, produce r i p e sperm a t 118 mm TI-, and have f u l l y devejoped spermatophores a t 155 mm TL (Perez-Far fante 1969). The small e s t r i p e female recorded by Burkenroad (1939) was 135 mm long, and t h e minimum l e n g t h o f r i p e females i n t h e no r the rn Gu l f o f Mexico was 140 mm (S t . Amant and L indner 1966). I n F lo r i da , t h e ova r i es of wh i t e shrimp begin t o develop a t 110 mm i n f a l l and complete development when growth resumes i n t h e s p r i n g (Joyce 1965). Adu l t wh i te shrimp are powerfu l swimmers capable o f m i g r a t i n g g rea t d is tances ( w i t h c u r r e n t s ) and l i v i n g i n euphot ic l i t t o r a l zones a t r e l a t i v e l y h i gh l i g h t i n t e n s i t i e s (Young 1959). White shrimp catches i n bottom t r a w l s may a l so be low du r i ng t h e midday quiescent p e r i o d on t h e subs t ra te sur face (Wickham and M ink le r 1975). I n c o n t r a s t Veal e t a l . (1983) repo r ted t h a t s i nce wh i t e shrimp g e n e r a l l y burrow i n t o t h e bottom a t n i gh t , they may become more d i f f i c u l t t o catch. Bottom t r a w l catches a t n i g h t a l so might be lower when wh i t e shrimp are a c t i v e l y swimming i n t h e water column. Longshore cu r ren t s i n f l u e n c e movement p a t t e r n s along t h e southeast At1 a n t i c coast (Shipman 1983b). Numerous r e p o r t s from shrimpers i n d i c a t e d t h a t wh i t e shrimp sometimes school on t h e surface, p a r t i c u l a r l y i n l a t e f a l l and e a r l y w in te r du r i ng t h e i r sou the r l y o f f sho re m ig ra t i ons (Susan Shi pman , pers. comm.) . M i g r a t i o n

White shrimp along t h e southeast At1 a n t i c coast m i g r a t e southward d u r i n g autumn and e a r l y w in te r and then nor thward i n l a t e w in te r and e a r l y s p r i n g (L indner and Cook 1970; McKenzie 1981; Whitaker 1982; Shipman 1983b). More s p e c i f i c m ig ra t i ons repo r ted by Joyce (1965) showed a major s o u t h e r l y m i g r a t i o n from North Ca ro l i na t o Cape

Canaveral, F l o r i da , i n f a l l and a n o r t h e r l y m i g r a t i o n f rom t h e Cape i n sp r i ng . I n Georgia, a mark and recap tu re s tudy revea led t h a t 96% of t h e shrimp recovered i n t h e w i n t e r came f rom more s o u t h e r l y waters (Shipman 1983b) . A wh i t e shrimp tagged i n October o f f Nor th Ca ro l i na was recaptured 576 km southward of f F l o r i d a ' s east coast and one wh i t e shrimp tagged i n January o f f c e n t r a l F l o r i d a was recaptured 416 km t o t h e n o r t h o f f South Ca ro l i na (Anderson 1966). Whitaker (1981) suggested a c o r r e l a t i o n between shrimp m i g r a t i o n and 1 a t i t u d e w i t h a c t i v i t y being g rea tes t i n t h e more southern areas. I n nor theas t F lo r i da , wh i t e shrimp 120 t o 140 mm TL moved o f f s h o r e f rom August through A p r i l (Joyce 1965). Deta i l e d analyses of w h i t e shrimp sampling from June 1962 - June 1963 by Joyce (1965) revea led t h a t w h i t e shrimp caught off Cape Canaveral, F l o r i d a had migra ted f rom more no r the rn nu rse ry areas i n December and January, suggest ing t h a t Cape Canaveral i s t h e southern l i m i t o f commercial wh i t e shr imping a long F l o r i d a ' s east coast. Southward movements o f 10 t o 20 km per day d u r i n g f a l l were suggested by Joyce (1965) f o r schoo l ing w h i t e shrimp along t h e nor theas t F l o r i d a coast . Movements o f i n d i v i d u a l shrimp tagged i n Georgia waters and recaptured of f F l o r i d a ranged from 1.8 t o 6.9 n a u t i c a l m i l e s per day (Shipman 1983b). Of fshore migran ts make up t h e va luab le s p r i n g f i s h e r y f o r a d u l t females i n Georgia, South Carol ina, and Nor th Ca ro l i na i n years f o l l o w i n g r e l a t i v e l y m i l d w i n t e r s (McKenzie 1981).

White shrimp emigra t ion f rom e s t u a r i e s i s governed l a r g e l y by body s ize, age, and envi ronmental c o n d i t i o n s ( K l ima e t a1 . 1982; Shipman 1983b). Low water temperatures (<18" C) and s p r i n g t i d e s a t f u l l moon s t imu la ted mass movements f rom South Ca ro l i na e s t u a r i e s (Whi taker 1982). Dur ing ebb t i des , wh i t e shrimp tend t o school and m ig ra te near t h e sur face a t n i g h t (Benson 1982). W i 11 i ams (1958) suggested t h a t muddy subs t ra te i s n o t s t r o n g l y p r e f e r r e d d u r i n g emigra t ion

f rom e s t u a r i e s t o t h e sea. Emigrat ion was delayed i n South Ca ro l i na and Georgia when unusua l l y low f reshwater i n f l o w caused h igh s a l i n i t i e s (Shipman 1983b). I n South Carol ina, shrimp congregated i n t h e deeper channels ( s t a g i n g areas) as water temperatures dec l i ned t o about 9°C (McKenzie 1981). Recent s tud ies showed t h a t wh i t e shrimp movements o f f s h o r e i n f a l l and w i n t e r a re t r i g g e r e d by water temperature dec l i nes i n e s t u a r i e s i n t h e south At1 a n t i c (Shipman 1983b), and Lou is iana (Whi te and Boudreaux 1977) . P r e c i p i t a t i o n , s p r i n g t i d e s , and s t rong t i d a l exchanges associated w i t h no r th - e a s t e r l y storms a l so i n f l u e n c e t h e t i m i ng and magnitude o f emi g r a t i on f rom inshore waters (Shipman 1983b).

GROWTH

Juven i l e wh i t e shrimp grow du r i ng summer and fa1 1, grow s low ly over w in te r , and then resume growth as water temperatures r i s e i n t h e e s t u a r i e s du r i ng A p r i l and May. Spr ing growth i s about equal t o t h e summer growth of 18 t o 30 mm per month. S i m i l a r growth r a t e s were c a l c u l a t e d from mark- r ecap tu re s tud ies i n Georgia from 1978 t o 1981 (Shipman 1983b). The r a t e of inc rease i n weight i s r e l a t i v e l y low among the small wh i te shrimp, h ighes t i n mid-sizes, and decreasing among the 1 arger ones (Kutkuhn 1962).

Fo l low ing two mysis stages and two p o s t l a r v a l stages, young w h i t e shrimp 7 mm long en te r t h e e s t u a r i e s where t h e i r growth r a t e i s about 1.2 mm per day (Wi l l i ams 1965). J u v e n i l e shrimp i n t h e south A t l a n t i c qrow 1.0 t o 2.3 mm Der day o r 28 t o 64-mm per month ( ~ c ~ e n z i e 1981).

Young w h i t e shrimp 13 t o 68 mm l ong (mode 33 mm) f i r s t appear i n Georgia 's upper creek and marsh areas i n June; by J u l y t h e mode increases t o 43 mm and t h e range from 13 t o 103 mm ( H a r r i s 1974). White shrimp i n Georgia r i v e r s and sounds are about 78 mm l ong i n Ju ly , 108

mm i n August, 130 mm i n September, and 146 mm i n October. Dur ing w i n t e r t h e 1 engt h - f requency mode dec 1 i nes when growth stops and t h e l a r g e r shrimp m ig ra te southward o f f sho re . White shrimp t h a t overwintered i n i nsho re waters grew f rom a mqdal l e n g t h o f 118 mm t o 162 mm i n June, 176 mm i n August, and 180 mm i n September. Modal leng ths i n t h e o f f s h o r e popu la t i on were 143 mm i n March, 172 mm i n June, and 181 mm i n August. The growth o f tagged a d u l t w h i t e shrimp o f bo th sexes up t o 15 days a f t e r r e l ease was g rea tes t i n summer a t 0.41 mm per day compared w i t h 0.14, 0.10, and 0.13 mm per day i n f a l l , w in te r , and spr ing, r e s p e c t i v e l y ( Shi pman 1983b). Anderson (1966) repo r ted t h a t wh i t e shrimp were 80 mm l ong w i t h i n 2 months a f t e r ha t ch inu i n May, 110 mm by 3 months, 130 mm by 4 months, 145 mm by 5 months, and 155 mm by 6 months (November). They grew s l o w l y from November through March, b u t resumed growth i n spr ing ; 1-year-o ld spawners were 173 mm long i n May.

Female shrimp grow f a s t e r and reach l a r g e r s i zes than males ( E t z o l d and Christmas 1977). I n nor theas t F l o r i d a t h e l a r g e s t female sampled by Joyce (1965) was 192 mm l ong and t h e l a r g e s t male was 175 mm long; most shrimp l onge r than 115 mm were females. Anderson (1966) repor ted females as l ong as 197 mm i n t he Caro l inas and males as 1 ong as 182 min.

Mor ta l i t y

Few wh i t e shrimp l i v e as l ong as one year (Anderson 1966); however, mark and recap tu re s tud ies showed t h a t a few l i v e d as l ong as 27 months i n M i s s i s s i p p i ( E t z o l d and Christmas 1977), more than 17 months i n Georgia (Shipman 1983b), and as l ong as 4 years (average 18 months) i n Texas ( K l ima e t a l . 1982). Because o f t h e u s u a l l y sho r t l i f e span, t h e abundance o f wh i t e shrimp would be expected t o f l u c t u a t e w ide l y f rom year t o year bu t apparen t ly compensating f a c t o r s a r e a t work; e.g., i n 1977 a f t e r a massive w i n t e r k i l l i n Georgia coas ta l

waters, when t h e numbers o f spawning wh i t e shrimp were reduced t o 7% o f normal, subsequent r ec ru i tmen t i n t o t h e f i s h e r y was o n l y 40% below normal (Music 1979). For wh i t e shrimp i n t h e south A t l a n t i c f i s h e r y , instantaneous m o r t a l i t y r a t e s (McKenzie 1981) were 0.02 t o 0.25 ( f i s h i n g ) , 0.21 t o 0.56 ( n a t u r a l ) , and 0.24 t o 0.80 ( t o t a l ) . Weekly m o r t a l i t i e s ranged f rom 13 t o 51%; t h e lower r a t e s were nearer t o r e a l i t y f o r bo th j u v e n i l e s and a d u l t s (McKenzie 1981).

Hurr icanes cause major losses o f wh i t e shrimp i n t he Gu l f o f Mexico. A hu r r i cane s t r i k i n g t h e Lou is iana coast i n summer 1957 destroyed l a r g e numbers o f wh i t e shrimp when s a l i n i t i e s increased, cover and food supp l i es were destroyed, d i s p e r s a l and s t rand ing were excessive, and tu rbu lence i n e s t u a r i e s was h igh (Kutkuhn 1962). Hur r i cane Car la caused a 61% drop i n t h e 1961 Lou is iana ca t ch o f wh i t e shrimp and Hurr icane Cami l l e caused an 88% drop i n p roduc t ion i n M i s s i s s i p p i i n August 1969 ( B a r r e t t and G i 1 l e s p i e 1973). Sudden c o l d f r o n t s and subsequent d e c l i n e s i n water temperatures have caused m o r t a l i t y and reduced rec ru i tmen t o f wh i t e shrimp i n south A t l a n t i c shal low inshore waters; two consecut ive m i 1 d w in te r s may be requ i red t o support s p r i n g harves ts i n South Caro l ina (Whi t a k e r 1983a).

Diseases and Pa ras i t es

The e f f e c t o f diseases and p a r a s i t e s on wh i t e shrimp m o r t a l i t y i s n o t we1 1 known ( B a r r e t t and G i 1 l e s p i e 1973). A 99% 1 oss o f egg p roduc t i on was a t t r i b u t e d t o a m i c r o s p o r i d i an p a r a s i t e i n f e c t i o n o f wh i t e shrimp gonads (Gunter 1956), ye t t h e next y e a r ' s p roduc t i on was as h igh as t h a t o f t he preceeding year. V i b r i o i n f e c t i o n o f male wh i t e shrimp p m e d egg f e r t i l i z a t i o n under 1 aboratory cond i t i ons (M idd led i t ch e t a l . 1980). L i t e r a t u r e rev iews o f diseases and pa ras i t es o f penaeid shrimp show t h a t v i ruses, bac te r ia , fung i , protozoa, helminths, and nematodes o f t e n

i n f e c t shrimp (L indner and Cook 1970; Couch 1978; Overst reet 1978). Diseases and pa ras i t es ranked a f t e r p redat ion and p e r i o d i c phys ica l catastrophes as l i m i t i n g f a c t o r s i n na ture and a f t e r n u t r i t i o n and reproduc t ion requirements i n m a r i c u l t u r e (Couch 1978). Symbionts may be r e l a t e d t o shrimp k i l l s du r i ng 1 ow oxyqen cond i t i ons (Overs t ree t 1978). A par -as i t i c cestode, ~ r o c h r i s t a n e l i a penaei , i n f e c t i n g the hepatopancreas o f a d u l t s h r i m p i s o f some concern i n t he M i s s i s s i p p i Sound; however, from an economi c s tandpoint , m i c r o s p o r i d i an protozoans t h a t cause a "co t ton" appearance i n t h e musculature o f shrimp are t h e most t h rea ten ing (Christmas e t a1 . 1976). I n Georgia i n 1978-81, m i c r o s p o r i d i an pa ras i t es were observed i n 3.9% o f 33,350 wh i te shrimp captured f o r tagging. Lower recovery r a t e s o f tags from in fec ted than f rom un in fec ted shrimp suggested h igher m o r t a l i t y among t h e i n fec ted shrimp (Shipman 1983b). Hut ton e t a l . (1959) suggested t h a t i n f e c t e d shrimp may be more suscep t i b l e t o p reda t i on and disease.

THE FISHERY

The c h a r a c t e r i s t i c s o f t he wh i te shrimp f i s h e r y i n t he south A t l a n t i c -- i n c l u d i n g processing, rnarket i ng, economics, and s o c i o l o g i c a l aspects -- were repor ted by McKenzie (1981). The shrimp i n d u s t r i e s along t h e A t l a n t i c coast o f Nor th Carol ina, South Carol ina, Georgia, and F l o r i d a are based main ly on whi te, brown, and p ink shrimp. F l o r i - da 's f i s h e r y inc ludes the rock shrimp (S i cyon ia b r e v i r o s t r i s ) . White shrimp c o n t r i b u t e d 58% o f t he t o t a l 1957-80 ca tch i n t h e f o u r States. The low was 31%, f o l l o w i n g t h e severe w in te r o f 1976-77, and t h e h igh was 76% i n 1973. White shrimp con t r i bu ted an average of 81% t o t h e ca tch along t h e east coast of F lo r ida , and 78% o f t h e t o t a l catch i n Georgia, 58% i n South Carol ina, and 6% i n North Carol ina. The average annual wh i t e shrimp 1 andings i n 1976-1980 (mi 11 ions of pounds, heads o f f ) were 0.16 f o r Nor th Carol ina, 2.47 f o r South

Carol ina, 3.41 f o r Georgia, and 1.63 along the east coast o f F l o r i d a (McKenzie 1981).

The average annual wh i t e shrimp landings i n 1976-80 were 13% below the average o f t he 1957-80 landings. 'The decrease was a t t r i b u t e d t o t he severe w in te rs o f 1976-77 and 1977-78. The Georgia Department of Natura l Resources (1983) repor ted a 34% drop i n 1981 from the 1971-80 wh i te shrimp landings. The drop was caused by unusua l ly low w i n t e r water temperatures i n 1980-81. Annual catches i n Georgia and F l o r i d a have been r e l a t i v e l y steady, averaging near 18% and 29%, respec t i ve l y , about the c o e f f i c i e n t o f v a r i a t i o n f o r t he 195 7-80 (24-year) average; t h i s average v a r i e d 54% i n South Ca ro l i na and 114% i n North Ca ro l i na (McKenzie 1981). I n t h e South A t l a n t i c , wh i t e shrimp 1 andings (heads-o f f ) i n 1957-80 ranged from a low o f 3.2 m i l l i o n 1b i n 1977 t o a h igh of 12.2 m i l l i o n 1b i n 1971. Although f l u c t u a t i o n s i n abundance are n a t u r a l and are expected even when environmental f a c t o r s appear f avo rab le (McKenzie 1981) , a1 t e r a t i o n o f h a b i t a t s by p o l l u t i o n o r physical causes i n numerous es tua r i es are becoming ser ious f a c t o r s i n f l u e n c i n g shrimp produc t ion (E tzo ld e t a1 . 1983).

White shrimp enter t h e commercial f i s h e r y when the g rav id shrimp congreqate o f f t h e c e n t r a l and southward coast o f South Ca ro l i na i n A p r i l o r May and remain i n t he South Caro l ina f i s h e r y through June o r e a r l y Ju l y . I n Georgia, the wh i te shrimp f i s h e r y season opens i n June i n t e r r i t o r i a1 offshore waters. Juveni les en ter t h e coas ta l f i s h e r y i n August i n South Carol ina, Georgia, and nor theas t F lo r i da . I n North Carol ina, t hey are caught ma in ly i n t h e f a l l i n t h e area from Southport t o Cape Fear. The f i s h e r y cont inues through mi d- December i n South Caro l ina and t o t h e end of December i n Georgia and no r the rn F l o r i d a (McKenzie 1981). Catches i n nearshore waters o f Georgia are lowest i n June and peak i n August and September (Music 1979). Catch per u n i t o f e f f o r t

was h i ghes t i n no r t he rn Georg ia 's o f f sho re waters i n l a t e summer and f a l l and land ings peaked i n September and October (Ga. Dep. Nat. Resour. Coasta l Resources Div., Data Manage. Sect ion, pers. comm.) . As water temperatures drop, w h i t e shr imp move southward and a re caught i n coas ta l waters o f extreme southern Georgia i n January (Music 1979); however, some may be caught as l a t e as February depending on t h e da te o f c l osu re o f t e r r i t o r i a l waters (Susan Shi pman, pers. comm.) . Joyce (1965) repor ted t h a t abundance peaked i n December and January i n no r t he rn F l o r i d a from St. August ine t o Cape Canaveral .

>lost commercial shr imp ca t ch i s made w i t h i n 9 km o f t h e coas t ( E t z o l d e t a l . 1983) on t r a w l a b l e bottoms w i t h i n t h e 11-m depth contour . The b read th o f t h e Cont inen ta l S h e l f w i t h i n t h e 11-m (6-fathom) con tour i s g r e a t e s t a long t h e no r t he rn and c e n t r a l Georgia s h e l f , bu t i s narrower a long t h e coas t o f no r t heas t F l o r i da , South Caro l ina, and Nor th Caro l ina . About 99% o f Nor th C a r o l i n a ' s w h i t e shr imp ca t ch was taken i n i t s t e r r i t o r i a l waters. For t h e o t h e r s ta tes , t h e percentages were 90% f o r South Caro l ina, 85% f o r F l o r i d a , and 59% f o r Georgia. I n Georgi a, two 3-year s t u d i e s (1974-77, 19 78-81) o f month ly shr imp d i s t r i b u t i o n and abundance were made by us i ng 30-n in t r a w l samples t o eva lua te shr imp a t 36 s t a t i o n s i n e s t u a r i e s and nearshore waters ou t t o 4.8 km (Music 19 79; Shi pman ,1983a). The ca t ch o f w h i t e shr imp per hour averaged 50 1b and 32 l b i n sounds (52% and 44% o f t o t a l ) , 44 1b and 34 I b i n creeks (46% and 48% o f t h e t o t a l ) , and 1.5 1b and 5 1b i n o u t s i d e waters (2% and 7% o f t h e t o t a l ) .

Freshwater i n f l o w i s t h e dominant f a c t o r i n f l u e n c i n g abundance, d i s t r i bu- t i o n , and growth o f w h i t e shr imp (McKenzie 1981). Dur ing t h e drought and low f reshwater i n f l o w i n 1980, t h e shr imp moved f u r t h e r up es tua r i es , which lengthened t h e i r r es i dency t h e r e and

increased m o r t a l i t y . The lower 1 andings i n 1980-81 were caused by low f reshwater i n f l o w and low w i n t e r water temperatures.

S ince j u v e n i l e wh i t e shr imp l i v e i n coas ta l wetlands, t h e areas o f such wetlands are use fu l measures o f p o t e n t i a1 abundance (Turner 1977). The areas o f coas ta l wet1 and a re 79,826 ha i n Nor th Caro l ina, 204,146 ha i n South Caro l ina, 192,508 ha i n Georgia, and 47,631 ha i n no r t heas t F l o r i d a (McKenzie 1981). None o f t h e f o u r s t a t e s pe rm i t commerci a1 t r a w l i n g i n des ignated nursery areas. A p o s i t i v e r e 1 a t i o n between t h e 1962 and 1963 f a l l w h i t e shr imp commercial 1 andings i n F l o r i d a w i t h r e l a t i v e abundance o f J u l y inshore samples was r e p o r t e d by Joyce (1965). Produc t ion es t imates o f o f f s h o r e harves ts i n Alabama and M i s s i s s i p p i a l s o have shown a s t r ong r e l a t i o n t o i nsho re abundance o f j u v e n i l e s (S t . Amant and L indner 1966; Loesch 1976). Christmas and E t z o l d (1977) concluded t h a t subsequent year r e c r u i t m e n t i s n o t a major c o n s i d e r a t i o n i n management because i t i s l a r g e l y independent o f t h e abundance o f paren t s tock; t he re fo re , management would be aimed toward maximimum sus ta ined y i e l d f rom t h e c u r r e n t y e a r ' s r ec ru i tmen t . I n Georgia, t h e r e was 1 i t t l e r e 1 a t i o n between t h e s i z e o f t h e f a l l wh i t e shr imp land ings and s a l i n i t y , abundance o f j u v e n i l e s i n August, and abundance of g r a v i d shr imp i n t he preceding s p r i n g (Shipman 1983a).

Spor t and noncommercial b a i t shr imp f i s h e r i e s a r e d i f f i c u l t t o eva lua te because n o t a l l a re 1 icensed (McKenzie 1981). Recrea t iona l ca t ch has been es t imated t o equal 10% o f t h e south A t l a n t i c commercial ca t ch ( E t z o l d e t a l . 1983). The most impor tan t da ta are f rom boat r e g i s t r a t i o n s . I n 1973, 44% o f t h e 16,780 r e g i s t e r e d r e c r e a t i o n a l boat owners i n 11 eas te rn South Ca ro l i na coun t i es caught an es t imated t o t a l o f 371 mt o f shrimp. O f t h e 15,888 shr imp ing c r a f t owners l i censed i n Nor th Ca ro l i na i n 1978, 46% were s p o r t f i shermen who caught as much as 3% o f

t h e commercial catch. I n Georgia i n 1982-83, t h e r e were 1,479 b a i t shrimping 1 icenses, 76 commercial b a i t 1 icenses, and 1,959 commercial food shr imping 1 icenses. I n 1980-81, 127 non- commercial l i censes were issued f o r t h e S t . Johns River , F l o r i d a a l l ow ing spo r t f ishermen t o t ake up t o 50 1b per day of shrimp by t r a w l i n g i n inshore waters o n l y on weekends and ho l idays .

Although shrimp are important b a i t f o r spo r t f i s h i n g i n North and South Carol i na, t he 1 i ve-ba i t i n d u s t r y i s r e 1 a t i v e l y small (McKenzie 1981). I n F l o r i d a t he commercial b a i t f i s h e r y landed and so ld 22.3 m i l l i o n l i v e shrimp annual l y i n 1972-80 (McKenzie 1981).

ECOLOGICAL RClLE

White shrimp convert d e t r i t u s , p l a n t ma te r i a l , microorganisms, macroinver tebrates, and f i s h p a r t s i n t o use fu l p r o t e i n f o r carn ivores (e.g. o the r inver tebra tes , f i s h , and man). Naupl ii subs i s t on yo l k granules u n t i l t hey reach t h e protozoea I stage (McKenzi e 1981) . White shrimp l a r vae feed on zooplankton and phytoplankton; wh i t e shrimp protozoea feed on green a1 gae, diatoms, o r copepods (Dobkin 1961). I n a l abo ra to ry t e s t , c u l t u r e d algae were fed t o protozoea and newly hatched b r i n e shrimp up t o the mysis stage (Cook and Murphy 1969). Ea r l y stages o f wh i te shrimp l a r vae feed on p lank ton and suspended d e t r i t u s (Christmas and Etzo l d 1977).

Juven i le and adu l t wh i t e shrimp are ben th i c omnivores; t h e major d i f f e r e n c e s i n food s e l e c t i o n are t h e k inds and a v a i l a b i l i t y of m a t e r i a l s selected. Juveni l e and adu l t penaeids are benth ic omnivores t h a t feed l a r g e l y a t n igh t , except i n t u r b i d waters (McKenzie 1981). Fecal p e l l e t s o f f i s h and i nve r teb ra tes can be an important food i tem of j u v e n i l e shrimp. Lindner and Cook (1970) noted t h a t wh i t e shrimp were s e l e c t i v e p a r t i c u l a t e feeders. Major food repor ted i n t h r e e s tud ies were

d e t r i t u s , c h i t i n , pa r t s o f annel ids and gastropods, f i s h par ts , bryozoans, sponges, cora ls , f i 1 aments of algae, and vascular p l a n t stems and r o o t s (Christmas and E tzo ld 1977). L i p i d s supp l ied by annel ids i n t h e d i e t were important f o r ovar ian matura t ion (M idd led i t ch e t a l . 1980).

Cannibal ism i s common among j u y e n i l e and adu l t wh i te shrimp (Perez-Farfante 1969), bu t McKenzie (1981) suggested t h a t t h e cannibal ism repo r ted i n t h e l i t e r a t u r e was r e l a t e d t o crowding i n aquaria. B o t t i n o e t a l . (1980) found t h a t body f a t t y acids i n shrimp were i n f l uenced by d i e t . Food convers ion r a t i o s o f 1.8 and 1.9 (i.e., 1.8 o r 1.9 1b o f food y i e l d 1 1b o f shrimp) were repo r ted f o r wh i t e shrimp fed i n two marine ponds a t Marifarms, Inc. , Panama C i t y , F l o r i d a (Brown 1977). Assimi l a t i o n e f f i c i e n c y i n j u v e n i l e wh i t e shrimp may reach 80% t o 85% f o r a v a r i e t y o f p l ant and animal m a t e r i a l s (McKenzie 1981). White shrimp were an important food f o r many marine and e$ tua r i ne f i s h (Gunter 1956; Perez-Farfante 1969; L indner and Cook 1970; McKenzie 1981 ; Benson 1982) . Larva l and j u v e n i l e shrimp were important food items f o r 13 o f 21 j u v e n i l e f i s h species captured from seagrass beds i n F l o r i d a es tua r i es o f t h e Gul f o f Mexico (Carr and Adams 1973).

White shrimp are i nva luab le i n t h e food chains o f coas ta l 'waters. They r e c y c l e bas ic n u t r i e n t s by feed ing on organ ic ma t te r and microorganisms i n sediments (Odum 1971; Carr and Adams 1973). Concentrat ions along t h e Lou i s i ana coast are g rea tes t where subs t ra tes are h igh i n organic content and where water temperatures and s a l i n i t i e s are f avo rab le ( B a r r e t t and G i 1 l e s p i e 1973; Gaidry 1974). Kutkuhn (1966) i 1 l u s t r a t e d t h e dependence of w h i t e shrimp on t h e es tua r i ne environment . Juveni l e s t o l e r a t e lower s a l i n i t i e s than do many o ther f i s h and s h e l l f i s h ; t h i s s a l i n i t y t o l e rance reduces compet i t ion between shrimp and

f i s h and may be as impor tan t as f ood supp ly f o r t h e growth and s u r v i v a l o f t hese seasonal m ig ran ts (Hedgpeth 1963; Gunter 1967).

ENVIRONMENTAL REQUIREMENTS

Water temperature d i r e c t l y o r i n d i r e c t l y i n f l u e n c e s w h i t e shr imp spawning, growth, h a b i t a t se l ec t i on , osmoregul a t ion , movement, m ig ra t i on , and m o r t a l i t y . Sp r i ng water temperature inc reases t r i g g e r spawning, and r a p i d water temperature d e c l i n e s i n f a 1 1 po r t end t h e end o f spawning (L i ndne r and Anderson 1956). Growth i s f a s t e s t i n summer and slow o r n e g l i g i b l e i n w i n t e r . Water temperatures below 20°C i n h i b i t growth of j u v e n i l e shr imp ( E t z o l d and Christmas 1977) and growth i s v i r t u a l l y n i l a t 16°C (S t . Amant and L, indner 1966). Growth r a t e s inc rease r a p i d l y as temperatures inc rease above 20" C . Increased water temperature a f f e c t s m o l t i n g r a t e ( ~ G r e z - ~ a r f ante 1969). Good c o r r e l a t i o n between hea t i ng - degree-days and c a t c h l e f f o r t r a t i o f o r penaeid shr imp was s i m i l a r t o c o r r e l a t i ons of y i e l d -pe r -hec ta re versus 1 a t i t ude (Turner 1977). Temperature and f ood supp ly l i m i t e d t h e growth o f w h i t e shr imp p o s t l a r v a e more than d i d s a l i n i t y d i f f e r e n c e s between 2 and 35 pp t (Ze in -E ld i n 1964).

Severe w i n t e r s i n 1939-40, 1966, 1976-77, and 1977-78 caused mass m o r t a l i t y and reduced catches i n t h e South A t l a n t i c w h i t e shrimp f i s h e r y ( McKenzi e 1981; Sh i prnan 1983a; Whit aker 1983a). The Georg ia Department o f Na tu ra l Resources (1983) r e p o r t e d a 34% drop i n w h i t e shr imp l and ings i n 1981 and a 99% drop i n 1981 s p r i n g ca t ch o f r o e shrimp a f t e r t h e unusua l l y c o l d 1980-81 w i n t e r . White shr imp a re more t o l e r a n t o f h i gh temperatures and l e s s t o l e r a n t o f low temperatures than e i t h e r brown o r p i nk shrimp ( E t z o l d and Christmas 1977). Among pos t la rvae , brown shrimp were more r e s i s t a n t than

w h i t e shr imp t o h i ghe r temperatures.

White shr imp m o r t a l i t y was r e p o r t e d a t water temperatures o f 8°C and lower (Joyce 1965). M o r t a l i t y o f w h i t e shrimp i s t o t a l a t 3°C o r lower, r ega rd less of s a l i n i t y . White shr imp s u r v i v a l a t low temperatures depends on ambient temperature, t h e r a t e o f temperature dec l i ne , t h e d u r a t i o n o f low temperatures and s a l i n i t y (Joyce 1965) . The impact o f low water temperature and low s a l i n i t y on w h i t e shr imp was d iscussed by Music (1979) and Shiprnan (1983a). Adu l t w h i t e shr imp ( > 90 mm long) may be more s u s c e p t i b l e than j u v e n i l e s t o c o l d temperatures (Whi taker 1983a). Wiesepape (1975) found t h e 24-h LC50 ( tempera tu re caus ing 50% mor ta l i ty i n 24 h ) t o be 36' and 37°C f o r w h i t e shrimp acc l imated a t 29" and 34"C, r e s p e c t i v e l y . Post 1 arvae and 30-mm long j u v e n i l e s have s i m i l a r b u t h i ghe r r e s i s t a n c e t imes than 50-mm juven i l es .

S a l i n i t y

Adu l t w h i t e shr imp spawn o f f s h o r e where s a l i n i t i e s a re a t l e a s t 27 pp t . The l a r v a e move shoreward and become second-stage pos t l a r vae as they e n t e r e s t u a r i e s on f l o o d t i d e s . J u v e n i l e w h i t e shrimp moved 160 km upstream i n t o water o f l e s s than 1.0-ppt s a l i n i t y waters i n t h e St . Johns R iver , F l o r i d a (Joyce 1965). J u v e n i l e w h i t e shr imp have even been recovered f rom Lake Monroe Power St a t i on f i 1 t e r screens l oca ted 270 km f rom t h e mouth o f t h e St . Johns R i ve r -- e s p e c i a l l y when low r a i n f a l l and low r i v e r s tages caused reve rse t i d a l f l o w (Edwin Joyce, pers. cbmm., February 1984). The h i gh ca lc ium i o n concen t ra t i ons i n t he St. Johns R iver may e x p l a i n the r e l a t i v e ease w i t h which mar ine species e n t e r and remain i n low s a l i n i t y waters (Joyce 1965). The lowest s a l i n i t y i n which w h i t e shrimp were recorded i n t h e noy thern G u l f o f Mexico was 0.42 p p t ( Perez-Farf ante 1969). Al though f i e l d s t u d i e s i n d i c a t e t h a t j u v e n i l e w h i t e shr imp p r e f e r low s a l i n i t i e s , l a b o r a t o r y

s tud ies have revealed t h a t wh i te shrimp appear t o t o l e r a t e a wide range o f s a l i n i t i e s ; t hey have been successful l y reqred a t s a l i n i t i e s o f 18 t o 34 pp t ( Perez-Farf ante 1969). McKenzi e (1981) c i t e d several s tud ies i n which f a s t growth was repor ted f o r wh i t e shrimp a t s a l i n i t i e s of 7 t o 15 ppt.

White shrimp i n Georgia move toward h igher s a l i n i t y waters as sexual development progresses, and most spawn o f f s h o r e i n t h e sea ( H a r r i s 1974).

Tem~era tu re - S a l i n i t y I n t e r a c t i o n s

Temperature-sal i n i t y t o 1 erance ranges f o r wh i te shrimp vary a t d i f f e r e n t l i f e stages, bu t t he i n t e r a c t ions are more pronounced a t t he extremes of to le rance. For example, Couch (1978) repo r ted t h a t broken-back syndrome (do rsa l separat ion o f t h i r d and f o u r t h p l e u r a l p l a tes on abdominal) appears a f t e r sudden drops i n s a l i n i t y ( f r om 15 pp t t o 3 p p t ) i n c o l d water (8°C). The c r i t i c a l thermal maxima fo r wh i t e shrimp are in f luenced l a r g e l y by acc l ima t i on temperatures, and t o a l esse r ex ten t by s a l i n i t y and s i z e of t e s t animal (Laney 1973). Freshwater i n f l o w may a f f e c t coas ta l water temperatures, which i n t u r n a f f e c t t h e growth r a t e s (Whi te and Boudreaux 1977) and m i g r a t i o n o f wh i te shrimp (Shipman 1983b). Spr ing spawning of wh i t e shrimp co inc ides w i t h a r a p i d r i s e i n bottom water temperatures i n h igh s a l i n i t y o f f sho re waters (McKenzie 1981).

Substrate

White shrimp p r e f e r shal low, muddy-bottom substrate. Landings o f shrimp along the Louis iana coast were h ighes t i n areas where subs t ra tes were h i g h l y o rgan ic ( B a r r e t t and G i l l e s p i e 1973; Gaidry 1974). A r e l a t i v e h igher 1 i nea r c o r r e l a t i o n (RZ = 0.69) between i n t e r t i d a l land area and average annual shrimp catch along Lou i s i ana ' s inshore reg ions was repo r ted by Turner (1977). The r e 1 a t i on between inshore catches and hectares of vegetated es tua r i ne h a b i t a t

i n t he nor theas tern Gu l f o f Mexico (Tampa Bay, F lo r i da , t o Mob i le Bay and Perdido Bay, Alabama) a l so showed a s t rong c o r r e l a t i o n ( R ~ = 0.64). A d i r e c t r e 1 a t i onsh ip between commercial shrimp landings and i n t e r t i d a l vegetated areas and degrees l a t i t u d e was repo r ted by Turner (1977). The annual landings (kg lha) i n 1955-64 were 19.7 i n North Carol ina, 7.9 i n South Carol ina, 13 i n Georgia, and 22.4 i n east F lo r i da . White shrimp undoubtedly composed most o f t he landings except i n North Carol ina. Southward f a 1 1 m i g r a t i o n probably account f o r t h e h igh landings from F l o r i d a waters. The area o f nearshore s o f t sediments c o r r e l a t e we1 1 w i t h wh i te and brown shrimp d i s t r i b u t i o n from Pamlico Sound, North Caro l ina t o nor thern F l o r i d a (McKenzie 1981).

Temporal and s p a t i a l s h i f t s by brown, white, and p ink shrimp he lp reduce d i r e c t i n t e r s p e c i f i c compet i t ion e s p e c i a l l y f o r c e r t a i n subs t ra te (Lassuy 1983). White shrimp burrow l ess deeply i n t o muddy subs t ra tes and are more a c t i v e i n d a y l i g h t than are brown o r p ink shrimp. Staggered seasonal recru i tment o f brown and wh i te shrimp i n t o south A t l a n t i c es tua r i es would reduce compet i t ion (Baisden 1983).

Other Environmental Considerat ions

The l oss o f nursery grounds has been considered t h e major t h r e a t t o t h e wh i te shrimp f i s h e r y i n t h e Gulf of Mexico because t h a t i s where shrimp are most vu lnerab le t o h a b i t a t d is tu rbance (Gunter 1956). Studies i n F lo r ida , Louisiana, and Texas i d e n t i f i e d 1 andf i 11, dredging, and impoundments as major detr iments t o shrimp produc t ion (Christmas and E tzo ld 1977; E tzo ld e t a l . 1983). Because o f t h e l oss of r i c h o rgan ic m a t e r i a l along bulkheads, shrimp abundance the re was reduced t o about 118 t h a t o f nearby una l te red shore l ines (Mock 1967). About 18,171 ha of wetlands, 3.5% o f t he t o t a l , were l o s t from t h e South A t l a n t i c coast between 1954 and 1968 (McKenzie 1981). Manmade

cana ls i n Lou i s i ana e s t u a r i e s have i nc reased s a l i n i t y and adverse ly a f f e c t e d w h i t e shr imp s u r v i v a l and growth (B ig1 ane and LaFleur 1968). Inc reased s a l i n i t i e s have favored brown shr imp over w h i t e shr imp i n t h e c e n t r a l - no r t he rn G u l f o f Mexico (Chr is tmas and E t z o l d 1977). The e f f e c t s o f p e s t i c i d e s and p o l l u t i o n on shr imp h a b i t a t a long t h e g u l f coas t a re a l s o o f concern ( B i g l a n e and LaF leur 1968; Christmas and E t z o l d 1977). Severa l exampl es o f w h i t e shr imp l osses t o p e s t i c i d e s a long t h e South C a r o l i n a coas t were g i ven by McKenzie (1981) and

t h e t o x i c i t i e s and b i o l o g i c a l e f f e c t s o f p e s t i c i d e s , heavy meta ls , pet ro leum products , and chemotherapeut ic chemica ls were g i ven by Couch (1978). Trawl ca tches o f w h i t e shr imp dropped below seasonal averages when d i s s o l v e d oxygen was below 3.0 mg/l i n a l t e red , e u t r o p h i c cana ls assoc ia ted w i t h hous ing developments i n West Bay, Texas ( T r e n t e t a1 . 1976). M a i n t a i n i n g s u i t a b l e nu r se r y grounds u l t i m a t e l y may dec i de t h e f u t u r e o f t h e shr imp resources o f t h e g u l f coas t (Chr is tmas and E t z o l d 1977) and south A t l a n t i c (McKenzie 1981; E t z o l d e t a l . 1983).

LITERATURE CITED

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B a r r e t t , B.B., and M.C. G i l l e s p i e . 1973. Pr imary f a c t o r s which i n f l u e n c e commerci a1 shrimp produc t ion i n coas ta l Louis iana. La. Dep. W i l d l . Fish. Tech. B u l l . 9. 28 pp.

Benson, N.G., ed. 1982. White shrimp Penaeus s e t i f e r u s . Pages 7-9 i n L - i f e h i s t o r y requi rements 3 se lec ted f i n f i s h and s h e l l f i s h i n M i s s i s s i p p i Sound and adjacent areas. U.S. F ish W i l d l . Serv. B i 01. Serv. Program FWS/OBS-81/51. 9 7 PP*

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Christmas, J.Y., W. Langley, and T. VanDevender. 1976. I n v e s t i g a t i o n s o f commerc ia l ly impor tan t penaeid shrimp i n M i s s i s s i p p i . Gu l f Coast Res. Lab., Ocean Springs, Miss. 66 PP.

Clark, S.H., and C.W. C a i l l o u e t . 1975. Die1 f l u c t u a t i o n s i n catches of j u v e n i l e brown and w h i t e shrimp i n a Texas es tua r i ne canal . Cont r ib . Mar. Sc i . 19:119-122.

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Couch, J.A. 1978. Diseases, paras i tes , and t o x i c responses o f commercial penaeid shrimps o f t h e Gu l f o f Mexico and s o u t h , A t l a n t i c coasts o f Nor th America. U.S. N a t l . Mar. F ish. Serv . Fish. B u l l . 76(1) : 1-44.

Dobkin, S. 1961. E a r l y developmental stages o f p i nk shrimp, Penaeus duorarum, f rom F l o r i d a U.S. F ish W i l d l . Serv. F ish. B u l l . 190(61) :321-349.

Duronslet, M.J., J.M. Lyon, and F. Marul l o . 1972. V e r t i c a l d i s t r i b u t i o n o f p o s t l a r v a l brown, Penaeus aztecus. and white. P. s e t i f e r u s , shrimp &ring immiqra t ion throuah a t i d a l Dass. ., ., Trans. Am. F ish. SOC .

Etzold, D.J., and J.Y. Christmas. 1977. A comprehensive summary o f t h e shrimp f i s h e r y o f t h e Gu l f of Mexico Un i t ed States; a r e g i o n a l management plan. Gu l f Coast Res. Lab. Tech. Rep. Ser. 110. 2 , P a r t 2. 20 PP*

Etzold, D.J., C.Y. Christmas, and V. Blomo. 1983. Chapter I. Analys is o f envi ronmental and demand f a c t o r s on shrimp produc t ion i n t h e g u l f and south A t l a n t i c Un i t ed States. Pages 1-205 i n W.D. Chauvin, ed. Assessment 5 shrimp i n d u s t r y p o t e n t i a l s and c o n f l i c t s . Vol. I. Shrimp Notes Inc., New Or1 eans, La.

Gaidry, W.J., 111. 1974. C o r r e l a t i o n s between inshore s p r i n g w h i t e shrimp popu la t i on d e n s i t i e s and o f f s h o r e ove rw in te r i ng popul a t ions . La. W i l d l . Fish. Comm. Tech. B u l l . 12. 16 PP.

Georgia Department o f Na tu ra l Resources. 1983. Georgi a 1 andi ngs, annu a1

Gi les, J.H., and G. Zamora. 1973. Cover as a f a c t o r i n h a b i t a t s e l e c t i o n by j u v e n i l e brown (Penaeus aztecus) and wh i t e (P. s e t i f e r u s l shrimD. Trans. Am. t i s h . SOC . 102(1) :144-145.

Gunter, G. 1956. P r i n c i p l e s o f shrimp f i s h e r y management. Proc. Gu l f Caribb. Fish. I n s t . 8:99-106.

Gunter, G. 1967. Some r e l a t i o n s h i p s o f e s t u a r i e s t o t h e f i s h e r i e s o f t h e Gu l f o f Mexico. Pages 621-638 i n G.A. Lau f f , ed. Es tuar ies . A m 7 Assoc. Adv. Sci. Publ. 83.

Ha r r i s , C.D. 1974. Observat ions on t h e w h i t e shrimp (Penaeus s e t i f e r u s ) i n Georqi a. G a . r N a t ; Resour. ~ o n t r i b . Ser. No. ' 27:54 pp.

Hedgpeth, J.W. 1963. Bi 01 og i c a l aspects, e s t u a r i e s and lagoons. Chapter 23, Pages 693-729 i n J.H. Hedgpeth, ed. T r e a t i s e o n m a r i n e ecology and paleoecology. Geol . Soc. Am. Mem. 67.

Hutton, R.F., F. Sogandares-Barnal, B. Eldred, R. M. Ing les, and K.D. Woodburn. 1959. I n v e s t i g a t i o n s on t h e p a r a s i t e s and diseases o f sa l twa te r shrimps (Penaeidae) of spo r t s and commerci a1 importance i n F l o r i d a . F la. Board Conserv. Mar. Lab. Tech. Ser. No. 26. 38 pp.

Johnson, M.C., and J.R. F i e l d i n g . 1956. P r o ~ a q a t i o n o f t h e wh i t e shrimp, ~ e n a e i s s e t i f e r u s (L inn . ) i n c a ~ t i v i t v . Tul ane Stud. Zoo1 .

Joyce, E.A., J r . 1965. The commercial shrimps o f t h e nor theas t coast o f F l o r i d a . F l a. Board Conserv. Mar. Lab. Pro f . Pap. Ser 6. 224 pp.

Klima, E.F., K.N. Baxter, and F.J. Pa te l la , J r . 1982. A rev iew of

t h e o f f sho re shrimp f i s h e r y and the 1981 Texas c losure . Mar. Fish. Rev. 44:16-30.

Kutkuhn, J.H. 1962. Gul f o f Mexico commercial shrimp popu la t ions - t rends and c h a r a c t e r i s t i c s , 1956-59. U.S. F i s h W i l d l . Serv. F ish. B u l l . 62:343-402.

Kutkuhn, J.H. 1966. The r o l e o f es tua r i es i n t h e development and perpetua t ion o f commercial shrimp resources. Pages 16-36 i n R.F. Smith, A.H. Swartz, a n T W.H. Massmann, eds. A symposium on es tua r i ne f i s h e r i e s . Am. F ish. Soc. Spec. Publ. 3.

Laney, R.W. 1973. A comparison o f t h e c r i t i c a l thermal maxima o f j u v e n i l e brown shrimp (Penaeus aztecus aztecus Ives) a n i t e shrimD

~ -

(Penaeus s e t i f e r u s (Linnaeus). M.S. Thesis. North Ca ro l i na S ta te Un i ve rs i t y , Raleigh, N.C. 63 pp.

Lassuy, D.R. 1983. 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 (Gulf of Mexico) . Brown shrimp. U.S. F ish W i l d l . Serv. FWS/OBS-82111.1. 15 pp.

Lawrence, A.L., Y. Akamine, B.S. Midd led i tch , G. Chamberlain, and D. Hutchins. 1980. Matura t ion and reproduct i on o f Penaeus s e t i f e r u s i n c a p t i v i t y . Proc. World M a r i c u l t . Soc. 11:481-487.

Lawrence, A.L., M.A. Johns, and W.L. G r i f f i n . 1983. Shrimp m a r i c u l t u r e s t a t e of t h e a r t . Texas A&M Univ. . .

Sea Grant Col lege Progr. TAMU-SG- 84- 502 : 12 pp.

Lindner, M.J., and W.W. Anderson. 1956. Growth, migra t ions , spawning and s i z e d i s t r i b u t i o n of shrimp Penaeus s e t i f e r u s . U.S. F i sh W i l d l x l i s h . B u l l . 56:554-645.

Lindner, M.J., and H.L. Cook. 1970. Synopsis of b i o l o q i c a l da ta on t h e wh i te shrimp ~enaeus s e t i f e r u s (Linnaeus) 1767. F i s h e r i e s Synopsi s 101. FA0 Fish. Rep. 4: 1439-1469.

Loesch, H.C. 1976. Shrimp popul a t i o n d e n s i t i e s w i t h i n Mobi le Bay. Gu l f Res. Rep. 5(2):11-16.

McKenzie, M.D., ed. 1981. P r o f i l e o f t h e penaeid shrimp f i s h e r y i n t h e south A t l a n t i c . South A t l a n t i c F i shery Management Counci 1, Charleston, S.C. 321 pp.

Midd led i tch , B.S., S.R. M iss le r , H.B. Hines, J.P. McVey, A. Brown, D.G. Ward, and A.L. Lawrence. 1980. Met abol i c prof i 1 es o f penaeid shrimp: d i e t a r y l i p i d s and ovar ian matura t ion . J. Chromatogr . 195: 359-368.

Mock, C.R. 1967. Natura l and a l t e r e d es tua r i ne h a b i t a t s o f penaeid shrimp. Proc. Gu l f Caribb. Fish. I n s t . 19:86-98.

Music, J.L. 1979. Assessment of Georgia's shrimp and crab resources. Ga. Dep. Nat 1. Resour. Contr ib . Ser. 30:l-75.

Odum, W.E. 1971. Pathways o f energy f low i n a south F l o r i d a estuary. U n i v e r s i t y o f Miami Ph.0. D i sse r ta t i on . D i sse r t . Abst. Univ. Mich. 70-18, 156:1898B.

Overstreet, R.M. 1978. Marine ma1 adi es? Worms, germs, and o ther symbionts f rom t h e nor thern Gu l f o f Mexico. M i s s i s s i p p i - Alabama Grant Consort i um MASGP-78-021. 104 pp.

Pearson, J.C. 1939. The e a r l y l i f e h i s t o r i e s o f some American Penaeidae, c h i e f l y t h e commercial shrimp Penaeus s e t i f e r u s (L inn.) . U.S. B u ~ . B u l l . 49:l-73.

~ 6 r e z - ~ a r f ante, I. 1969. Western At1 a n t i c shrimp; o f t h e genus Penaeus. U.S. F i sh W i l d l . Serv. r i s h . u l 1 . 67(3) ~461-591.

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Shipman, S. 1983a. Survey o f Georgia 's major mar ine resources. Pages 1-154 (Chapter I ) i n S. Shipman, V. Baisden, a x H. Ashley. Studies and assessment o f Georgi a ' s mar ine f i s h e r i e s resources 1977-1981. Ga. Dep. Nat. Resour. Completion Rep. P.L 88-309 P r o j . 2-319-R. 503 pp.

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SO1 73 -101

REPORT DOCUMENTATION .IL"~-* NO-

PAGE FWS/OBS-82/11.27* I I num"c* -*- *- L m). o d LUC(~IO

Species. P r o f i l e s : L i f e H i s t o r i e s and Environmental Requirements o f Coastal Fishes and Inver tebra tes (South A t l a n t i c ) -- White

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Shrimp 7. ~ ~ a l

Robert J. Muncv

P.O. rawer BX '

M i s s i s s i p p i State U n i v e r s i t y M iss i ss i pp i State, MS 39762-5603

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8% I ~ s o d r c Owew1acnm kame DM 16atas* Nat iona l Coastal Ecosystems Team U.S. Army Corps of Engi neers F i sh and W i l d l i f e Service Waterways Experiment S ta t i on

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

U.S. Department o f the 1 n t e r i o r P.O. B&X 631 Uashington, DL 20240 Vicksburg, MS 39180

I & *b.tnc( (Urnit r00 ' Species p ro f i l es are l i t e r a t u r e sumnaries o f the taxonomy, morphology, range, l i f e h i s to ry , and environmental requirements o f aquat ic species. They are prepared t o a s s i s t i n environmental impact assessment. The wh i te shrimp, Penaeus set i ferus, i s the most impor tan t comnercial species i n t h e Southeastern Uni ted States. It serves an important eco log i ca l r o l e as food f o r o ther l a rge i nveceb ra tes and f ishes. Major b a i t i ndus t r y i s i n nor theast F l o r i d a and Georgia. .Spawning occurs of fshore w i t h i n 9-m depth contour where s a l i n i t i e s are a t l e a s t 27 ppt. I n spring, pos t l a r va l shrimp move w i t h t i d a l cur ren ts i n t o inshore es tuar ine waters. Juveni le wni t e shrimp p re fe r shal low organic- r i c h substrate w i t h low s a l i n i t i e s (1-10 pp t ) . Nearshore s o f t sediment areas co r re l a ted w e l l w i t h whi te and brown shrimp d i s t r i bu t i ons . Water temperature in f luences spawning, growth, .hab i ta t se lec t ion , emigrat ion, and mor ta l i t y . Low w in te r temperatures have g r e a t l y a f fec ted surv iva l , recrui tment, and harvest i n the South A t l a n t i c f i she ry . Ma in ta in ing su i t ab le nursery grounds i s a major concern f o r the f u tu re o f t he f i shery .

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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

REGION 5 REGION 6 Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service One Gateway Center P.O. Box 25486 Newton Corner, Massachusetts 02158 Denver Federal Center

Denver, Colorado 80225

REGION 7 Regional Director U.S. Fish and Wildlife Service 101 1 E. Tudor Road Anchorage, Alaska 99503

DEPARTMENT OF THE INTERIOR U.S. FISH A I D WILDLIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has respon- sibility for most of our.nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water 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.