REFEJtEPdCE COPY Do Not Remove from the Library

U. S. Fish and Wildlife Service

Biological Report 82 (1 1-34) National wetiarld3 -n"r

April 1985 700 Cajun Dome Boulevard TR EL.82-4 Lafayette, Louisiana 70506

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

STRIPED MULLET

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers

T h i s i s one o f t h e f i r s t r e p o r t s t o be p u b l i s h e d i n t h e new " B i o l o g i c a l R e p o r t " s e r i e s . T h i s t e c h n i c a l r e p o r t s e r i e s , p u b l i shed b y t h e Research and Deve lopment b r a n c h o f t h e U.S. F i s h and W i l d l i f e S e r v i c e , r e p l a c e s t h e "FWS/OBS1' s e r i e s p u b l i s h e d f r o m 1976 t o September 1984. The B i o l o g - i c a l R e p o r t s e r i e s i s d e s i g n e d f o r t h e r a p i d p u b l i c a t i o n o f r e p o r t s w i t h an a p p l i c a t i o n o r i e n t a t i o n , and i t c o n t i n u e s t h e f o c u s o f t h e FWS/OBS s e r i e s on r e s o u r c e management i s s u e s and f i s h and w i l d l i f e needs.

B i o l o g i c a l Report 82(11.34) TR EL-82-4 A p r i l 1985

Species P r o f i l e s : L i f e H i s t o r i e s and Environmental Requirements o f Coastal F ishes and I nve r teb ra tes (South F l o r i d a )

STRIPED MLILLET

Mark R. C o l l i n s Department of Zoo1 ogy U n i v e r s i t y o f F l o r i d a

Ga inesv i l l e , FL 32611

P r o j e c t Manager L a r r y Shanks

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 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

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 D i v i s i o n o f B i o l o g i c a l Serv ices

Research and Development F i s h and Wild1 i f e Serv ice

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

Th i s s e r i e s should be referenced as f o l l o w s :

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 environmental requi rements of coasta'i- f i shes and i nve r t eb ra tes . U. S. F i s h W i l d l . Serv. B i o l . Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

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

C o l l i n s , M. R. 1985. Species p r o f i l e s : 1 i f e h i s t o r i e s and envi ronmenta l requirements 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 F l o r i d a ) - - s t r i p e d mu l l e t . U.S. F i s h W i l d l . Serv. B i o l . Rep. 82(11.34). U.S. Army Corps o f Engineers, TR EL-82-4. 11 pp.

PREFACE

This species p r o f i l e i s one o f a ser ies on coasta l aquat ic organisms, p r i n c i p a l l y f i s h , o f spor t , commercial , o r eco log ica l importance. The p r o f i l e s are designed t o 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 csnprehensive sketch o f the b i o l og ica l c h a r a c t e r i s t i c s and environmental requ i re - m n t s o f the species and t o descr ibe how populat ions o f the species may be e q e c t e d t o reac t t o environmental changes caused by coasta l development. Each p r o f i l e has sect ions on taxonomy, 1 i f e h i s t o r y , eco log ica l r o l e , environmental requirements , and economic importance, i f appl i cab l e. A t h ree - r i ng b inder i s used f o r t h i s ser ies so t h a t new p r o f i l e s can be added as they are prepared. 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 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 the f o l 1 owing addresses.

I n f o m a t i o n Transfer Special i s t Nat ional Coastal Ecosys tems Team U.S. F ish and W i l d l i f e Serv ice NASA-Sl i d e l 1 Computer Compl ex 1010 Gause Boulevard Sl i d e l 1 , LA 70458

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

CONVERSION TABLE

m i l 1 ime te rs (mn) cent imeters ( an) meters (m) k i 1 m e t e r s ( km)

2 square meters (m ) square k i 1 m e t e r s ( km2) hec tares (ha)

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

m i l l igrams (mg) grams ( g ) k i l ograms ( k g ) 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

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

!Y To Obtain

inches inches fee t m i 1 es

square f e e t square m i l es acres

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

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

1.8(OC) + 32 Fahrenhei t degrees

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

inches 25.40 i nches 2.54 f e e 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 l es ( m i ) 1.852

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

ga l 1 ons ( g a l ) 3.785 cub ic f e e t ( f t 3 ) 0.02831 acre- f e e t 1233.0

ounces (02) 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 thermal u n i t s (B tu ) 0.2520

m i l 1 imeters centimeters meters meters k i l m e t e r s k i 1 m e t e r s

square meters hectares square k i lometers

1 i t e r s cub ic meters cubic meters

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

Fahrenhei t degrees 0.5556("F - 32) Cels ius degrees

CONTENTS

.................................................................. PREFACE ......................................................... CONVERSION TABLE

ACKNOWLEDGMENTS ..........................................................

.............................................. NOMENCLATLlRE/TAXONOMY/RANGE ........................................... MORPHOLOGY/IDENTIFICATION A I D S ........................................... REASON FOR I N C L U S I O N I N S E R I E S

. ............................................................ L I F E HISTORY S p a w n i n g ............................................................... L a r v a e and J u v e n i l e s . . ................................................. A d u l t s . ................................................................

................................................... GROWTH CHARACTERISTICS F I S H E R I E S ................................................................

S p o r t .................................................................. C o m m e r c i a l . ............................................................ M a n a g e m e n t . ............................................................

ECOLOGICAL ROLE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................................... ENVIRONMENTAL REQUIREMENTS

............................................................ T e m p e r a t u r e S a l i n i t y . ..............................................................

....................................................... D i s s o l v e d O x y g e n .................................................................. D e p t h

. ........................................................ LITERATURE C I T E D

Page

i i i i v v i

1 1 3 3 3 3 4 4 5 5 5 6 6 7 7 8 8 8

9

ACKNOWLEDGMENTS

I am g r a t e f u l f o r t he rev iew o f the manuscript by J. Shireman o f t he Univer- s i t y o f F lo r i da .

Figure 1. S t r i ped mu l le t .

STRIPED MULLET

S c i e n t i f i c name . . Mugil cephalus L i nnaeus 1758

Pre fe r red common name . . . s t r i p e d mu1 l e t (F igure 1 )

Other common names . . . grey mu1 l e t , b lack mu1 l e t

Class . . . . . . . . . . Osteichthyes Order . . . . . . . . . . Perciformes Family . . . . . . . . . . . Mugi 1 i dae

Geographic Range: Coastal waters o f a1 1 seas, roughly between 42' N and 42' S. I n the western A t l a n t i c from B r a z i l t o Nova Scot ia (Hoese and Moore 1977). Abundant along the south F l o r i d a - coast (Figure 2).

MORPHOLOGY/IDENTIFICATION AIDS

Dorsal f i n I V + I spines, 8 s o f t rays; anal f i n I 1 1 spines, 8 rays (11, 9 i n j uven i l es ) ; pec tora l f i n 16-17 rays; caudal f i n 18-20 rays; l a t e r a l

scale count 38-42. Anal and second dorsal f i n s a1 most scale less; o r i g i n o f second dorsal behind o r i g i n o f anal f in ; pec tora l f i n shor ter than d i s - tance from l a s t spine o f f i r s t dorsal t o o r i g i n o f second dorsal f i n ; head somewhat wider than deep; mouth te rmi - nal and small, t ee th inconspicuous; no 1 a t e r a l 1 i ne v i s i b l e, bu t 1 ongi t u d i nal dark s t r i p e s on the sides o f elongate body; body b l u i sh-gray d o r s a l l y and whi te v e n t r a l l y ; scales c y c l o i d i n young, feeb ly c teno id i n adul ts ; d i s - t i n c t a x i l l a r y scale a t pectoral f i n ; g i l l raker number increas ing w i t h size; prominent adipose eye1 i d w i t h on ly a narrow s l i t over the pup i l , cover ing the preorb i t a l a n t e r i o r l y and running tw ice as f a r behind the eye as i n f r o n t (Hoese and Moore 1977). The s i x Mugil species repor ted from south F l o r i d a sometimes are d i f f i c u l t t o separate taxonomical l y ; however Rivas (1980) constructed a key t h a t i s use- f u l f o r t e n t a t i v e i d e n t i f i c a t i o n . The two species most o f t e n encountered, s t r i p e d m u l l e t and whi te mul le t , may be separated by anal f i n ray counts o f 8 and 9, respect ive ly .

T. PETERSBURG

PALM BEACH

CAPE ROMAN

THOUSAND ISLAND

M I L E S

K I L O M E T E R S

F i g u r e 2. S t r i p e d rnu1:Iet a r e d i s t r i b u t e d a l o n g t h e e n t i r e c o a s t o f t h e Sou the rn Fl o r i da Region.

2

REASON FOR INCLUSION IN SERIES

S t r i ped mul le t , perhaps the most widespread and abundant inshore t e l e - o s t (Odum 1970), supports a valuable commercial f i s h e r y along the g u l f coast o f F l o r i d a (Rivas 1980). M u l l e t are primary consumers t h a t feed la rge- l y on r e l a t i v e l y minute l i v i n g and dead vegetable matter. They are an ecol og ica l l y important component i n the f low o f energy through es tuar ine communities. I n many p a r t s o f the wor ld s t r i p e d mu1 l e t are impor tant i n f i s h cu l tu re .

LIFE HISTORY

Spawning

Because o f the s c a r c i t y o f data on s t r i p e d mu1 1 e t spawning, specula- t i o n s on the l o c a t i o n s o f spawning grounds are based l a r g e l y on the ap- pearance o f larvae. M u l l e t have been repor ted t o spawn inshore (Breder 19401, a long beaches (Gunter 1945), 8 t o 32 km o f f sho re (Broadhead 1953), and i n water deeper than 40 m (Anderson 1958). I n the G u l f o f Mexico, mu1 1 e t were observed spawning 65 t o 80 km of fshore i n water 1,000 t o 1,800 m deep (Arnold and Thompson 1958). These observations i n d i c a t e t h a t m u l l e t spawn over a wide range o f coastal waters b u t the cu r ren t consen- sus i s t h a t most spawn of fshore.

The spawning season o f s t r i p e d mu1 1 e t usual l y begins i n October, peaks i n November-December, and ends i n February (Anderson 1958; Ri-vas 1980). Along F l o r i d a s west coast, s t r i p e d mu1 1 e t spawn from October through May a t Cedar Key and from December through J u l y a t Bayport ( K i l b y 1949). I n A u s t r a l i a some fe- males spawn only i n a l t e r n a t e years (Thomson 1955), and evidence from Louis iana (Shireman 1975) suggests t h i s may a1 so be t r u e f o r mu1 l e t i n U .S. waters. The estimated fecund i t y o f s t r i p e d m u l l e t i s 0.5 t o 2.0 m i l - l i o n eggs per female, depending on the

s i ze o f the female (Broadhead 1953). A fecundity value o f 648 f 62 eggslg body wei gh t was ca l cu la ted by Shehadeh e t a l . (1973).

Eggs are transparent, straw- colored, non-adhesive, spher ical , and w i thou t ex terna l marking. The eggs average 0.72 mn i n diameter, con ta in an o i l g lobu le 0.28 mn i n diameter, a re p o s i t i v e l y buoyant, and hatch about 48 h a f t e r f e r t i l i z a t i o n (Thomson 1963 1.

Larvae and Juveni 1 es

Larvae average 2.4 mn long ( a l l lengths i n t h i s r e p o r t are t o t a l 1 engths unl ess i nd i ca ted otherwise) a t hatching and have no mouth, pa i red f i n s , o r branchia l skeleton. A f t e r 5 days they are about 2.8 mn long, the jaws are we1 1 -defined, the i n t e r n a l organs become organized, and f i n ' buds have s t a r t e d developing (Thomson 1963). Morphological and. m e r i s t i c development and growth cont inue u n t i l the la rvae are about 16-20 nn standard l eng th (SL), a t which t ime they m i - g ra te t o inshore waters and es tua r ies ( K i l b y 1949; Anderson 1958). The m i - g ran t la rvae have two spines and n ine rays i n the anal f i n ( the "Querimana" stage) u n t i l they reach 35-45 mn SL, a t which t ime the f i r s t ray fuses i n t o a t h i r d spine and the adipose eye1 i d becomes apparent. The m u l l e t i s then considered a j uven i 1 e (Anderson 1958).

Juven i le s t r i p e d m l i l l e t 40 t o 69 mn SL reach a " d e f i n i t i v e s ta te " of osmoregul a to ry capabi 1 i ty . For exam- ple, they t o l e r a t e s a l i n i t i e s o f 0 t o 35 ppt, whereas they could n o t have prev ious ly surv ived i n freshwater. Juven i les spend the r e s t o f t h e i r f i r s t year o f l i f e i n coastal waters, s a l t marshes, and es tuar ies , and o f t e n move t o deeper water i n the f a l l when the adu l t s migra te o f fshore t o spawn, y e t l a r g e nunbers o f imnature m u l l e t overwinter i n estuar ies. A f t e r t he f i r s t year, m u l l e t i n h a b i t the sea, s a l t marshes, estuar ies, o r freshwater ri vers (Nordl i e e t a1 . 1982). Appar-

ent ly in some instances, females great ly outnumber the males i n brack- i sh and freshwater hab i ta t s (Shi reman 1975; Col 1 ins 1981 1.

Adul ts

The length a t which mullet mature ranges from about 20G t o 300 m SL; the females mature a t a s l i g h t l y l a rg - e r s i z e than the males. A1 though some mature i n t h e i r second year of l i f e , most mature i n t h e i r t h i r d year (Broadhead 1953, 1958; R i vas 1980 ) . On the northern gulf coast of Florida, the growth r a t e and s i z e a t maturity of stri ped mu1 1 e t increased progres- s ive ly from west t o eas t . Mullet may 1 i ve 4 years o r more (Rivas 1980) ; the maximum age reported i s 13 years (Thomson 1963).

Mature mullet migrate offshore t o spawh i n the f a l l and winter, often in large schools. Mullet t h a t mature in freshwater e i t h e r migrate t o the sea t o spawn or resorb t h e i r gonads (Shi reman 1975). Swi m i ng speed dur- ing migration is much greater than t h a t predicted t o be energet ical ly optimal, probably because of the in- creased hydromechanical ef f ic iency provided by school ing and the selec- t i ve force of heavy predation duri ng spawning migrations (Peterson 1976). Fishing f o r mullet i s heaviest during spawning mi g ra t i ons when school s a re large , the condition of the f i s h i s excel l e n t , and the prized roe (gonads) a re l a rges t .

In a mullet tagging study, 2,779 recoveries from 12,647 tagged mu1 1 e t revealed t h a t 91% were recovered with- in 32 km of the re lease point; only 2% were recovered more than 160 km away. These r e s u l t s together with other tagging s tudies and meris t ic analyses (Broadhead 1953; Broadhead and Mefford 1956; De Sylva e t a l . 1956), indicate t h a t Florida gulf coast mullet are divided i n t o three 1 oosely k n i t populations, the south Florida population extending northward t o the Steinhatchee area. East coast

mu1 l e t commonly mi gra te much fu r the r (up t o 560 k m ) , usually in a southerly d i rec t ion , and a re considered a s ing le population. Fish returning from spawning migrations on the gulf coast usually have l e s s f a t reserves and more eroded f i n s and les ions on the body than do p rem ig ra to r y mu1 l e t (M. R . C o l l i n s , U n i v e r s i t y o f F l o r i d a , G a i n e s v i l l e ; unpubl ished data) .

GROWTH CHARACTERISTICS

Growth r a t e s of stri ped mu1 1 e t along the gulf coast of Florida in- crease from west t o e a s t along the panhandle and t o the south along the peninsula, possibly due t o a s l i g h t increase in coastal water temperatures (Broadhead 1958). In a tagging study, Broadhead (1953) found t h a t growth during spring and summer i s twice t h a t of f a l l and winter: 7.3 i. 1.8 mm/quarter f o r the f i r s t and fourth quar ters of the year , 16.4 i. 2.7 mm/quarter f o r t h e second, and 19.1 f 2.7 m / q u a r t e r f o r the th i rd . The mean fork lengths of mu1 l e t from four gulf coast areas fo r each year of l i f e a re given in Table 1.

Table 1. Mean f o r k l eng ths (mm) o f s t r i p e d m u l l e t a t va r i ous ages from f o u r g u l f coas t areas o f F l o r i d a as c a l c u l a t e d from sca le a n a l y s i s (Broadhead 1958).

Year o f 1 i f e

Area 1 2 3 4

Pensacola 142 207 263 --

Apa lach ico la 134 207 271 --

Cedar Key 175 258 307 --

Homosassa 178 269 319 366

Females are l a r g e r and grow s l i g h t l y f a s t e r than males o f the same age (Broadhead 1958). I n the Cedar Key and Crys ta l R ive r areas, m u l l e t longer than 28 cm SL are predominantly females ( C o l l i n s 1981). I n the Gu l f o f Mexico, the growth i n l eng th o f mu1 l e t g radua l ly slows as the f i s h become la rger , and reaches an asymp- t o t e a t an average length o f 600 inn TL, a t probably 5-6 years o f age (Rivas 1980). Length-weight r e l a t i o n - ships f o r s t r i p e d m u l l e t i n the g u l f a re given i n F igure 3.

FISHERIES

Spor t

S t r i ped mu1 l e t along the F l o r i d a g u l f coast a re used as b a i t f o r a wide v a r i e t y of f i shes and are regarded as an e x c e l l e n t food f i s h . I n f r e s h and brack ish waters, mu1 l e t a re sometimes caught w i t h hook and l i n e . Earthworms may be used f o r b a i t , b u t oatmeal and chicken l a y i n g mash are more popu- l a r . M u l l e t from freshwater o f t e n have a muddy o r hydrogen s u l f i d e - l i k e

L L N O T H I N I N C H E S I I . 10 1 1 8 , I, I. 1 1 <, L I I I I I I I I I

1'" I. BOTH SEXES NOT SPAWNIN0 I , I

2. FEMALES SPAWNlNO

I. MALE SPAWNING

B O O "'t , I:::;

l ~ ~ ~ ~ x ~ ~ ~ ~ ~ . ~ ~ ~ ~ ~ . ~ ~ ~ . ~ ~ . . l " " " - " " " " " " " " " " " " . . " - " " ". 'zcP.;:::.:;:~"-"... "" ' " " I . . . . " " . " " .

L E N G T H I N M U .

Figure 3. Length-weight re1 a t ionsh ips o f s t r i p e d m u l l e t from F l o r i d a ' s g u l f coast i n 1951 (Broadhead 1953).

taste. I n brack ish and sa l twater areas, mu1 l e t are sometimes snagged w i t h t r e b l e hooks, bu t most o f t e n some type o f n e t i s used. Sport fishermen usua l l y use a cas t n e t o r seine, b u t some use sho r t lengths o f g i l l nets o r t r a n e l nets.

Coinnercial

I n general, spor t and commercial mu1 l e t fishermen employ the same types o f nets, b u t commercial seines, g i l l nets, and trammel nets are usua l l y much longer. Seines are now used more on the east coast than on the west coast because o f the abundance o f smooth bottoms near sandy beaches on the east coast. Nets w i t h l a r g e s t meshes are used i n the f a l l when mul- l e t are g rav id and f a t . Many g i l l ne ts and trammel nets are longer than 500 m and are usua l l y s e t from a boat. When m u l l e t are i n open water e i t h e r n e t i s s e t t o completely enc i r - c l e the school; when the school i s near a shore l ine the n e t i s se t i n a h a l f - c i r c l e from shore t o shore; and i n i n t e r t i d a l coves o r creeks the nets are used t o completely block o f f an area, ca tch ing the m u l l e t as they leave w i t h the ebb t i de . F i s h i n g may be c a r r i e d ou t day o r n igh t , and some- times two boats work together, combining t h e i r nets and e n c i r c l i n g a school from opposi te d i r e c t i ons. When seines o r nets are used, many m u l l e t escape by jumping over the f l o a t l i n e and, i n the case o f g i l l and t r a n e l nets, they may avoid the net. Small m u l l e t escape through the mesh.

The commercial landings o f mu1 l e t were no t repor ted by species u n t i l 1958. Since then, F l o r i d a has pro- duced 80% t o 90% o f the U.S. s t r i p e d m u l l e t catch from the Gul f of Mexico (Rivas 1980). The annual F l o r i d a catch o f s t r i p e d mu1 1 e t i n 1958-1981 f l u c t u a t e d from a high o f 35 m i l l i o n I b i n 1964 t o a low o f 1 6 . 8 m i l l i o n 1b i n 1976 (Table 2) . Gul f coast land- ings con t r i bu ted about 80% o f t he catch. M u l l e t a lso are important b a i t f i s h , espec ia l l y i n the s p o r t

Table 2. Annual ca t ch o f s t r i p e d mu1 l e t ( m i 11 ions o f pounds) i n F l o r i - da, 1958-81. Data compiled from annual ca tch s t a t i s t i c s , Nat iona l Marine F i she r i es Serv ice, Washington, D. C.

Year Catch Year Catch Year Catch

ly, minimum lengths o f 9 t o 11 inches FL (depending on l o c a l ordinances) are the on ly regu la t i ons (Rivas 1980). A1 though 1 andi ngs and f i s h i n g i ntensi - ty f o r s t r i p e d m u l l e t decl ined from 1964 t o 1977, ca tch -pe r -un i t -o f -e f fo r t and the average s i ze o f the f i s h re - mained unchanged. This r e l a t i o n s h i p suggests t h a t the l ack o f demand r a t h e r than a s c a r c i t y o f m u l l e t l i m - i t s the f i s h i n g (Rivas 1980).

ECOLOGICAL ROLE

f i s h e r y f o r b i l l f i sh. They commonly b r i n g a h igher p r i c e as b a i t than as food f i s h .

Most (65%) s t r i p e d m u l l e t are marketed i n the Southeast Un i ted States. The remainder are marketed i n 1 arge northeastern, m i dwestern, and Cal i f o r n i a c i t i e s . O f the t o t a l catch, about 25% i s marketed fresh, 63% frozen i n the round, 6% smoked, and 6% as roe. Market ing i n o the r forms has been attempted, bu t f a i l e d because mu1 1 e t t u r n ranc id w i t h i n 60 days and consumer acceptance i s poor outs ide o f the southeast (Cato e t a l . 1976; Rivas 1980).

Management

Closed seasons and o ther regula- t i o n s have been used t o manage the F l o r i d a m u l l e t i n the past, b u t most have s i nce been e l i m i nated. Current-

There has been some discrepancy concerning the d i e t o f j u v e n i l e s t r i p e d m u l l e t bu t most authors now agree t h a t l a r v a l m u l l e t p r i m a r i l y ea t microcrustaceans (De S i l va 1980). The f a c t t h a t 1 arvae, successfu l ly reared t o 20 mn SL, were fed e n t i r e l y on animal mater ia l i 11 us t ra tes the depen- dence o f l a r v a l and p o s t l a r v a l mu1 l e t on zooplankton (Nash e t a l . 1974). I n F l o r i d a ' s Ind ian R iver lagoon the stomach contents o f near ly 400 s t r i ped m u l l e t la rvae up t o 35 mn SL were examined. Larvae up t o 15 mn SL f e d almost e n t i r e l y on copepods (70%) and mosquito l a rvae (30%); those from 15 t o 25 mn SL fed on copepods (SO%), mosquito l a rvae (1581, and p l a n t de- b r i s (35%); and la rvae from 25 t o 35 mn SL fed p r i m a r i l y on p l a n t debr is (80%) and copepods (10%) (Har r i ngton and H a r r i ngton 1961 1. The p ropor t i on o f sand and d e t r i t u s i n the gut o f j u v e n i l e s increases w i t h length, i n d i - c a t i n g t h a t they tend t o take more food from the bottom as they grow o l d e r (De S i l v a and Wijeyaratne 1977). The exceptions are j u v e n i l e s t h a t feed from the water column o r surface when s u i t a b l e foods are plen- t i f u l there. For example, i n one study m u l l e t 35-80 mn long fed on a b l oom o f the d i no f 1 age1 1 a t e Krypto- per id in ium sp. (Odum 1968). The r a t e o f d iges t fon o r r e t e n t i o n t ime o f food i n j u v e n i l e s var ies w i t h both s a l i n i t y and body weight. I n S r i Lanka, a study o f j u v e n i l e mu1 l e t 10 t o 50 mn long showed t h a t d iges t i on r a t e i n - creases w i t h body weight and t h a t both

d iges t i on and inges t i on ra tes i n - creased w i t h s a l i n i t y from 1 pp t t o 30 pp t (Perera and De S i l v a 1978). Food evacuation t imes ranged from about th ree t o s i x hours.

Adu l t s t r i p e d mu1 l e t have been described as herbivorous, d e t r i - t i vorous, and " i n te r face" feeders. The d i e t and feeding behavior of m u l l e t may vary by l oca t i on , bu t t h e i r major food i s e i t h e r ep iphy t i c and benth ic microal gae, macrophyte d e t r i - tus, o r inorgan ic sediment p a r t i - c les. A1 though sediment p a r t i c l e s f u n c t i o n as a g r ind ing paste i n the g i zza rd - l i ke p y l o r i c p o r t i o n o f the stomach, some small p a r t i c l e s are r i c h i n adsorbed micro-organi sms and are s e l e c t i v e l y ingested f o r t h e i r food va l ue (Odum 1970 ) .

M u l l e t commonly feed by sucking up the top l a y e r o f sediment, which i s r i c h i n d e t r i t u s and microalgae, p r i - m a r i l y diatoms i n the Cedar Key area ( C o l l i n s 19811, and by grazing on epiphytes and ep i fauna from seagrasses and o ther substrates. They a l so i n - gest sur face scum when l a r g e concen- t r a t i o n s o f microalgae are present a t the a i r -water i n t e r f a c e (Odum 1970), and feed on swarming polychaetes (Nereis succinea) i n the water column

and MigTarese 1978). I n some freshwater h a b i t a t s s t r i p e d mu1 1 e t feed p r i m a r i l y on the epiphytes and epifauna o f aquat ic macrophytes and on benth ic f i lamentous green algae (Co l l i n s 1981 1, b u t they a l so i nges t sediment f o r tri t u r a t i o n (g r i nd ing ) . The t ime o f most in tense feeding ap- pa ren t l y var ies w i t h locat ion . I n a l l F l o r i d a h a b i t a t s s tud ied by Odum (1970) feeding va r ied w i t h the he ight o f the t i de , whereas i n sa l twater (Cedar Key) and freshwater (Crys ta l R ive r ) s i t e s s tud ied by C o l l i n s (1981) feeding was s t r i c t l y d iu rna l and unre- l a t e d t o t i d a l stage.

The major predators o f j u v e n i l e and a d u l t m u l l e t are f i shes and b i r d s (Thomson 1963). I n F lo r i da , sharks sometimes feed heav i l y on l a r g e mul-

l e t . Mu1 l e t up t o 35 cm long are fed on by spot ted seat rout (Cynoscion nebulosus) (Breuer 1957). A1 thougT the d i e t o f s t r i p e d m u l l e t over laps those of a v a r i e t y o f aquat ic species, compet i t ion has n o t been documented.

P a r a s i t i c i n f e c t i o n s and i n f e s t a - t i o n s are common i n mu1 l e t . O f near ly 300 a d u l t mu1 l e t from freshwater and sa l twa te r s i t e s on F l o r i d a ' s g u l f coast, no i n d i v i d u a l s were found w i th - ou t paras i tes on e i t h e r the g i l l s o r body surface (M. R. Co l l i ns , Univ. o f Fla., Gainesvi l l e ; unpubl i shed data). S t r i ped m u l l e t serve as de- f i n i ti ve o r intermediate hosts f o r many paras i tes i n c l u d i n g f l age1 1 ates, c i 1 i a tes , myxospori d i ans, monogenean and d i genean trematodes, nematodes, acanthocephalans, leeches, argu l ids , copepods, and isopods (Paperna 1975; Overs t reet 1978; Paperna and Over- s t r e e t 1981 ). Extensive paras i te - induced m o r t a l i t i e s o f mu1 l e t i n the w i l d have no t apparent ly been repo r t - ed.

ENVIRONMENTAL REQUIREMENTS

Temperature

An ana lys i s o f the worldwide d i s t r i b u t i o n o f s t r i p e d mu1 l e t sug- gests t h a t m u l l e t are n o t permanent r e s i dents i n waters w i t h temperatures below 1 6 O ~ o r where water temperatures f a i l t o reach 18O~. On F l o r i d a ' s g u l f coast, young s t r i p e d m u l l e t l i v e i n s a l t marsh pool s a t temperatures from 1 3 O ~ t o 34.5'~ ( K i l b y 1949). Water temperatures probably regu la te the l eng th o f t ime t h a t young m u l l e t s tay i n estuar ies. M u l l e t l e s s than 50 mn SL p re fe r temperatures from 30.0° t o 32.5O~ and mu1 l e t from 50 t o 130 mn SL p r e f e r temperatures from 19.5~ t o 2 0 . 0 ~ ~ . For a l l s izes o f f i s h the temperature se lec ted tends t o decrease as s a l i n i t y increases. I n experiments w i t h Hawaiian s t r i p e d mu1 l e t 78-122 mn SL, the upper l e t h a l tem e ra tu re was oP between 2 7 . 0 ~ and 29.6 C f o r f i s h accl imated a t 21°c i n f reshwater and

the lower l e t h a l tem e ra tu re was be- g tween 1 0 . 2 ~ and 15.0 C (Sy lves ter e t a1 . 1974). The upper 1 e tha l tem era- I t u r e f o r m u l l e t accl imated a t 26 C i n sa l twater (32 p p t ) ranged from 29.0° t o 33.0°c, and the lower l e t h a l temperature ranged from 1 0 . 4 ~ t o 14. OOc. The c r i t i c a l thermal maximum (CTM) o f Hawaiian m u l l e t (70 t o 125 mn SL) i s a f fec ted both by acc l imat ion temperature and by t ime o f day (Syl ves ter 1975). Mean CTM increased from 38.5O t o 41.3O~ f o r f i s h a c c l i - mated a t 20.0' and 29.0°c, respec- t i v e l y , and mean CTM i s about 0.5O~ higher a t midday than i n morning o r evening. C r i t i c a l thermal maxima and minima were no t ava i l ab le f o r a d u l t mu l l e t .

Sal i n i ty

Adu l t m u l l e t have been repor ted from waters w i t h a s a l i n i t y ranging from 0 p p t ( C o l l i n s 1981) t o 75 p p t (Simmons 1957). Using f i s h induced t o spawn i n the laboratory, Sy lves ter e t a1 . (1975) found t h a t egg su rv i va l was h ighest a t a s a l i n i t y o f 32 p p t ( t he h i ghest sal i n i ty tested), whereas the g r e a t e s t s u r v i v a l o f l a r v a e was a t 26 p p t i n t e s t s f rom 24 t o 36 p p t . When j u v e n i l e m u l l e t are 40-70 mn SL they reach a d e f i n i t i v e s t a t e o f osmoregu- 1 a to ry capabi 1 i ty and can to1 e ra te s a l i n i t i e s from freshwater t o f u l l seawater (Nordl i e e t a1 . 1982).

D i ssol ved Oxygen

Mu1 l e t 1 arvae apparent ly cannot surv ive i n d issolved oxygen (DO) con- c e n t r a t i o n s below 4 ppm (Sy lves ter e t a l . 1975). Over a range o f 1.0 t o 8.0 ppm DO, eggs incubated i n the labora- t o r y f o r 48 h had a s u r v i v a l r a t e o f 0%-3% a t concentrat ions 4.5 ppm and below, and 851-901 f o r 5.0 ppm and

above. Larvae were he ld from one t o fou r days i n concentrat ions o f 4.0-7.9 ppm DO, and those h e l d f o r 4 days had a mean su rv i va l o f 0-8s a t 4.0-5.4 ppm, 21% a t 6.4 ppm, and 84% a t 7.9 ppm. Although 7.9 ppm i n t h i s i n - stance i s 146% satura t ion , there was no evidence o f gas bubble disease. No s p e c i f i c data on oxygen requ i rements were found f o r a d u l t mul le t , b u t pre- 1 iminary experiments w i t h caged f i s h show t h a t they surv ive a t an oxygen concentrat ion o f 4.4 ppm DO a t 2 9 ' ~ and a s a l i n i t y o f 28 p p t (M. R. Co l l i ns , Univ. o f Fla., Ga inesv i l le ; unpublished data). I n a study o f schools o f m ig ra t i ng mul le t , DO was 29% lower a t the t r a i l i n g than the 1 eadi ng edges, suggesting t h a t the cont inua l breaking up and reforming o f the school and changing o f p o s i t i o n w i t h i n the school a1 low f i s h a t the center and t r a i l i n g edge access t o water h igher i n DO (McFarland and Moss 1967).

M u l l e t l i v e i n a wide range o f habi t a t s and depths and spawn p r imar i - l y i n r e l a t i v e l y deep cool coastal waters. The la rvae move inshore t o shal low waters along beaches and i n s a l t marshes. I n Hawaii, schools o f m u l l e t l e s s than 50 mn SL were always found i n waters o f minimal depth, i n c l u d i n g the swash zone and t i d e pool s, despi te near - le tha l tempera- tu res (Major 1978). Juven i les greater than 50 mn SL p r e f e r s l i g h t l y deeper waters beyond the swash zone, b u t dur ing f l o o d t i d e s they may move i n t o shallow waters vacated by smal ler mul le t . The preference o f f i s h l e s s than 50 mn SL f o r extremely shallow water apparent ly permits them t o es- cape most predators and t o feed w i th - ou t ser ious compet i t ion (Major 1978).

LITERATURE CITED

Anderson, W. W. 1958. Larval devel- opment, growth, and spawning o f s t r i p e d m u l l e t (Mu il cephalus) along the south A t + a n t i c coast o f the Un i ted States. U.S. F i s h Wi ld l . Serv. Fish. B u l l . 58:501- 519.

~ r n o i d , E. L., and J. R. Thompson. 1958. Offshore spawning of the s t r i p e d m u l l e t Mu il ce halus i n the Gulf of &o%peia 1958; 130-132.

Bishop, J. M a y and J. V. Miglarese. 1978. Carnivorous feeding i n a d u l t s t r i p e d mu1 1 e t . Copei a 1978:705-707.

Breder. C. M.. J r . 1940. The spawn- i n g o f ~ u g i l ce halus on the ~l or- + ida west coast o ~ e i a 1940:138-

Breuer, J. P. 1957. Ecological sur- vey o f B a f f i n and Alazan Bays, Texas. Publ. I ns t . Mar. Sci. Univ. Tex. 4:135-155.

Broadhead, G. C. 1953. Invest iga- t i o n s o f the black mu1 l e t , Mu il

t l o r i d a . Fla. I+ cephalus L., i n n o r t west

Board Conserv. Tech. Ser. No. 7. 33 pp.

Broadhead, G. C. 1958. Growth o f the b lack mul le t , (Mu il cephalus L.) i n west and n o r t west k l o r i da. Fla.

+i Board Conserv. Tech. Ser.

No. 25. 29 pp.

Broadhead, G. C., and H. P. Mefford. 1956. The m i g ra t i on and expl o i ta - t i o n o f the b lack mul let , Mugil

ce halus L. i n F lo r i da , as deter- k rom tagging dur ing 1949- 1953. Fla. Board Conserv. Tech. Ser. No. 18. 31 pp.

Cato, J. C., P. B. Youngberg, and R. Raulerson. 1976. Production, pr ices, and marketing: an econom- i c ana lys is of the F l o r i d a m u l l e t f ishery. Pages 10-62 i n J. C. Cato and W. E. M c M u l l o ~ h , eds. Economics, b io logy , and food techno1 ogy o f mu l l e t . F la. Sea Grant Program Rep. No. 15.

Co l l i ns , M. R. 1981. 'The feeding p e r i o d i c i t y of s t r i p e d mu1 l e t , Mugil cephalus L., i n two F l o r i d a hab i ta ts . J. F i s h B i o l . 19:307- 315.

De S i lva , S. S. 1980. B io logy of j u v e n i l e grey mu l le t : a sho r t review. Aquaculture 19:21-36.

De S i lva , S. S., and M. J. S. Wijeyaratne. 1977. Studies on the b io logy of young grey mul le t , Mu il ce halus L. 11. Food and &hng*cul t u r e 12: 157-167.

De Sylva, D. P., H. B. Stearns, and D. C. Tabb. 1956. Populat ions o f the black m u l l e t (Mu il ce halus L.) i n F lo r i da . &+kia Conserv. Tech. Ser. No. 19. 45 PP*

Gunter, G. 1945. Studies on marine f i shes o f Texas. Publ. I ns t . Mar. Sci. Univ. Tex. 1:l-190.

Harr ington, R. W., and E. S. Harr ing- ton. 1961. Food se lec t i on among

f i shes invading a h i gh subt rop ica l s a l t marsh: from onset o f f lood- i n g through the progress o f a mosqui t o brood. Ecol ogy 42:646- 666.

Hoese, H. D., and R. H. Moore. 1977. Fishes o f the Gul f o f Mexico-Texas, Louisiana, and adja- cent waters. Texas A&M U n i v e r s i t y Press, College Sta t ion . 327 pp.

K i lby , J. D. 1949. A p re l im ina ry r e p o r t on the young s t r i ~ e d m u l l e t ~ u g i 1 cephal us L: i n ' two g u l f coastal areas o f F lo r i da . 0. J. F la . Acad. Sci . 11:7-23.

Major, P. F. 1978. Aspects o f estua- r i n e i n t e r t i d a l ecology o f juve- n i l e s t r i ~ e d mul le t . Muail c e ~ h a - lus , i n ~ a w a i i . U:S.N~~~F#&K FiSh. Serv. Fish. B u l l . 76:299- 313.

McFarland, W. N., and S. A. Moss. 1967. I n t e r n a l behavior i n f i s h school s. Science 156:260-262.

Nash, C. E., C. M. Kuo, and S. C. McConnel . 1974. Operational

Overstreet , R. M. 1978. Marine mala- d ies? Worms, germs, and o ther symbionts from t h e nor thern Gu l f of Mexico. Miss.-Ala. Sea Grant Consort. Publ . MASGP-78-021. 140 PP.

Paperna, I. 1975. Paras i tes and diseases of t h e grey m u l l e t (Mugi 1 idae) w i t h specia l reference t o t h e seas o f t he Near East. Aquaculture 5: 65-80.

Paperna, I., and R. M. Overstreet . 1981. Paras i tes and diseases o f mu l l e t s (Mugil idae). Pages 411- 493 i n 0. H. Oren, ed. Aquacul- t u r e o f grey mu1 1 ets. In terna- ti onal B i o l o g i c a l Program 26. Cambridge U n i v e r s i t y Press, Cambridge, England.

Perera, P. A. B., and S. S. De S i l va . 1978. Studies on the b io logy o f young grey m u l l e t (Mu il ce halus) d igest ion. Mar. B+~&T=JB).

Peterson, C. H. 1976. Cru is ing speed dur ing m ig ra t i on o f t he s t r i p e d m u l l e t Mugil cephalus: an evolu- ti onary response-To predat ion. Evo lu t i on 30: 393-396.

procedures fo r r e a r i n g . larvae o f the grey m u l l e t (Mu il cephalus Rivas, L. R. 1980. Synopsis o f know- . Aquaculture 3:&. 1 edge on the taxonomy, b io logy ,

d i s t r i b u t i o n . and f i s h e r v o f the

Nord l ie , F. G., W. A. Sze l is towsk i , and W. C. Nord l ie . 1982. Onto- penesi s o f osmotic resu l a t i o n i n t h e s t r i p e d mu1 l e t Mu -il ce halus L. J. F ish . B i o l . ' 2 & b E - -

Odum, W. E. 1968. Mu1 l e t grazing on a d i n o f l age1 1 a te bloom. Chesa- peake Sci . 9:202-204.

Odum, W. E. 1970. U t i l i z a t i o n o f the d i r e c t grazing and p l a n t d e t r i t u s food chains by the s t r i ~ e d mu1 l e t Mu il ce ha1 u; Pages 222-240 i n A. 78Z'E- ed. Marine f o x chains. 01i;er and Boyd, Ltd., Edinburgh, Scot1 and.

Gu l f o f ~ e x i c o mu1 1 e t s " (Pisces: Mugil idae). Pages 34-53 - i n M. F landor fer and L. Skupien, eds. Proceedings o f a workshop f o r p o t e n t i a l f i s h e r y resources o f the northern Gul f o f Mexico. Miss.- Ala. Sea Grant Consort. Publ. MASGP-80-012.

Shehadeh, Z. H., C. M. Kuo, and K. K. M i l i sen. 1973. Induced spawning o f grey m u l l e t Mu il ce halus L. w i t h f r a c t i o n a t e a+- sa mon - p i t u i t a r y ex t rac t . J . F ish B i o l . 5:471-478.

Shireman, J. V. 1975. Gonadal devel- opment o f s t r i p e d mu1 l e t (Mugil

ce halus) i n f resh water. Prog. h u t. 37~205-208.

Simnons, E. G. 1957. Ecological survey of the upper Laguna Madre o f Texas. Publ. I n s t . Mar. Sci. Univ. Tex. 4:156-200.

Sylvester , J. R. 1975. C r i t i c a l thermal maxima o f three species o f Hawaiian es tuar ine f i s h : a com- pa ra t i ve study. J. F i sh B i o l . 7:257-262.

Sylvester , J. R., C. E. Nash, and C. E. Emberson. 1974. Pre l iminary study o f temperature to1 erance i n j u v e n i l e Hawaiian m u l l e t (Mu il

Prog. Fish- d- u t

Sylvester , J. R., C. E. Nash, and C. R. Emberson. 1975. S a l i n i t y and oxygen to lerances o f eggs and 1 arvae of Hawaiian s t r i p e d mu1 1 et , Mu il cephalus L. J. F i sh B i o l . h - 6 2 9 .

Thomson, J. M. 1955. The movements and migra t ions o f mu1 l e t (Mugil cephal us L. ). Aust. J. Mar. Freshwater Res. 6:328-347.

Thomson, J. M. 1963. Synopsis o f b i o l o g i c a l data on the grey mu1 l e t Mugi 1 cephal us ~ i n n a e u s 1758. Aust. ' m . 0 . Div. F i s h Oceanogr. F ish. Synop. 1. 66 pp.

F l o r i d a ) - - S t r i ped Mu1 1 e t 7. Aufhor(s) I 8. Perlonn8ng Oqanlrat lon Rapt. NO.

SO271 -101

Clark R. C o l l i n s

F l o r i d a Coope ra t i ve F i s h and Wi ld1 i f e Research U n i t 11 7 New ins -Ze ig l e r Hal 1 11. ~ o n t r m c t ( ~ ) or t r a n t ( ~ ) no.

REPORT DOCUMENTATION '.-REPOUT No, ' 2.

P A G E i B i o l . Rep. 82(11.34)*

U n i v e r s i t y o f F l o r i d a G a i n e s v i l l e , FL 32611

3. Racbonent's Accession NO.

12. Spnsoring Orgeniratlon Name and Address

N a t i o n a l Coas ta l Ecosystems Team U.S. Army Corps o f Eng ineer F i s h and W i l d l i f e S e r v i c e Waterways Exper iment Sta. U.S. Dep. o f t h e I n t e r i o r P.O. Box 631 Washington, DC 20240

15. Supplementay Holes

4. Title and Subtntle I Recar( Date 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 A p r i l 1985 Requirements of Coas ta l F i shes and I n v e r t e b r a t e s (South

*U.S. Army Corps o f Eng ineers Repo r t No. TR EL-82-4

16. Abstract (Urn i t 200 words)

Species p r o f i l e s a r e l i t e r a t u r e summaries o f t h e taxonomy, m o r p h o l o ~ y , d i s - t r i b u t i o n , l i f e h i s t o r y , b i o l o g i c a l and p h y s i c a l env i ronments , and e n v i r o n - menta l r equ i r emen ts o f c o a s t a l spec i es o f f i s h e s and a q u a t i c i n v e r t e b r a t e s . They a r e des igned t o a s s i s t i n env i r onmen ta l impac t assessment and p e r m i t r ev i ew . The s t r i p e d m u l l e t (Mug i l c e h a l u s ) i s a v a l u a b l e f o o d f i s h and b a i t f i s h . About 31 m i l l i o n pounds Pr were anded i n F l o r i d a i n 1981. S t r i o e d ~ u l l e t a r e caught w i t h hook and l i n e , trammel n e t s , g i l l n e t s , and se ines , and by snagging. Some s t r i p e d m u l l e t spawn a l o n g beaches, b u t t h e y u s u a l l y spawn o f f s h o r e i n r e l a t i v e l y deep ( t o 1,800 m) c o o l wa te r s . J u v e n i l e s spend t h e i r f i r s t y e a r i n s a l t marshes, e s t u a r i e s , and c o a s t a l wa te rs . S t r i p e d m u l l e t u s u a l l y a v o i d w a t e r c o l d e r t h a n 16O C.

17. Document Analysis a. D a u r i p l o o

F i shes E s t u a r i e s Feed ing Growth

1 b. Identlhsn/Qpen.Endcd Terms

S t r i p e d mu1 l e t L i f e h i s t o r y Mug i l cephalus Spawning Sal i n i t y r equ i r emen ts H a b i t a t r equ i r emen ts Temperature r e q u i rements

c. COSATl F$eld/Cmup - , 19. Stcursty Class (Thcs Ream) 21. No of Pages

Unl i m i t e d -- : U n c l a s s i f i e d - ' 20. Sccurnt Class (This Page) ( ~ n c l ' a s s i f i e d

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