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U. S. Fish and Wildlife Service Nntinnnl Wptlnndc R ~ c p n r r h Conk*
Biological Report 82 (1 1.52) 700 Cajun Dome Boulevard TR EL-82-4
June 1986 Lafayette, Louisiana 70506
Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Florida)
GRAY, LANE, MUTTON and YELLOWTAIL SNAPPERS
Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers
B i o l o g i c a l Report 82(11.52) TR EL-82-4 June 1986
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 F lo r i da )
GRAY, LANE, MUTTON, AND YELLOWTAIL SNAPPERS
Stephen A. Bortone and
James L. W i 11 iams Department o f B io logy
U n i v e r s i t y o f West F l o r i d a Pensacola, FL 32514
P r o j e c t Manager C a r r o l l Cordes P r o j e c t O f f i c e r John Parsons
Nat ional Coastal Ecosystems Team U.S. F i sh and W i l d l i f e Service
1010 Gause Boulevard S l i d e l l , LA 70458
Performed f o r
Coastal Ecology Group Waterways Experiment S ta t i on U.S. A r m y Corps o f Engineers
Vicksburg, MS 39180
and
Nat ional Coastal Ecosystems Team Research and Development 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 Washi ngton, DC 20240
T h i s s e r i e s should be re fe renced as f o l l ows :
U. S. F i s h and W i l d l i f e Serv ice. 1983-19-. Species p r o f i l e s : l i f e h i s t o r i e s and environmental requi rements o f coas ta l f i s h e s and i nve r teb 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 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 ows :
Bortone, S.A., and J. L. Wi l l i ams. 1986. Species p r o f i l e s : l i f e h i s t o r i e s and envi ronmenta l requi rements o f coas ta l f i s h e s and i n v e r t e b r a t e s (South F lo r ida) - -g ray , lane, mutton, and y e l l o w t a i 1 snappers. U. S. F i s h W i l d l . Serv. B i o l . Rep. 82(11.52). U. S. Army Corps o f Engineers, TR EL-82-4. 18 pp.
PREFACE
This species p r o f i l e i s one o f a se r i es on coasta l aquat ic organisms, p r i n c i p a l l y f i s h , o f spor t , commercial, o r eco log ica l importance. The p r o f i 1 es are designed t o p rov ide coasta l managers, engineers, and b i o l o g i s t s w i t h a b r i e f comprehensive sketch o f t h e b i o l o g i c a l c h a r a c t e r i s t i c s and environmental requirements o f t he species and t o descr ibe how populat ions o f t h e species may be expected t o r e a c t t o environmental changes caused by coasta l development. Each p r o f i l e has sect ions on taxonomy, 1 i f e h i s t o r y , eco log ica l r o l e , environmental requirements, and economic importance, i f app l icab le . A t h r e e - r i n g b inde r i s used f o r t h i s se r i es so t h a t new p r o f i l e s can be added as they are prepared. Th is p r o j e c t i s j o i n t l y planned and f inanced by t h e U.S. Army Corps o f Engineers and t h e U.S. F i sh and W i l d l i f e Service.
Suggestions o r quest ions regard ing t h i s r e p o r t should be d i r e c t e d t o one of t he f o l l o w i n g addresses.
I n fo rma t ion Trans fer Specia l i s t Nat iona l Coastal Ecosystems Team U.S. F i s h and W i l d l i f e Serv ice NASA-Slidell Computer Complex 1010 Gause Boulevard S l i d e l l , LA 70458
U.S. A r m y Engineer Waterways Experiment S t a t i o n A t ten t i on : WESER-C Post O f f i c e Box 631 Vicksburg, MS 39180
CONVERSION TABLE
M e t r i c t o U.S. Customary
m i l 1 i i ne te rs (m) cent imeters (on) meters (m) k i 1 m e t e r s ( km)
2 square meters (m ) 10.76 square k i 1 m e t e r s ( km2) 0.3861 hec ta res (ha) 2.471
l i t e r s ( 1 ) cub ic ineters (m3) cub ic meters
To Obta in
inches inches f e e t m i l es
square f e e t square n i l e s acres
gal 1 ons cub ic f e e t acre- f e e t
m i l l iyrams (mg) 0.00003527 ounces grams ( g ) 0.03527 ounces k i l oyrarns (kg ) 2.205 pounds m e t r i c tons ( t ) 2205.0 pounds m e t r i c tons 1.102 s h o r t tons k i 1 ocal o r i e s ( kcal ) 3.965 B r i t i s h thermal u n i t s
Ce ls ius degrees 1.8(OC) + 32 Fahrenhe i t degrees
U.S. Customary t o M e t r i c
inches 25.40 inches 2.54 f e e t ( f t ) 0.3048 fathoms 1.829 m i l e s ( m i ) 1.609 n a u t i c a l m i l e s ( m i ) 1.852
square f e e t ( f t 2 ) acres 2 square m i l e s (mi )
ga l 1 ons ( gal ) 3.785 cub ic f e e t ( f t 3 ) 0.02831 a c r e - f e e t 1233.0
ounces (02) 28.35 pounds ( l b ) 0.4536 s h o r t tons ( t o n ) 0.9072 B r i t i s h thermal u n i t s ( B t u ) 0.2520
m i l 1 imeters cen t imete rs ineters meters k i l o m e t e r s k i 1 oqe te rs
square meters hectares square k i l o m e t e r s
1 i t e r s c u b 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 e s
Fahrenhei t degrees 0.5556(OF - 32) Ce ls ius degrees
CONTENTS
Page
PREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i i i CONVERSION TABLE . . . . . . . . . . . . . . . . . . . . . . . . . . . . i v ACKNOWLEDGMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v i
NOMENCLATURE/TAXONOMY/RANGE . . MORPHOLOGY/IDENTIFICATION A I D S
G r a y S n a p p e r . . . . . . . . . . . . . . L a n e S n a p p e r
. . . . . . M u t t o n S n a p p e r . . . . Y e l l o w t a i l S n a p p e r
REASON FOR INCLUSION I N SERIES L I F E HISTORY . . . . . . . . .
. . . . . . . . . S p a w n i n g F e c u n d i t y a n d E g g s . . . .
. . . . . . . . . . L a r v a e . . . . . P r e - A d u l t P h a s e
A d u l t s . . . . . . . . . . . . . . GROWTH CHARACTERISTICS
COMMERCIAL AND SPORT FISHERY . . . . . . . . . ECOLOGICAL ROLE
ENVIRONMENTAL REQUIREMENTS . . T e m p e r a t u r e L i m i t s . . . . S a l i n i t y . . . . . . . . .
. . . . . . . . . S u b s t r a t e D e p t h . . . . . . . . . .
LITERATURE C I T E D . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
ACKNOWLEDGMENTS
We thank W i l l i am Seaman o f t he U n i v e r s i t y o f F lo r i da , Gainesvi l l e , and personnel from t h e F l o r i d a Sea Grant f o r he lp i n o rgan i z i ng t h i s repor t . We a l so thank H.D. Hoese and R.H. Moore f o r permiss ion t o use Table 2. Add i t i ona l l y , we thank C h u r c h i l l Grimes, Nat ional Marine F i she r i es Service, Panama C i t y , F lo r i da , Robert Shipp, Un i ve rs i t y o f South Alabama, Mobi le , and Wil lam Seaman f o r rev iewing t h e manuscript.
Gray Snapper
Figure 1. Gray, lane, mutton, and y e l l o w t a i l snappers. Adapted from i l l u s t r a - t i o n s i n Evermann and Marsh (1902).
GRAY, LANE, MIJTTON, AND YELLOWTAIL SNAPPERS
S c i e n t i f i c name. . . Lut janus gr iseus ( L i nnaeus)
P re fe r red common name. . Gray snapper (F igu re 1 )
Other common names . . . . . Mangrove snapper, g r e y snappe r , 1 o w y e r , mango s n a p p e r , b l a c k s n a p p e r , cabal 1 e ro te , pargo p r i e t o , pargo , p a r g o d i e n t o n , pargo de p i e d r a , p a r g o moreno , v i v a n e a u s a r d e g r i s e , aquadera ( C e r v i g o n 1966; Starck 1971; F i sche r 1978)
S c i e n t i f i c name . . Lut ianus synagr is (Linnaeus)
P re fe r red common name. . Lane snapper ( F igure 1)
Other common names . . Candy s t r i p e r , r a i n b o w snapper, pa rgo guanapo, p a r g o b i a j a i b a , v i v a n e a u gazou,
a r i a c o ( C e r v i g o n 1966; F i s c h e r 1978)
S c i e n t i f i c name . . . Lut janus anal i s (Cuv ie r )
P re fe r red common name. . . . . Mutton snapper (F igu re 1)
Other common names . . . Mutton f i sh , k i n g snapper, pargo cebadai, pargo c r i o l l o , vivaneau sorbe ( C e r v i g o n 1966, F i s h e r 1978)
S c i e n t i f i c name . . Ocyurus chrysurus ( Bl och)
P re fe r red common name. . . Y e l l o w t a i l snapper (F igu re 1)
Other common names . . . Rabir rubia, v i v a n e a u quaue j a u n e ( C e r v i gon 1966; F i scher 1978)
C l ass . . . . . . . . . Osteichthyes Order . . . . . . . . . Perc i formes Fami ly . . . . . . . . . Lut jan idae
Geographic range: The f.our species o f snappers i n h a b i t t h e t r o p i c a l and sub t rop i ca l coas ta l waters o f t h e western A t l a n t i c Ocean, and i n t h e summer they i n h a b i t waters o f t h e temperate zones as f a r n o r t h as Massachusetts (North Caro l ina f o r t h e lane snapper). Along t h e con t i nen ta l she1 f o f pen insu lar F l o r i d a , t h e f o u r species a re associated w i t h t h e hard bottoms (Figure 2). Juven i les are more broad ly d i s t r i b u t e d , o f t e n i n h a b i t i n g shal low es tuar ies .
The f o u r snapper species' i n d i v i d u a l desc r i p t i ons a r e f o l l owed by a summary o f t h e i r morphol o g i c a l characters (Table 1 ) and a key t o the L u t j a n i d a e o f t h e G u l f o f M e x i c o (Table 2) which should enable i d e n t i - f i c a t i o n o f most s p e c i e s o f f sou th F l o r i d a as w e l l . There i s some d i s p a r i t y between Table 1 and Table 2 i n t he data on numbers o f g i l l rakers.
Gray Snapper
D o r s a l f i n , 10 s p i n y r a y s , 1 4 s o f t r a y s ; ana l f i n , 3 s p i n y r a y s , 7 -8 s o f t r a y s . Number o f p o r e d s c a l e s a l o n g l a t e r a l l i n e , 43 -47 . Number o f g i l l r a k e r s on f i r s t g i l T arch, 9-10 on upper p o r t i o n , 21-22 on l o w e r . Body somewhat e l o n g a t e ; n o t s t r o n g l y c o m p r e s s e d ; h e a d p r o f i l e nea r l y s t r a i g h t o r s l i g h t l y convex f r o m s n o u t t o nape. Head l e n g t h i s 33% t o 38% o f s t a n d a r d l e n g t h (SL ) ; body dep th i s 31% t o
l ~ a r g e l y ex t rac ted from Jordan and Evermann (18981, Anderson ( 1 9 6 7 ) , Bohl ke and Chap1 i n (19681, Randal 1 (19681, and Hoese and Moore (1977 ) . See the re fe rences f o r e x p l a n a t i o n s of a b b r e v i a t i o n s and measurements.
38% o f SL. S n o u t l e n g t h 33% o f h e a d l e n g t h ; e y e , 2 1 % . M o u t h 1 arge, max i1 1 a r y r e a c h i n g f r o n t o f p u p i l , 31% o f head 1 e n g t h . B o t h j a w s w i t h a' n a r r o w b a n d o f v i l l i f o r m t e e t h ; u p p e r j a w a l s o w i t h f o u r c a n i n e s a n t e r i o r l y , t w o o f t h e s e e n l a r g e d . Vomer w i t h v i l l i f o r m t e e t h i n an arrow-shaped p a t c h w i t h a m e d i a l p o s t e r i o r p r o j e c t i o n . P r e o p e r c l e f i n e l y s e r r a t e above w i t h c o a r s e r s p i n e s a t i t s d e e p l y e m a r g i n a t e d a n g l e . do rsa l s p i n e s s t r o n g ; f o u r t h s p i n e l o n g e s t , 40% o f head l e n g t h . S o f t 'dorsal rounded, t h e 9 t h and 1 0 t h r a y s l o n g e s t . Cauda l m a r g i n a t e . A n a l f i n somewhat a n g u l a t e , i t s second s p i n e equa l t o o r s l i g h t l y l o n g e r t h a n t h i r d . P e c t o r a l s s h o r t , f i n l e n g t h n e a r l y e q u a l t o d i s t a n c e f r o m s n o u t t o u p p e r preoperc l a r margin.
C o l o r i n l i f e : Body and f i n s va r i ab le : gray t o green-brown , g e n e r a l l y w i t h r e d d i s h t i n g e s . m e d i a n f i n s d a r k e r , o f t e n edqed w i t h w h i t e o r y e 1 l o w . P e c t o r a l s n e a r l y c o l o r l e s s . No b l a c k s p o t on body s i d e . Young w i t h a b l a c k b a r f r o m t i p o f s n o u t t h r o u g h e y e , o f t e n w i t h a b l u e s t r e a k b e n e a t h eye. Young o f t e n w i t h a p a t t e r n o f n a r r o w p a l e b a r s 1 a t e r a l l y . J u v e n i l e s and a d u l t s commonly b e a r o r a n g e o r d a r k b r o w n d o t s o n centers o f sca les on s i d e s , f o r m i n g rows o f spots.
Lane Snapper
D o r s a l f i n , 1 0 s p i n y r a y s , 12-13 s o f t r a y s ; ana l f i n , 3 s p i n y r a y s , 8 - 9 s o f t r a y s . Number o f
.pored s c a l e s a1 ong 1 a t e r a l - 1 i ne, 47-52. Number o f g i l l r a k e r s on' f i r s t g i l l a r c h , 6 - 7 o n u p p e r po r t i on , 13-14 on lower. Body oblong and compressed; head p r o f i l e n e a r l y s t r a i g h t from snou t t o nape. Head l eng th 38% o f SLY body d e p t h 34% t o 42% o f SL. Snout l eng th 33% o f head
PETERSBURG
OKEECHOBEE
CAPE ROMAN
@ Coaatal distribution
Figure 2. The d i s t r i b u t i o n o f t he p re fe r red h a b i t a t o f a d u l t snappers (hard bottoms i n c l u d i n g both na tu ra l and a r t i f i c i a l r e e f s ) o f f t he south coast o f F lo r ida .
Table 1. Summary o f morphologica l charac te rs o f f o u r snappers.
Species o f snapper
Character Gray Lane Mutton Y e l 1 owta i 1
Dorsal f i n e l ementsa X, 14 X, 12-13 I X - X I , 13-14 X, 12-14
Anal f i n elementsa 111, 7-8 111, 8-9 111, 8 111, 8-9
Pored scales i n l a t e r a l 1 i n e
G i l l r ake rs 9-10+21-22 6-7+13-14 6-8+12-13 21-22 ( t o t a l )
Head l e n g t h b 33%-38% 38% 34%-38% 37%
Body depthb 31%-38% 34%-42% 31%-40% 30%-34%
P o s t e r i o r body spo t Absent Sl i g h t l y Equal t o Absent 1 arger pupi 1 than eye, diameter, one- f o u r t h en ti re1 y be1 ow above 1 a t e r a l 1 a t e r a l 1 i n e 1 i n e
Ho r i zon ta l body s t r i p e Absent Absent Absent Present
a Roman numerals i n d i c a t e number o f sp iny rays , a r a b i c numerals s o f t rays .
b ~ i v e n as a percentage o f s tandard leng th .
l e n g t h ; e y e s m d i l , 2 0 % o f h e a d leng th . M a x i l l a r y reach ing t o f r o n t o f eye, 36% o f head l e n g t h . Nar row band o f v i l l i f o r m t e e t h i n each jaw. The upper jaw a1 so bears f o u r c a n i n e tee th , two o f them en1 arged. V i 11 i- f o r m t e e t h on vomer i n an a r r o w - s h a p e d p a t c h w i t h o u t a m e d i a l p o s t e r i o r p r o j e c t i o n . P r e o p e r c l e f i n e l y s e r r a t e above w i t h c o a r s e r s p i n e s a t i t s a n g l e , w h i c h i s emarginate. D o r s a l s p i n e s s l ender ; t he f o u r t h i s l o n g e s t , 37% o f head l e n g t h . S o f t d o r s a l s o m e w h a t angul a t e p o s t e r i o r l y , the e i g h t h ray . 1 o n g e s t . C a u d a l f i n m o d e r a t e 1 y
emarg ina te . Anal f i n rounded, n o t angul ate; i t s second s p i n e s t r o n g e r t h a n t h i r d , b u t o f e q u a l 1 e n g t h . -.
Pectora l f i n s 26.3% t o 30.3% o f SLY reaching f r o n t o f anal f i n .
C o l o r i n l i f e : D i f f u s e b l a c k spo t o r b l o t c h s l i g h t l y l a r g e r t h a n eye below s o f t p o r t i o n o f dorsal f i n , t h e l a t e r a l 1 i n e t r a n s e c t i n g i t s 1 owermost f o u r t h . Body s i 1 v e r y v i n k t o r e d w i t h seven o r e i g h t y e 1 l o w s t r i p e s ; wavy d i a g o n a l o r y e l l o w l i n e s o n l a t e r a l s u r f a c e a b o v e l a t e r a l l i n e . C a u d a l f i n r e d t o p ink , edges dusky. Anal, p e l v i c , and
T a b l e 2. Key t o t h e s n a p p e r s of t h e Gulf o f Mexico (Hoese and Moore 1977).
Dorsal s p i n e s X; l a t e r a l 1 ine s c a l e s fewer than 00 . . . . . . . . . . 2 . . . Dorsal sp ines XII ; l a t e r a l 1 i n e s c a l e s nore than 50. .Vermil ion
snapper , Rhomboplites aurorubens.
Sca les on s o f t dorsa l and anal f i n s , with l a s t rays of t h e s e f i n s n o t produced; dorsa l rays 12-14; g i l l r ake r s on lower l i n b of
. . . . . . . . . . . . . f i r s t a rch few (7 -10) o r numerous (19-21) 3 No s c a l e s on s o f t dorsa l anal f i n s , with l a s t rays produced a s
f i l a m e n t s (which may be broken o f f ) ; dorsal rays 10-11; g i l l . . . r a k e r s on lower limb of f i r s t a rch 16-17 Wenchman,
P r i s t ipomoides aqu i l o n a r i s .
G i l l r a k e r s on lower 1 imb of f i r s t a rch few !7-10); caudal lobes . . . . . . . . . . . . n o t produced i n a d u l t s ; no p terygoid t e e t h . 4
G i l l r ake r s on lower 1 imb of f i r s t a rch numerous, 19-21; caudal . . . l obes produced in a d u l t s : p terygoid t e e t h p resen t Yellowtail
snapper , Dcyurus chrysurus .
Dorsal rays usua l ly 14, r a r e l y 13. . . . . . . . . . . . . . . . . . . 5 Dorsal r ays normally 12 . . . Lane snapper, Lutjanus svnaqr i s .
Vomerine too th patch with p o s t e r i o r extens ion . . . . . . . . . . . . 6 Vomerine too th patch wi thout p o s t e r i o r extens ion . . . . . . . . . . . 8
Gi l l r a k e r s on lower 1 imb of f i r s t a rch 7-8; anal rays usua l ly 8 . . . 7 G i l l r ake r s on lower 1 imb of f i r s t a rch 9-10; anal rays usua l ly 9
( 8 i n 11% o f Gulf of i-lexico specimens) . . . Red snapper , Lut janus campechanus.
Body comparat ively s t o u t , g r e a t e s t depth 36%-43% ( u s u a l l y 37%-42%) o f s tandard l e n g t h ; pec to ra l f i n in a d u l t s longer than d i s t a n c e from t i p o f snout t o pos ter iormost edge of p reoperc le ( i n j u v e n i l e s pec to ra l f i n a s i n Lut janus g r i s e u s ) . . . . . . . .
Body comparatively s l e n d e r , with g r e a t e s t depth 31-38% (usual 1 y 32-37%) of s t andard l eng th ; pec to ra l f i n in a d u l t s a s lonq a s d i s t a n c e from t i p of snout t o pos ter iormost edge of p reoperc le Gray snapper , Lutjanus g r i s eus .
L a t e r a l - l i n e s c a l e s 45-47; body depth 29%-32% of s t andard l e n g t h ; caudal wi thout pointed lobes . . . Cubera snapper, Lutjanus cyanopterus .
La te ra l -1 i n e s c a l e s 47-51; body depth 31-$0? o f s tandard l e n g t h ; caudal with pointed lobes . . . Mutton snapper, Lutjanus anal i s .
Sca les l a r g e a wi th 39-44 ( u s u a l l y 40%-43%) t r a n s v e r s e rows between " s c a l e bone" and base of caudal f i n ; 40-45 pored s c a l e s i n
. . . l a t e r a l l i n e ; 5-7 rows between base of dorsa l and l a t e r a l l i n e Schoolmaster, Lut.ianus apodus.
S c a l e s small , with 45-49 t r a n s v e r s e rows hetween " s c a l e hone" and . . . base of caudal ; 46-48 pored s c a l e s in l a t e r a l 1 ine nog
snapper , Lut.ianus jocu.
en1 arged scal e above and behi n d t h e n i l 1 s l i t .
d i s t a l p o r t i o n o f dorsal f i n s ye1 low. P r o x i m a l p o r t i o n s o f d o r s a l f i n reddish.
Mutton S n a ~ ~ e r
Dorsal f i n , 9 t o 11 ( u s u a l l y 10) s p i n y r a y s , 13-14 s o f t r a y s ; a n a l f i n , 3 sp iny rays, 8 ( r a r e l y 7 ) s o f t rays . Number o f po red s c a l e s a l o n g l a t e r a l l i n e , 47-51. Number o f g i l l r a k e r s on f i r s t g i l l a r c h , 6 - 8 o n upper por t ion , 12-13 on l o w e r . Body d e e p a n d c o m p r e s s e d ; . d o r s u m i s e l e v a t e d w i t h a s t e e p s t r a i g h t p r o f i l e t o t i p o f s n o u t . H e a d l e n g t h i s 34% t o 3 8 % o f SL, b o d y d e p t h 3 1 % t o 4 0 % o f SL . S n o u t 1 e n g t h 35% o f h e a d 1 e n g t h ; e y e smal l , 18.5% o f head l e n g t h . Mouth moderate; maxi1 1 a r y b a r e l y r e a c h i n g f r o n t o f eye, 37% o f head 1 e n g t h . Upper and l o w e r j a w s w i t h a n a r r o w hand o f v i l l i f o r m t e e t h ; s i x c a n i n e t e e t h i n uppe r jaw, f o u r e n l a r g e d . Vomer w i t h v i l l i f o r m t e e t h i n a w e d g e - s h a p e d p a t c h w i t h o u t a p o s t e r i o r l y d i r e c t e d m e d i a1 p r o - j e c t i o n . C o a r s e s e r r a t i o n a1 o n q e n t i r e edge o f p r e o p e r c l e; n o t c h a t ang le s h a l l o w . D o r s a l s p i n e s weak and s l e n d e r , d o r s a l o u t 1 i n e n e a r l y s t r a i g h t ; f o u r t h s p i n e 1 onqes t , 37% o f h e a d l e n g t h . M a r g i n o f s o f t do rsa l angul a te . Caudal f i n deep1 y e m a r g i n a t e . A n a l f i n a n g u l a r p o s t e r i o r l y , s i m i l a r t o s o f t d o r s a l m a r g i n ; s e c o n d a n d t h i r d s p i n e s e q u a l l y s t r o n g . P e c t o r a l s 1 ong , reach ing j u s t p a s t ana l o r i g i n , 26% t o 31% o f SL.
C o l o r i n 1 i f e : There a r e b o t h b a r r e d and p l a i n c o l o r phases w h i c h c o r r e s p o n d t o r e s t i n g and swimming a c t i v i t i e s , r e s p e c t i v e l y . A d u l t 01 i v e above, w h i t e l a t e r a l l y , a n d r e d d i s h be low. A l l f i n s w i t h r e d , c a u d a l m a r g i n b l a c k . S n o u t w i t h i r r e g u l a r d i a g o n a l b l u e 1 i n e c o n t i n u i n g i n back o f eye; a n o t h e r i r r e g u l a r b l u e l i n e r u n s f r o m m i d d l e o f m a x i l l a r y t o e y e .
L a t e r a l s u b d o r s a l s p o t e n t i r e l y above l a t e r a l 1 i n e , equa l t o p u p i l d i a m e t e r . L a t e r a l s u b d o r s a l s p o t i n j u v e n i l e s equa l t o eye and l o w e r h a l f p r o j e c t i n g b e l o w l a t e r a l l i n e .
Y e l l o w t a i l Snapper
D o r s a l f i n , 1 0 s p i n y r a y s , 12-14 s o f t r a y s ; ana l f i n , 3 s p i n y r a y s , 8 - 9 s o f t r a y s . Number o f p o r e d s c a l e s a1 o n g 1 a t e r a l 1 i ne , 48-49. Number o f g i l l r a k e r s o n f i r s t g i l l a r c h , 2 1 - 2 2 . B o d y f u s i f o r m ; c o m p a r a t i v e 1 y e l o n g a t e dorsum o n l y s l i g h t l y e l e v a t e d ; nape t o d o r s a l f i n g e n t l y cu rved . Head l e n g t h 37% o f SL, body d e p t h 30% t o 34% o f SL. Snout l e n g t h 33% o f head 1 enath, eye 20% o f head 1 ength. Pre- o r b i t a l narrow, 15% o f head l e n g t h a t i t s n a r r o w e s t w i d t h . M o u t h i n comparison w i t h o t h e r snapper s m a l l , o b l i q u e , t h e l o w e r j a w p r o j e c t i n g . M a x i l l a r y extending t o f r o n t o f eye, 38% o f head 1 ength. Upper jaws w i t h a narrow band o f v i l l i f o r m t e e t h w i t h f i v e o r s i x somewhat s m a l l c a n i n e t e e t h l a t e r a l 1.y. L o w e r j a w w i t h l a r g e r v i l l i f o r m t e e t h . T e e t h o n vomer i n a r row-shaped p a t c h w i t h a m e d i a l p o s t e r i o r l y d i r e c t e d p r o - j e c t i o n . Preoperc l e weak1 y s e r r a t e , t e e t h a t a n g l e a 1 s o s h a l l o w l y emarg ina te . D o r s a l s p i n e s s t r o n a ; f i f t h i s longes t , 42% o f head l eng th . D o r s a l f i n n o t d e e p l y e m a r g i n a t e . Caudal f i n long, deeply forked; upper l o b e l o n g e r . P e c t o r a l s 1 o n g , r e a c h i d g anus; 83% o f head l e n g t h . T h i r d anal sp ine l onge r than second.
C o l o r i n l i f e : A p r o m i n e n t y e l l o w m i d - l a t e r a l s t r i p e b e g i n n i n g on snout and b r o a d e n i n g p o s t e r i o r l y t o e n t i r e depth o f cauda l pedunc le . E n t i r e cauda l f i n y e l l o w . Y e l l o w spots on body above y e l l o w l a t e r a l s t r i p e . Background body c o l o r g r a y t o o l i v e above, wh i t e below. D o r s a l f i n y e l l o w d i s t a l l y , p a l e a t b a s e a n t e r i o r l y . A l l o t h e r f i n s p a l e ye1 1 ow t o c l ear.
REASON FOR INCLUSION I N SERIES
T h e s e f o u r s n a p p e r s p e c i e s compose a major p o r t i o n o f b o t h t h e s p o r t and c o m m e r c i a l c a t c h e s o f F l o r i d a ' s snapper f i s h e r y . They a r e g e n e r a l l y a s s o c i a t e d w i t h r e e f s , on hard o r l i v e bottoms. They a re h i g h l y sought a f t e r f o r human food. These snappers a re a l s o an impor tan t s h a l l ow-water component o f t h e s o u t h F l o r i d a r e e f and n e a r - r e e f i c h t h y o - fauna. Gray snappers, i n p a r t i c u l a r , a r e associated w i t h e s t u a r i n e a r e a s a s a d u l t s , a n d e s p e c i a l l y a s j u v e n i l e s . T h e s e s n a p p e r s a r e i m p o r t a n t m e m b e r s o f m a r i n e , n e a r s h o r e r e e f , o r r e e f - 1 i k e b io topes .
LIFE HISTORY
Spawning
Most snappers, i n c l u d i ng t h e s e f o u r s p e c i e s , s p a w n i n g r o u p s ( W i c k l u n d 1 9 6 9 ; S t a r c k 1 9 7 1 ; Thompson and Munro 19741, u s u a l l y o f f s h o r e (Croker 1962; S p r i n g e r and Woodburn 1960). Th i s may i n d i c a t e an a c t u a l o f f s h o r e movemen t d u r i n g r e p r o d u c t i o n a s s u s p e c t e d f o r a n u m b e r o f o t h e r t r o p i c a l r e e f - assoc ia ted spec ies (Johannes 19781, o r p a r t o f a tendency f o r l a r q e r , 01 der ( i .e., sexual 1 y mature) coas ta l f i s h e s t o occur i n deeper w a t e r t h a n j u v e n i l e s . Unconfirmed o b s e r v a t i o n s i n d i c a t e t h a t g r a y snapper spawn a t dusk as p a r t o f a d a i l y a c t i v i t y c y c l e and may h a v e p e a k s p a w n i n g p e r i o d s t h a t f o l l o w a l u n a r c y c l e ( S ta rck 1971).
Seasonal spawning p a t t e r n s v a r y among t h e s e f o u r 1 u t j a n i d s p e c i e s , b u t g e n e r a l l y , o n c e t h e y a t t a i ~ s e x u a l m a t u r i t y , t h e y h a v e a p r o t r a c t e d s p a w n i n g p e r i o d w i t h seasonal peaks (Erdman 1976 ) . The
l e n g t h o f t he spawning p e r i o d may be even l o n g e r f o r f i s h i n t h e warmer p a r t s o f t h e i r r ange . No d a t a a r e a v a i l a b l e on t he t ime o f spawning o f mut ton snapper.
I n d i v i d u a l g r a y s n a p p e r p r o b a b l y s p a w n m o r e t h a n o n c e d u r i n g a s p a w n i n g s e a s o n ( S t a r c k 1 9 7 1 ; B a s h i r u l l a h 1 9 7 5 ) . O f f V e n e z u e l a , t h e s p a w n i n g s e a s o n began i n J u l y and ended i n Oc tobe r ( G u e r r a - C a m p o s a n d B a s h i r u l 1 a h 1975).
I n Cuba, l a n e s n a p p e r spawn from March t h r o u q h September, w i t h spawning peaks i n J u l y and A u g u s t (Rodriguez-Pi no 1962; Druzhi n i n 1970 ) . Erdman ( 1 9 7 6 ) r e p o r t e d a spawning peak i n May o f f Puer to Rico, and Munro e t a1 . (1973) n o t e d m a t u r e e g g s i n l a n e s n a p p e r c a u g h t i n February o f f Jamaica.
Ye1 1 o w t a i l snapper were sexual 1 y mature from March t o May i n nearshore areas o f f Jamaica, b u t o f f s h o r e t h e y apparen t ly spawned year r ound (Munro e t a l . 1 9 7 3 ) . E r d m a n ( 1 9 7 6 ) i n d i c a t e d a d e c l i n e ii~ s p a w n i n g a c t i v i t y among s p e c i m e n s c a u g h t du r i ng November through December o f f Puerto Rico. P iedra (1969) r e p o r t e d t h a t ye1 1 o w t a i l snapper femal es o f f Cuba had r i p e gonads f r o m March t o August.
Age a t matura t ion i s a p p a r e n t l y unknown f o r these s p e c i e s , and most s t u d i e s r e 1 a t e m a t u r i t y t o 1 e n g t h . Thompson and Flunro (1974) w r o t e t h a t l ane snapper ma1 es and females a t t a i n sexual m a t u r i t y a t 18 cm f o r k 1 e n g t h ( F L ) . Ye1 1 o w t a i l s n a p p e r ma1 e s become s e x u a l l y m a t u r e a t 26 cm FL and females a t 29 t o 3 1 cm FL. Gray snapper sex r a t i o s have been repo r ted as equal o f f F l o r i d a ( S t a r c k 19711, b u t o f f Cuba a r a t i o o f two f ema les t o o n e m a l e h a s b e e n o b s e r v e d ( G u e r r a - C a m p o s a n d B a s h i r u l l a h (1975 1.
H y b r i d i z a t i o n o f y e 1 l o w t a i l snapper w i t h l a n e snapper has been r e p o r t e d ( R o d r i q u e z - P i no 1961 ; = L u t j a n u s a m b i g u u s ) . Y e l l o w t a i l snapper have a l s o been t h o u g h t t o h y b r i d i z e w i t h t h e d o g s n a p p e r (Lut janus jocu) as s t a t e d by Jo rdan - and- -~vermann (1898; = Lut janus $1 u t jano ides) .
Fecundity and Eggs
Fecundity est imates repor ted f o r a l l f ou r snapper species v a r y w i d e l y (Tab1 e 3 ) . A ~ a r t i a l reason fo r t h i s v a r i a t i o n may be the r e l a t i o n between fecund i t y and f i s h s ize, as noted f o r the gray snapper (Guerra-Campos and B a s h i r u l l ah 1975)) when fecund i t y = 0.00299 x t o t a l l e n g t h (TL, mm)13.26; f ecund i t y = 1478.82 x we ight ( grams)0.973. Gray snapper eggs are s m a l l (be tween 0 . 4 a n d 0 .6 mm i n d i a m e t e r ) w i t h one c e n t r a l o i l g l o b u l e ; eggs a r e a p p a r e n t l y n o t adhesive (Starck 1971; Guerra-Campos and B a s h i r u l l ah 1975). Lane snapper eggs are l a r g e r (between 0.7 and 0.8
mm ~ o d r i g u e z - P i no 1962). Erhard t ( 1 9 7 6 ) e x a m i n e d t h e m i . c r o s c o p i c anatomy o f lane snapper ovar ian egg membranes; they genera l ly agreed w i t h desc r i p t i ons given f o r those o f o ther perc i fo rm f i shes (Hayashi 1972). Most 1 u t j a n i d species produce pe lag i c eggs (Breder and Rosen 1966).
Larvae
L i t t l e i s known o f l u t j a n i d larvae, as they tend t o be p lank ton ic a t a l eng th l ess than 10 mn TL ( Barans and Pow1 es 1977). Laboratory-reared l a rvae o f gray snapper described by Richards and Saksena (1980) were s i m i l a r i n appearance t o o the r described snapper l a r v a e ( i e . , red snapper Lut janus campechanus by C o l l i n s e t al . 1980; vermi l i o n snapper Rhombo 1 i tes auroru bens by La roc* Accordina t o Richards and Saksena (1980:515), ' 'pigmentation was sparse i n the larvae, being conf ined t o the ven t ra l m i d l i n e o f the t a i l , gut, p e l v i c f i n rays, c l e i t h r a l symphysis, b r a i n and on the base o f the dorsal
Table 3. Fecundi ty est imates (no. o f eggs) f o r f o u r species o f snappers. (SL= standard length, TL = t o t a l length , FL = f o r k length) .
Species and fecund i ty est imates F i s h l e n g t h (mm) Source
Gray 590,000 315 SL 1,086,682-5,900,010 488-660 TL
Lane 347,416-994,787
Mutton 1,365,975
Y e l l o w t a i l 99,660-1,472,594
Starck 1971 Guerra-Campos and
B a s h i r u l l ah 1975
225-335 FL Rodriguez-Pi no 1962
512 FL Rojas 1960
292-382 FL Piedra 1969
f i n . " Gray snapper l a r vae t rans- formed t o j u v e n i l e s a t a s i z e between 6.3 and 9.6 mm SL (Richards and Saksena 1980).
a d a i l y a c t i v i t y p a t t e r n a s s o c i a t e d w i t h n o c t u r n a l f e e d i n g h a b i t s , and ye1 l o w t a i l snapper a re semi pe lag i c wanderers over the r e e f h a b i t a t (Moe 1972).
Pre-Adul t Phase
GROWTH CHARACTERISTICS D a t a a r e s c a r c e o n t h e l i f e
h i s t o r y o f j u v e n i l e s o r p re -adu l t s 03 these fou r snapper species ( ~ h o m p s o n and Munro 1 9 7 4 ) . P r e - a d u l t g r a y s n a p p e r o f t e n o c c u p y h a b i t a , t s d i f f e r e n t f r o m t h a t o f a d u l t s . Juven i l es are usua l l y associated w i t h s h a l l ow Tha l a s s i a g r a s s beds, man- g r o v e r o o t s , j e t t i e s , and p i l i n g s (S tarck 1971; Thompson and Munro 1974). Juven i l es u s i n g t h e i n s h o r e areas o f south F l o r i d a f o r n u r s e r i e s are c o n s p i c o u s l y absen t f r om March t h r o u q b May, b u t a r e a b u n d a n t i n t h e s e a r e a s f r o m J u l y t h r o u g h September ( S p r i n g e r and M c E r l e a n 1962) . Pre-adul t mutton snapper a1 so use inshore grass beds ( S p r i n g e r and McErl ean 1962 ) .
Adul t s
A d u l t g r a y and 1 ane s n a p p e r s t e n d t o occupy a w i d e v a r i e t y o f h a b i t a t s . These j n c l ude n a t u r a l and a r t i f i c i a l hard-bottom substrates, as w e l l as sof t -bot tom areas i n c e r t a i n p a r t s o f t h e i r r a n g e s . M u t t o n snapper a r e o f t e n a s s o c i a t e d w i t h sand, sea g r a s s , and c o r a l r u b b l e . Ye1 1 ow ta i l snapper most o f t e n occupy w a t e r j u s t above t h e h a r d - b o t t o m a r e a s w i t h w h i c h t h e y a s s o c i a t e (Randal l 1967). A d u l t snapper t e n d t o remain i n an area once they become e s t a b l i s h e d . T a g g i n g s t u d i e s i n d i c a t e 1 i t t l e movement b y a d u l t snapper ; o n l y 2 o f 1 3 t a g g e d and r e c o v e r e d g r a y s n a p p e r s h o w e d evidence o f moving more than 5 nmi i n 4 y e a r s . M u t t o n and y e l l o w t a i l snapper showed l i t t l e movement 262 d a y s a f t e r r e 1 ease ( B e a u m a r i a g e 1969). Gray snapper tend t o move i n
L e n g t h - w e i g h t e q u a t i ons h a v e been determined on ly f o r the l ane and y e l l ow ta i l snapper, a1 though S t a r c k (1971) p r e s e n t e d a s c a t t e r p l o t o f t h e l e n g t h s and w e i g h t s o f g r a y snapper. The length-wei gh t equat ions f o l l o w : l ane snapper -- weight (g ) = 4 .965 X 10'5 l e n q t h (mm T L ) 2.80 (Rodriguez-Pino 1962) and weight ( ) = 1.02 x 1 - leng th (mm TL)2.65q4 (Manooch and Mason 1984); and y e l l o w t a i l snapper -- weight (g) = 1.45 x 10-2 l e n g t h (mm TL)3.032 (Thompson and Munro 1974). The est imated ranges o f maximum lengths (mm, TL) f o r t he f o u r species (Manooch and Mason 1984; Thompson and Munro 1974; Bohl ke and Chap1 i n 1968) a re as f o l l ows :
Gray snapper 500-900 Lane snapper 375-512 Mutton snapper 642-750 Ye1 l o w t a i 1 snapper 688-750.
Scales, o t o l i t h s , and v e r t e b r a e have a l l been s u c c e s s f u l l y used t o d e t e r m i n e t h e a g e o f s n a p p e r s ( B o r t o n e and H o l l i n g s w o r t h 1 9 8 0 ) . Scales have been usefu l f o r aging the g r a y snapper ( C r o k e r 1962; S t a r c k 1971; Thompson and Munro 1974) and o t o l i t h s f o r a g i n g g r a y and l a n e snappers (Manooch and Plason 1984 ; Rodriguez-Pi no 1962; A1 egria-C and de Menezes 1970; Claro-Madruga and Bustamente Po la 1977). There are a p p a r e n t d i f f i c u l t i e s i n a q i n g y e l l o w t a i l s n a p p e r w i t h o t o l i t h s (Thompson and Munro 1974); Piedra. (1969) found ver tebrae t o be use fu l , however, and n o t e d t h a t t w o r i n q s equal ed one annul us.
Growth among s n a p p e r s v a r i e s considerably (Table 4 ) . The l a c k of general agreement on t h e i r growth r a t e may be due t o d i f ferences i n methodologies. For exampl e , Cl aro-Madruga and Bustamente Pol a (1977) noted t h a t , although one annulus in gray snapper was produced each year (between February and July off Cuba, b u t between February and March off south Florida; Starck 1971), t h e f i r s t r ing was not an annulus. This f a c t alone would account f o r major di f ferences i n growth a s estimated by d i f f e r e n t researchers.
I n d i v i d u a l g r o w t h r a t e s determined from tagged- and- re1 eased
Table 4. Mean snapper by age Starck 1971; ( Rodri guez-Pi no Piedra 1969.
snapper a1 so indicated t h a t growth i s v a r i a b l e , p robab ly being s t r o n g l y i n f l u e n c e d by 1 o c a l h a b i t a t s and t h e i r abundance o f f o o d . S t a r c k (1971) found t h a t gray snapper growth v a r i e d from 1 . 6 t o 7 . 4 mmlmonth. Growth r a t e s d e r i v e d f rom back- c a l c u l a t e d l e n g t h s a t mean annulus formation i n d i c a t e a narrower range o f mean monthly growth r a t e s ( n o t including the f i r s t year of -growth): 3 .7 t o 4 .5 mmlmonth, g r a y s n a p p e r (Croker 1962; S t a r c k 1971 ; Cl a r o - Madruga and Bustamente Pol a 1977) ; 2 t o 4 . 0 mmlmonth l a n e s n a p p e r (Rodriguez-Pi no 1962; Alegria-C. and d e Menezes 1 9 7 0 ) ; 3 . 3 mmlmonth, ye1 1 owta i l s n a p p e r (Thompson and Munro 1974).
I back-cal culated fork lengths (mm) of gray, lane , and ye1 lowtai 1 group. Data a r e from the following sources: (1 ) estimated from
2) Croker 1962; (3 ) Claro-Madruga and Bustamente Pola 1977; (4 ) 1972; (5) estimated from Alegria-C. and de Menezes 1970; (6 )
Species Age e l l owtai group (1 1 ( 2 1 ( 3 1 - T
COMMERCIAL/SPORT FISHERY
The F l o r i d a 1 and ings o f g r a y , 1 a n e , m u t t o n , a n d y e 1 1 o w t a i l snappers a r e c o n s i d e r a b l e , b u t a r e exceeded by t h e r e d and v e r m i l i o n snappers ( N a t i o n a l M a r i n e F i s h e r i e s Serv ice 1970-79). The f o u r species a re g e n e r a l l y c a p t u r e d by hook and l i n e w i t h c u t b a i t , b u t a r e a1 so caught by f i s h t r a p s and spea r guns i n t h e southern p o r t i o n o f F l o r i d a . A summary o f F l o r i d a ' s c o m m e r c i a1 l a n d i n g s a n d v a l u e s ( T a b l e 5 )
i n d i c a t e s i nc reased c a t c h e s o f g r a y and l a n e snapper between 1 9 6 9 and 1978, whereas m u t t o n and ye1 1 o w t a i 1 snapper l and ings have f l u c t u a t e d and show no d i s c e r n i b l e t r e n d s i n t h e same 1 0 - y e a r p e r i o d . F i s h e r y management p l ans have been p r e p a r e d f o r t he r e e f f i s h resources a long the S o u t h A t l a n t i c U.S. c o a s t , w h i c h i n c l u d e s e a s t e r n F l o r i d a ; a n d t h e G u l f o f M e x i c o , w h i c h i n c l u d e s F l o r i d a ' s west coas t (South A t1 a n t i c F ishery Management Counci l 1982; G u l f o f Mexico F i shery Management Co unc i 1 1980 ) . Da ta s u m m a r i z e d i n t h e s e
Tab1 e 5. F l o r i d a commercial 1 andings (thousands o f pounds) and ( i n parentheses) dockside values (thousands o f U.S. do1 1 a rs ) f o r f o u r snapper species, 1969-1978 (Na t i ona l Marine F i s h e r i e s Serv ice 1970-1979).
Species Year Gray Lane Mutton Y e l l o w t a i l
p l a n s i n d i c a t e t h a t c a t c h t r e n d s w i t h i n t h e l a r g e r a r e a s w e r e g e n e r a l l y t h e s a m e a s t h o s e r e s t r i c t e d t o F l o r i d a , p r i m a r i l y because t h e m a j o r i t y o f snapper i n b o t h a r e a s a r e 1 anded i n F l o r i da . I n t he south A t l a n t i c p o r t i o n o f t h e f i s h e r y , y i e l d - p e r - r e c r u i t models i n d i c a t e d t h a t g ray and ye1 l o w t a i 1 snapper a re p robab ly be ing over- f i shed.
Spor t f i s h i n g f o r these s p e c i e s i s impor tan t b u t poor1 y documented. A r e c e n t s u r v e y ( N a t i o n a l M a r i n e F i s h e r i e s S e r v i c e 1980 ) , h o w e v e r , i n d i c a t e d t h a t i n 1979 r e c r e a t i o n a l f i shermen c a u g h t a b o u t 4.6 m i l l i o n g ray and o t h e r snapper I ( exc lud ing r e d and v e r m i l i o n snapper) from t h e South A t l a n t i c and G u l f o f Mexico. I f one assumes t h a t each f i s h averaged about 1.5 I b , t h e we igh t o f t h e 1979 r e c r e a t i o n a l c a t c h o f these snapper species was 6.9 m i l 1 i o n l b . Inasmuch as t h e t o t a l F l o r i d a c o m m e r c i a l c a t c h o f t h e s e f o u r species i n 1978 was o n l y 2 . 1 m i l 1 i o n Ib., one can assume t h a t F l o r i d a ' s s p o r t f i s h e r y s u b s t a n t i a l l y a f f e c t e d t h e f i s h stocks.
ECOLOGICAL ROLE
T h e s e s n a p p e r s p e c i e s a r e gene ra l l y c l a s s i f i e d as e u r y p h a g i c c a r n i v o r e s , b u t d a t a on f e e d i n g h a b i t s a re 1 im i t ed . Gray snapper a re pu rpo r ted t o f eed on p o r t u n i d c r a b s (34.8% frequency o f occurrence), sp ide r crabs (3.9%), and o t h e r crustaceans i n t h e Dry Tortugas area (Longley e t a l . 1925). Croker (1962) found t h a t g ray snapper f rom south F l o r i d a a t e f i s h e s (34% i n frequency, mos t l y anchovies) and crustaceans (79%), which i nc l uded g raps id crabs and penaeid shrimps. Juveni 1 e g ray snapper a t e p r i m a r i l y crustaceans (93% i n frequency, most ly amphipods and shrimps) and f i s h (5%) (S ta rck 1971). The a d u l t s tended t o e a t more
f i s h (20.6% t o 61.9% i n frequency) and fewer crustaceans t han j uven i l es . Simi 1 a r d i f f e r e n c e s between feed ing o f j u v e n i l e s and a d u l t s were repo r ted o f f Cuba on an a r t i f i c i a l r e e f (Val dez-Munoz and S i 1 va Lee 1977). The gray snapper d i e t v a r i e d cons iderab ly between b iotopes. Gray snapper a re appa ren t l y noc tu rna l p redators (S ta rck and Davis 1966), a t l e a s t as a d u l t s (S ta rck 1971), and fo rage away from t h e i r r e e f h a b i t a t s a t n i g h t , whereas j uven i 1 es appa ren t l y feed d i u r n a l l y among grass beds.
L i k e gray srlapper, l a n e snapper 1 i v e i n a wide v a r i e t y o f h a b i t a t s - - e.g., f rom co ra l r e e f s i n c l e a r water t o t u r b i d , b r a c k i s h w a t e r o v e r s o f t s u b s t r a t e s ; c o n s e q u e n t l y t h e y feed on a w i d e v a r i e t y o f o rgan i sms . A summary o f 1 ane snapper f o o d h a b i t s revealed t h a t t hey feed on a v a r i e t y o f c r u s t a c e a n s and f i s h ( R a n d a l 1 1967). Rodriguez-Pi no (1962) noted t h a t o f f Cuba t h e y a t e f i sh (32% b y vol m e ) , crustaceans (28%), anne l i d s (12%), mo l lusks ( I % ) , a lgae ( 2 ) , and u n i d e n t i f i e d ma te r i a l (26%).
Mutton snapper a re noc tu rna l and d iu rna l p redators on c rus taceans and f i s h , and occur over sand, seag rass , and cora l r ubb le subs t ra tes (Randal 1 1967). Crabs make up 44.4% o f t h e i r d i e t b y vo lume i n t h e C a r i b b e a n , f o l l o w e d b y f i s h ( 2 9 . 8 % ) , a n d gastropods (13%), as w e l l as octopods, he rm i t crabs, and shrimp.
J u v e n i l e ye1 1 o w t a i l snapper 1 i v e p r i m a r i l y among seagrasses , b u t b o t h j u v e n i l e s and adul t s o f t e n feed above t h e s u b s t r a t e . R a n d a l 1 ( 1 9 6 7 ) repo r ted t h a t a d u l t s e a t crabs (23.3% by volume), shrimp (16.2%), and f i s h 1 . 9 % O f f Cuba, y e 1 l o w t a i l s n a p p e r s t o m a c h p e r c e n t v o l umes c o n s i s t e d o f f i s h r e m a i n s ( 5 3 % ) , whole f i s h (32%), crustacean rema ins ( l o % ) , who le c r u s t a c e a n s ( 7 % ) and mol lusk remains ( 1 % ) ( P i e d r a 1 9 6 9 ) . Y e l l o w t a i l snapper have p r o b a b l y t h e
most d i ve rse food h a b i t s o f t h e f o u r Sal i n i ty snappers considered here.
Natura l p redators i n c l u d e sharks and o t h e r 1 a r g e r p r e d a t o r y f i shes, i n c l u d i n g o t h e r s n a p p e r s p e c i e s . F i s h i n g m o r t a l i t y , a s w e l l a s i n a d v e r t e n t f i s h i n g m o r t a l i t y o n j u v e n i l e s caused by s h r i m p t r a w l e r s over so f t -bo t tom areas, accoun ts f o r a l a r g e p r o p o r t i o n o f t h e i r t o t a l m o r t a l i t y ( G u l f o f Mex ico F i s h e r y Management Counci l 1980).
A r t i f i c i a l r e e f s a n d o t h e r s t r u c t u r e s a re s u i t a b l e h a b i t a t s f o r these species (Has t ings e t a1 . 1976; ~ a l d e z - ~ u n o z and S i l v a Lee 1977; H a s t i ngs 1979).
ENVIRONMENTAL REQUIREMENTS
Temperature L i m i t s
Tempera tu re 1 i m i t s f o r t h e i e species a r e a p p r o x i m a t i o n s based on f i e l d o b s e r v a t i o n s . R i v a s ( 1 9 7 0 ) r e p o r t e d t h e t e m p e r a t u r e r a n g e f o r g ray snapper as 18.3" t o 27.2"C, and t h e mean as 21.7"C; l a n e s n a p p e r , 16.1" t o 28.g°C, mean 24.OPC; a n d mut ton snapper, 18.9" t o 27.8"C, mean t o 24.8"C. Gray snapper have been c o l l e c t e d i n t h e 1 3 . 4 " t o 32 .5"C range (Spr inger and Woodburn 1 9 6 0 ) . The a v a i l a b l e data f o r g r a y snapper i n d i c a t e t he lower l e t h a l l i m i t t o be between 11" and 14°C ( S t a r c k 1971) . S p r i n g e r a n d W o o d b u r n ( 1 9 6 0 ) i n d i c a t e d t h a t t h e tempera tu re r a n g e f o r l a n e snapper i n t h e Tampa Bay a r e a was 15.0" t o 27 .5 "C . U n d e r 1 abora to ry cond i t i ons , t h e p r e f e r r e d upper temperature range f o r j u v e n i 1 e y e l l o w t a i l snapper was 24" t o 30°C a n d t h e u l t i m a t e u p p e r l e t h a l t e m p e r a t u r e was between 3 3 . 5 " a n d 34°C ( W a l l ace 1977 ) . The a c c l ima- t i o n t e m p e r a t u r e s e e m i n g l y h a d n o e f f e c t o n t h e u p p e : r l e t h a l temperature.
Gray snapper e n t e r f r e s h w a t e r 1 akes and s t reams i n s o u t h F l o r i d a (Gun te r and H a l l 1 9 6 3 ) . J u v e n i l e s o f a l l f o u r s n a p p e r s p e c i e s c o n s i d e r e d h e r e u s e e s t u a r i e s a s n u r s e r y a r e a s . Gray s n a p p e r h a v e been c o l l e c t e d a t a s a l i n i t y range o f 1.0 t o 35.0 pp t , and l a n e snapper a t 1 9 . 1 t o 3 5 . 0 p p t ( S p r i n g e r a n d W o o d b u r n 1 9 6 0 ) . A d u l t s 1 i v e g e n e r a l l y o f f s h o r e i n s a l i n i t i e s approaching 35 pp t .
Subs t ra te
A d u l t snappers o f these s p e c i e s g e n e r a l l y 1 i v e n e a r c o r a l r e e f s o r o t h e r h a r d - b o t t o m f e a t u r e s , a n d j u v e n i l e s i n inshore areas. J u v e n i l e gray snapperb a r e common i n s h a l l o w da te r qrass beds. Small adu l t s t e n d t o c o n g r e g a t e n e a r h a r d b o t t o m s , herea as m a t u r e a d u l t s t e n d t o a s s o c i a t e w i t h r e e f s ( n a t u r a l a n d a r t i f i c i a l ) i n deeper o f f s h o r e waters ( S t a r c k 1 9 7 1 ) . Lane snapper o f t e n occur i n grass beds a l o n g w i t h g r a y snapper (S ta rck 1971 ) , b u t a r e more abundant as j u v e n i l e s i n s o f t - and sand-bottom areas where s h r i m p i ng i s conducted. A d u l t s t e n d t o be f ound over mud- and sand-bo t tom a r e a s i n bays and channel s.
M u t t o n s n a p p e r f r e q u , e n t l y i n h a b i t open waters; bo th a d u l t s and. j u v e n i l e s may a s s o c i a t e w i t h g r a s s beds, h u t t h e a d u l t s a l s o l i v e i n o r near p a t c h r e e f s o f c o r a l and r o c k r u b b l e and sponge p a t c h e s . Ye1 1 ow- t a i l snapper are l e s s associated w i t h hard bottoms than t h e o the r snappers; they a r e found on pa tch r e e f s and on t h e o u t e r e d g e s o f d e e p e r c o r a l r e e f s . J u v e n i l e s a1 so l i v e among grass beds and f i n g e r c o r a l ( S t a r c k 1971).
Depth
From catch s t a t i s t i c s , Rivas (1970) and Thompson and Munro (1974)- repor ted depth d i s t r i b u t i o n s where l a r g e numbers o f adu l t s o f t he f o u r species were taken. Maximum depths repor ted by F i scher (1978), however, were much g rea te r (Table 6). General l y , 1 arger snapper i n h a b i t deeper areas than small e r snapper, a1 though the re are many exceptions.
Table 6. Water depths f o r gray, 1 ane, mutton, and ye1 1 owtai 1 snappers.
Depths (m) Range
Maxi mum Rivas Thompson & Fischer (1970) Munro (1974) (1978)
Gray 31-49 0- 10 180 Lane 29-59 100-120 - - Mutton 40-59 - - -- Ye1 1 ow-
t a i l -- 20-40 70
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50272 -101 -- REPORT DOCUMENTATION I. REPORT No.
3. Seclolcnt s - icerr ion N o
PAGE I B i o l o g i c a l Repor t 82111.5~)/2.- I i . -- - .- --- -- -- 4. Totie and Subt~t le
S ~ e c i e s P r o f i l e s : L i f e H i s t o r i e s and Envi ronmental Require-
/-
' 5. R e w r t Date
i June 1986
-- 12. Spon3ortns Organttatlon Name and Addreas
ments o f Coastal F ishes and I n v e r t e b r a t e s (South ~ 1 o r i d a ) - - Gray, Lane, Mutton, and Ye1 l o w t a i l Snappers
7. Author(%)
Stephen A. Bortone and James L. W i l l i ams 9. Performan# Organ~xntlon Name and Addreas
Department of B io logy U n i v e r s i t y o f West F l o r i d a Pensacola, FL 32514
Na t i ona l Coastal Ecosystems Team U.S. Army Corps o f Engineers F i s h and W i l d l i f e Se rv i ce Waterways Experiment S t a t i o n U.S. Department o f t h e I n t e r i o r P.O. Box 631 Washington, DC 20240 Vicksburg, MS 39180
6.
8. Performmg OrganlXat6on Reot. No
10. Pm~ect/Task/Work Un8t No.
11. Contract(C) or Grant(C) NO.
IC)
13. Type of Regor( & Perlod Covered -Ti I
IS . Supplementary Notes * ) U.S. Army Corps o f Engineers Report No. TR EL-82-4
16. Abstract (Limlt: 200 words)
These four snapper species have s i m i l a r morphologies as w e l l as l i f e h i s t o r y and env i ronmenta l requirements. Wi th some except ions they a r e r e s t r i c t e d t o t r o p i c a l and s u b t r o p i c a l coas ta l areas o f t h e western A t l a n t i c . Adu l t s o f t e n a s s o c i a t e w i t h c o r a l r e e f and o t h e r hard-bottom communities. Juven i l es and p re -adu l t s u s u a l l y i n h a b i t sha l low, i nsho re areas and a r e o f t e n found i n seagrass beds. Reproduct ion u s u a l l y takes p lace o f f s h o r e i n t h e warmer months o f t h e year. Fecund i ty i s p o s i t i v e l y r e l a t e d t o female body s i z e , b u t t h e r e i s s u b s t a n t i a l v a r i a t i o n b o t h w i t h i n and among t h e species. J u v e n i l e s o b t a i n a d u l t body p r o p o r t i o n s a t about 50 mm SL. Approximate t o t a l maximum leng ths o f a d u l t s a r e 900 mm f o r g ray snapper, 446 mm f o r l ane snapper, and 750 mm f o r mutton and y e l l o w t a i l snappers. O t o l i t h s a re t h e most p r e f e r r e d body p a r t used t o determine age. Most snappers a t t a i n sexual m a t u r i t y a f t e r 3-5 y e a r s a t a s i z e range o f 180-350 mm FL. Males mature a t a sma l l e r s i z e than females. Recent commercial catches of g ray and l ane snappers have been i nc reas ing , whereas mut ton and y e l l o w t a i l snappers show no c l e a r t rends. Spor t f i s h i n g probab ly has a s u b s t a n t i a l impact on t h e s tocks o f a l l f o u r species. A l l f o u r snappers feed predominant ly on smal l b e n t h i c f i s h e s and crustaceans, b u t y e l l o w t a i l snapper feed on more p e l a g i c animals. Upper water tempera ture t o l e r a n c e l i m i t s a r e 27.2-32.5 O C and lower l i m i t s a r e 11-14 OC.
17. Document Analys~s a. Deacnpton
Reefs Growth Coral L i f e cyc les Mar ine f i s h e s S a l i n i t y Sea grasses Tempera t u r e
b. Identlfien/Open~Ended Terms
Gray snapper Lu t j anus g r i s e u s Spawning Lane snapper Mut ton snapper Ye1 l o w t a i l snapper
c. COSATl Field/Grouo
I 19. Securmlv Class IT'lls Rcoortl ' U n c l a s s i f i e d , C --- I 20. iecurhtf C h a r Vhls Page) / t2. Price
Uhl i m i t e d re lease ! U n c l a s s i f i ~ d . . I
(See ANSI-239.18) OPTIONAL FORM 272 (4-771 IFormerlv NTlS-35) Department of Commrrce
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