NOAA Technical Report NMFS Circular 448 

Synopsis of Biological Data 

on the G ru nts 

Haemulon aurolineatum and 

H. plumieri (Pisces: 

Haemulidae) 

George H. Darcy 

February 1983 

FAO Fisheries Synopsis No. 133 

u.s. DEPARTMENT OF COMMERCE 

Malcolm Baldrige, Secretary 

National Oceanic and Atmospheric Administration 

John V. Byrne, Administrator 

National Marine Fisheries Service 

William G. Gordon, Assistant Administrator for Fisheries 
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material 

mentioned in this publication. No reference shall be made to NMFS, or 

to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that N;\IFS approves, recommends 

or endorses any proprietary product or proprietary material mentioned 

herein, or which has as its purpose an intent to cause directly or indirectly 

the advertised product to be used or purchased because of this NMFS 

publication. 
CONTENTS 

Introduction ........... . . .. .... . . . . . .... . ...... . .............................................. ......... . 

1 Identity .... . ........ .. . . ....... . ... . ....... ... .......... ... . ................................... ...... . 

1. 1 Nomenclature .......... . .. . ..... . .. . .. . ... . . .. . . . . ..... . . . ........ . .. ... .......................... . 

1. 11 Valid name . . . . . . . : . .. .................................... . . . ................................... . 

1. 12 Obj ective synonymy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. I 

1 .2 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3 

1.21 Affinities . ................ . ... . ................................... .. ....................... ..... 3 

1.22 Taxonomic status .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3 

1.23 Subspec ies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4 

1.24 Standard common names, vernac ul ar names. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4 

1. 3 Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4 

1.3 1 External morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4 

* 1.32 Cytomorphology 

1.33 Prote in spec ific ity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5 

2 Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5 

2. 1 Total area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5 

2.2 Differential distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6 

2 .21 Spawn , larvae and juvenil es . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6 

2.22 Adults. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7 

2 .3 Determinants of distribution changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7 

2.4 Hybridization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8 

3 Bionomics and life history .. . ........ . .. . . .. . . ... . . . ..................................................... 8 

3. 1 Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8 

3. 11 Sexuality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . .. . . . . .. 8 

3. 12 Maturity .. . . ........... .. .. . ................. . .................................................. 8 

3. 13 Mating. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . .. 9 

3. 14 Fertilization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... 9 

3. 15 Go nads ..................................................... .. ...... . ...... .. . ................. 9 

3. 16Spawning .......................................... .... .... .. .... ..... ..... ..................... 9 

3.17 Spawn ....................................................... .. . ...... ... ...................... II 

3.2 Preadult phase ....................... .. ................................. ............................ 11 

3.21 Embryonic phase .................... .. . ......................... ... ... ............. .. ........... 11 

3.22 Larvae and adolescent phase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .......... .. ... . 11 

3.3 Adult phase .................................. ..... ......... .............. ....... ... ... ............. 17 

3.3 1 Longevity ... . .... . .. . .. ..... ............................................................. . ..... 17 

3.32 Hardiness ..... ......... . . ... ................................................................... 17 

3.33 Competitors .. . ........................................ ..... .......... ........................... 19 

3 .34 Predators .................................... . ................... ..... ...... . . . ... .............. 19 

3.35 Parasites, diseases, inj ulies, and abnormal ities .................... .. ................................... 19 

3.36 Chemica l compositi on . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 

3.4 Nutrition and growth ..... . .. ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19 

3.4 1 Feeding . ... .. . . . . . . ... . . . . . . . . .. . . . . . . .. . . . . ... .. ....... . .. . . . ... .... . . . ... . .. . . ... ...... 19 

3 .42 Food . . .... . . .. . . . .... .. . . ... . .. ... . .. . . ... . ... .. .. ........ .. . .. . ... .... . . .... . .... . .. . 19 

3.43 Growth rate. . . . . . . . . . . . .. . . ......... ... ....................................................... 20 

*3 .44 Metabolism 

3.5 Behav ior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .... . .... 23 

3.5 1 Migrations and local movements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 

3.52 Schooling .. . .... . ............... .. . ....... ........... .... .......... ........................... 2-+ 

*3. 53 Responses to stimuli 

4 Population . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24 

4.1 Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24 

4 .11 Sex ratio ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24 

4 . 12 Age composition ............ . ... ... ....... ... .................................................... 24 

4.13 Size composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ........... 25 

4.14 Subpopulations .............. . ........................................... .... .. .. .. .............. 26 

4 .2 Abundance and density ...... . ........................................... ............................ 26 

4 .3 Natality and recruitment. ........ ..... . ........................................... .................... 28 

4 .31 Reproduction rates . ............................................ ... ....... ........................ 28 

4.32 Factors affecting reproduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 

4 .33 Recruitment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. 28 
o 3.4 em 

Figure I.-Adult Haemulon aurolineatum. (From Courtenay and Sahlman 1978.) 

o 6.0 em 

Fi~ure 2.-Adult Haemulon piumieri. (From Courtenay and Sahlman 1978.) 
1.2 Taxonomy 

1.21 Affinities 

Suprageneric 

Phylum Chordata 

Class Osteichthyes 

Superorder Acanthopterygii 

Order Perciformes 

Suborder Percoidei 

Family Haemulidae 

Systematics and distribution of the family Haemulidae were 

reveiwed by Konchina (1976), who listed several previously recognized families (Gaterinidae, Pomadasidae, Prisinpomidae, Plectorhinchidae , and Xenichthyidae) now included in Haemulidae. 

Although recent usage has favored the family name Pomadasyidae, 

Haemulidae is an earlier name which should be applied (Robins et 

al. 1980). 

Generic 

Several workers have reviewed the genus Haemulon, sensu lato. 

Jordan and Swain (1884) discussed the status ofthe genus and concluded that Bathystoma of Scudder (1863 as cited in Jordan and 

Swain 1884), in which the species aurolineatum is sometimes 

placed , should not be regarded as a distinct genus. Jordan and 

Fesler (1893) likewise did not recognize Bathystoma, but placed 

Haemulon along with lutjanids in the family Sparidae. Ginsburg 

(1948) reviewed the species of Bathystoma , a genus which he considered distinct based on a difference in number of dorsal spines 

(Bathystoma having 13 and Haemulon, sensu stricto, having 12). 

Arnov (1952) excluded Bathystoma from his review of Haemulon. 

Most recent authors (Courtenay 1961 ; Konchina 1976; Courtenay 

and Sahlman 1978) have considered Bathystoma to be a synonym 

of Haemulon. 

Courtenay (1961) listed synunyms of Haemulon Cuvier: 

Diabasis Desmarest, 1823 (preoccupied in Coleoptera) 

Haemulon Cuvier, 1829 

Haemylum Scudder, 1863 

Anarsostus Scudder, 1863 

Bathystoma Scudder, 1863 

Haemulum Cope, 1871 

Brachygenys Scudder, 1875 

The following description of the genus Haemulon is derived from 

Arnov (1952), Courtenay (1961), and Courtenay and Sahlman 

(1978): body oblong , compressed , perch-like ; head profile 

strongly convex; chin with 2 pores anteriorly and a central groove 

behind the symphysis of the lower jaw; teeth conical, in a narrow 

band in each jaw, the outer series enlarged , but no canines; no teeth 

on roof of mouth; posterior margin of suborbital not exposed; preopercJe with posterior margin slightly concave and rather flnely 

serrate, none of the serrae directed forward; opercle with 1 distinct 

spine; dorsal fin single, usually with 12 or 13 strong spines and 12-

18 soft rays; pectoral fins long; pelvic fins below base of pectoral 

fins, with 1 spine and 5 soft rays; anal fin with 3 strong spines and 

fewer than 11 soft rays; soft dorsal and anal fins densely scaled to 

their margins; caudal fin forked; scales ctenoid. small to moderate, 

extending onto entire head (except front of snout. lips, and chin); 

mouth usually red within. 

Specific 

Haemulon aurolineatum 

The following species diagnosis is from Courtena. (1961): POR'lI 

lateral line scales 50-52; caudal peduncle scales 22 (9-2-11): dor..al 

fin XII + I, 14-15, usually 15; anal fill III, 9, rarely 8. pectoral tIll 

17-18, usually 17; gill rakers 24-28; longitudmal scale fO\l. belov. 

lateral line parallel to long axis of body ; body slivery-white With 

yellow to brown stripe from behind preopercle to black blotch at 

base of caudal fill; pectoral fillS chalky; other flns pale grd) to 

chalky; juveniles with lateral stripe from belund eye to caudal 

peduncle continuous with oval spot when young. eparate from 

dumbbell-shaped spot in larger individuals. 

Adults of H. aurolineatum are easily separated from most other 

species of Haemulon because they possess 13 dorsal spines. Other 

Haemulon species, except for H. striatum and H. boschmae. have 

12 dorsal spines. The three species having 13 spines may be distinguished as shown in Table 1. 

Table I.-Distinguishing characters of three species of Haemulon from the 

western Atlantic. (From Courtena} and Sahlman 1978.) 

Character H. aurolineatum H. slriaIum H ho.\chmo( 

Scales around caudal 

peduncle 22 25-26 2327 

Scale rows below parallel to obltque most I) PJrJllel 

lateral line body axis to bod} a " 

Gill rakers in 

first arch 24-28 28-34 30 "If> 

Anal fin rays 9 7-8 7-"1 

(usuall) X I 

Body shape compressed. nor compre~~ed . Lomprc.."cd 

depth 32-36% depth 26-30 '1' depth 211 ,() 

SL SL SL 

Haemulor, plumieri 

The following species diagnosis is from Courtenay (1961): pored 

lateral line scales 48-51, usually 50-51; caudal peduncle scales 22 

(9-2-11); dorsal fill XI + I , 15-17, usually 16; anal fin III. 8-9. usually 9; pectoral flns 17 ; gill rakers 21-27, usually 25; cales above 

lateral line larger than those below; body silvery-white \I. ith numcrous blue and yellow stripes on head and body; pectoral fins chalky, 

other flns gray; black blotch often present beneath free margin of 

preopercle ; juveniles without lateral stripe; caudal spot round. po . 

terior to caudal base. 

Adults of H. plumieri can be separated from all other Haemulon 

species by the larger scales above the lateral line (Courtenay and 

Sahlman 1978). 

1.22 Taxonomic status 

The taxonomic status of H. aurolineatum has not al .... 3) been 

agreed upon . Jordan and Swain (1884) named a ~pecle • H. rlmator, which they distinguished from H. aurolineatum by Its deeper 

body shape and lighter yellow color. Jordan and Fe ler (\ 93) perpetuated thi separation. Ginsburg (1948) re\lcv.ed the pe Ie of 

"Bathystoma" and concluded that rimator and aurolmeatllm are 

sub pecies of H. aurolineatum. en u lato. and nam d thud ubspecie. angustum. Ginsburg .-eparatcd hIS thre ub peelC of 

aurolineatum on the ba i of bod) depth Courtena) 1 1 

reviewed the genu Haemulon in the .... e tern (Iantl and n Id-
ered rimator to be a synonym of aurolineatum. Courtenay (1 961) 

did not treat Ginsburg's subspecies as such in his review. Konchina 

(1976) also considered rimator to be a synonym of aurolineatum in 

his discussion of haemulid systematics and distribution. 

Haemulon aurolineatum and H. plumieri are generally considered morphospecies in recent literature. 

See 1.21 

1.23 Subspecies 

Haemulon aurolineatum 

Several subspecific names have appeared in literature. Ginsburg 

(1948) concluded that three allopatric subspecies exist: H. a. 

aurolineatum from the West Indies and South and Central America, 

H. a. rimator from the coastal United States, and H. a. angustum 

from Bermuda, although Ginsburg placed all of these in the genus 

Bathystoma rather than Haemulon. Ginsburg based his sePl!-ration 

of these subspecies primarily on body depth differences. Variations 

in gill raker counts, scale counts, and some fin ray counts were also 

noted by Ginsburg, though these appearto be clinal. Southern Florida populations, especially from Key West and the Tortugas, appear 

to approach the H. a. aurolineatum form of the West Indies (Ginsburg 1948). 

Hoese and Moore (1977) mentioned that northwestern Gulf of 

Mexico specimens of H. aurolineatum are sometimes separated 

into the subspecies (or species) rimator. 

Courtenay (1961) did not distinguish subspecies of H. aurolineatum. Most recent literature does not divide H. aurolineatum subspecifically. 

See 1.22 

Haemulon plumieri 

No subspecies are recognized. 

1.24 Standard common names, vernacular names 

The accepted common name of H. aurolineatum in the United 

States is tomtate (Robins et al. 1980), and standard FAO common 

names are: English, tomtate grunt; French, gorette tomtate; Spanish, ronco jen{guano (Courtenay and Sahlman 1978). 

The accepted common name of H. plumieri in the United States 

is white grunt (Robins et al. 1980), and standard FAO common 

names are: English, white grunt; French, gorette blanche; Spanish, 

ronco margariteno (Courtenay and Sahlman 1978). 

Other common names appearing in the literature are given in 

Table 2. 

1.3 Morphology 

1.31 External morphology 

Haemulon aurolineatum 

Courtenay (1961) and Herculano Soares and de Holanda Lima 

(1966, 1967) provided morphometric and meristic data as given in 

Table 3. 

Courtenay's data were gathered from specimens from Florida, 

the Gulf of Mexico, the Virgin Islands, and Bermuda; Hercu1ano 

Tabl e 2.-Common names of Haemuloll aurolillealum and H. plumieri appearing in the literature. 

Country or a rea Name Reference 

Haemu/oJl aurolineatwn 

United States 

Caribbean 

Brazil 

Venezuela 

Puerto Rico 

Portuguesespeaking 

countries 

Tom-tate. redmouth grunt, 

caesar, jem'guano 

Tom Tate. red mouthed grunt 

yellow Tom Tate 

seize, roncador 

xira 

cuji 

mulita. mula mariquita, 

maruca 

cotinga. cotinga 

chirra 

Jordan and Fesler 1893 

Hildebrand and Schroeder 1928 

Breder 1948 

Fiedler et al. 1947 

Manooch and Ba rans' 

Manooc h and Barans' 

Erdman 1974; Manooch and 

Barans' 

Brandao 1964 

Haemulon plumieri 

United States grunt , black grunt Hildebrand and Schroeder 1928 

common grunt Beebe and Tee-Van 1928 

Key West grunt, ruby red lips Moe 1963 

flannel mouth grunt Hastings 1972 

Puerto Rico 

Haiti 

cachicata Erdman 1974 

Cuba 

Venezuela 

Area not 

pecified 

ronCO-fonco 

ronco arara 

coro coro margariteno 

boar grunt. ronco grande. 

ronco blanco 

Beebe and Tee-Van 1928 

Sauskan and Olaechea 1974 

Gutierrez B. 1976 

Arnov 19S2 

'Manooch, C. S., Ill, and C. A. Barans. 1980. Distribution, abundance, age and 

growth of tomtate. Haemulon aurolinealwlI Cuvier. along the southeastern United 

States. Unpubl. manuscr. 

Table 3.-Morphometric a nd meristic da ta for Haemuloll aurolinealum . 

He rcul ano Soares and 

Courtenay de Holanda Lima 

( 196 1) (1966, 1967) 

Range Mode Range Mode 

Predo rsal distance (% SL) 37-43 40 

Preanal distance (% SL) 67-74 70 

Body depth (% SL) 27-38 34 

Head length (% SL) 32-38 36 34.S-38.6 

Snout length (% SL) 11-17 13 

Length of upper jaw (% SL) IS-23 17 

No. dorsal rays 14-16 IS 14- IS IS 

No. anal rays 7-9 9 9 9 

No. pectoral rays 16-18 17 17-18 18 

Scales above lateral line 6 6 6-7 6 

Scales below lateral line 11-13 12 10-13 II 

Scales around caudal peduncle 22 22 20-23 22 

Total lateral line scales 49-S2 SO,S I 47-S I 49 

Total gill rakers 24-28 26,27 26-28 27 

Soares and de Ho1anda Limas' data were from Brazilian specimens. 

Other detailed morphometric data appear in Herculano Soares and 

de Holanda Lima (1 966, 1967). Ginsburg (1948) compared meristics and morphometrics of several western Atlantic populations of 

H. aurolineatum. 

Body coloration silvery white and head dusky grayish brown; 

yellow to bronze rnidlateral stripe extends length of body from 

behind opercle to base of caudal fin; large dark brown or black spot 

present at base of caudal fin; mouth red within; peritoneum black 

(Courtenay 1961 ; Bohlke and Chaplin 1968). 
Haemulon plumieri 

Body elongate, compressed, back elevated; snout long, pointed, 

l.9-2.3 in head; eye 3.9-5.2 in head; mouth large, terminal; horizontal; maxillary reaching about middle of eye and 1.9-2.05 in 

head; gill rakers rather short, 1'4-15 on lower limb of first arch; 

scales moderate size, larger above lateral line; scale series very 

oblique; 4 scale rows between origin of dorsal fin and lateral line; 

scales extending on base of pectoral fins; soft parts of other fins 

densely scaled; dorsal fin long, scarcely notched; caudal fin 

forked, upper lobe longest; second anal spine the longest, reaching 

slightly past tip of third spine when deflexed; pectoral fins moderate, 1.3-1.6 in head (Hildebrand and Schroeder 1928). 

Distance from dorsal fin origin to pectoral fin origin 4.0-4.3 in 

SL; least depth of caudal peduncle 8.5-9.6 in SL; diameter of orbit 

3.2-5.2 in head ; length of pectoral fin 3.4-4.0 in SL; scales below 

lateral line in pectoral region not enlarged; preopercular serrae 

embedded in membranous border of preopercle (Arnov 1952). De 

Lamater and Courtenay (1974) examined H. plumieri scales using 

scanning electron microscopy. 

Body silvery-white, head bronze to yellow above; belly and 

underside of head white; series of dark blue stripes, margined with 

bronze on the head , run back onto the body; margin of each scale 

bronze, posterior edge often gray; in a darker phase, silvery-white 

in center of each scale may become bronze; membranes of spinous 

dorsal fin chalky to yellowish-white; soft dorsal , caudal, soft anal 

fins brownish gray ; mouth bright red within ; peritoneum black 

(Courtenay 1961). 

Coloration changeable, except that alternating blue and yellow 

stripes always present on head ; pattern on upper third of body dominated by white or blue spot on each scale resulting in a checkerboard pattern with yellow between the spots; often a broad dusky 

region on mid-side of body, and rarely a phase with several indistinct dark lateral stripes may be observed; a black blotch may be 

present beneath free edge of preopercle (Bohlke and Chaplin 1968). 

See l.21. 

1.33 Protein specificity 

Cequea R. and Perez (1971) conducted electrophoretic studies of 

hemoglobin, esterases, and general plasma proteins of several species in three genera of haemulids from Venezuela. Although hemoglobins and esterases can be used to separate the genera Haemuion , 

Anisotremus, and Orthopristis, species of Haemulon could not be 

detected electrophoretically. Intraspecific variations in proteins 

were observed in H. aurolineatum. 

2 DISTRIBUTION 

2. 1 Total area 

Haemulon aurolineatum 

Haemulon aurolineatum is distributed in waters of the continental shelf from Cape Cod, Mass., to Brazil, including Bermuda and 

the Gulf of Mexico (Fig. 3). Although several authors (Courtenay 

1961; Randall 1968; Bohlke and Chaplin 1968) listed Cape Cod as 

the northern end of its range, this species is apparently not common 

north of Chesapeake Bay (Hildebrand and Schroeder 1928; Courtenay and Sahlman 1978), though it is the most northerly ranging 

Haemulon species (Courtenay 1961). The southern limit of the 

00' rrr--'--'I-""""J~r:='~T""l~A""-'----'----""" 

. .,. 

"'. 

O'IR---~---F:-/ 

10·fR...---+---+li{·'·: 

Figure 3.-Distribution of Haemulon aurolineatum. (Based on Courtenay 

1961; Courtenay and Sahlman 1978.) 

range lies between Cape Branco and San Sebastian Island, Brazil 

(Konchina 1976). 

Haemulon aurolineatum is seldom taken on brown shrimp 

grounds in the western Gulf of Mexico (Hildebrand 1954), yet it is 

the most common fish on the Campeche pink shrimp grounds 

(Hildebrand 1955). It also occurs offshore in areas of bottom relief, 

such as the West Flower Garden Bank off Texas (Cashman 1973), 

the Florida Middle Grounds (Smith et al. 1975), on rocky reefs off 

Tampa Bay, Fla. (Springer and Woodburn 1960), and on shelf-edge 

and live-bottom habitats off the southeastern United States 

(Struhsaker 1969). 

Haemulon plumieri 

Haemulon plumieri is distributed in coastal waters from Chesapeake Bay to Natal, Brazil, including Bermuda, where it has been 

introduced (Beebe and Tee-Van 1933), the Bahamas, and the eastern Gulf of Mexico (Fig. 4). 

Haemulon plumieri occurs on offshore live bottom along the 

southeastern Atlantic coast of the United States (Anderson and 

Gehringer 1965; Struhsaker 1969; Bearden and McKenzie 1972; 

Barans and Burrell 1976), and in the Gulf of Mexico, where it also 

occurs on offshore natural and artificial reefs (Smith et al. 1979). 

around offshore platforms (Hastings et al. 1976). and near rock 

reefs (Springer and Woodburn 1960). It is only occasionally found 

on the Florida Middle Grounds (Smith et al. 1975). The distribution of H. plumieri in the northern Gulf of Mexico closely resembles that of H. aurolineatum (Hastings 1972). Haemulon piumieri 
.0'1rT"----,---.---:r:i'-::"'II5:T"NIr;;;;",---r--.., 

... 

ZO' 

0'11+--__ :----/ 

10'1I+---+---j-'i;. 

ZO'~--_1---~--~: 

'o'IH---_1---~--t; 

Figure 4.-Distribution or Haemuloll plumieri. (Based on Courtena} 1961; 

Courtenay and Sahlman 1978.) 

also occurs in shallow inshore areas of the eastern Gulf of Mexico 

(Courtenay 1961; Roessler 1970; Grimes 1971; Grimes and Mountain 1971; Wang and Raney 1971; Hastings 1972; Livingston 1976; 

Smith 1976), southeastern Florida (Davis 1967), in the Bahamas 

(Courtenay 1961; Bohlke and Chaplin 1968), and in the West 

Indies (Randall 1968; Billings and Munro)). 

2.2 Differential distribution 

2.21 Spawn, larvae and juveniles 

Haemulon aurolineatum 

Areas of spawn and larval concentration are not well known . 

Sauskan and Olaechea (1974) reported that H. aurolineatum 

spawns mainly offshore on Campeche Bank, at depths> 50 m, and 

the eggs are carried by currents from east to west across the Bank. 

Larvae apparently settle in shallow water during metamorphosis 

(Billings and Munro footnote 3). Although not identified to species, many haemulid larvae (probably Haemulon spp.) were collected by Houde et al. 4 during plankton sampling in the eastern Gulf 

lBilii ngs. V, and 1. Munro. 1974. The biology, ecology. and bionomics of Caribbean reef fishes. Ve. Pomadasyidae (grunts). Univ. We.t Indies, Sci. Rep. ODA! 

UWI Fi.h. 3, 128 p. 

4Houde, E. D.,J. C. Leak , C. E. Dowd , S. A. Berkeley, and W. 1. Richards. 1979. 

Ichthyoplankton abundance and diversity in the eastern Gulf of Mexico. Report to 

Bureau of Land Management under Contract No. AA550-CT7-28, 546 p. 

6 

of Mexico. These larvae were most common in late winter and 

spring and at stations where water depth was < 50 m. 

Ju veni les are locally common and arc found in a wide variety of 

habitats. Randall (1963) and Stone et al. ( 1979) reported them very 

abundant on artificial and natural patch reefs in the Virgin Islands 

and Florida, respectively. They arc abo found along rocky shores, 

j etties, and pilings (Jordan and Fesler 1893; Davis 1967 ; Wang and 

Raney /971 ; Hastings 1972; Smith 1976), on grass flats or backred areas (Re id 1954; Tabb and Manning 1961 ; Billings and 

Munro footnote 3), and In deeper water over live boltom and rock 

ridges, such as off the Carolinas (Manooch and Barans'). 

Although the basic distributions of Juvenile and adult H. aurolinealum are the same (Manooch and Barans footnote 5). some differences may ex ist. at least seasonall y, III response to depth and temperature. Sokol ova (1965) reported that juveniles were not caught 

with adu lts on eastern Campeche Bank, and Sauskan and Olaechea 

(1974) found that juveniles were more common and farther offshore on western Campeche Bank than on the eastern side . 

Manooch and Barans (Iootnote 5) noted that more juvenIles 

appeared in trawl catches north of lat. 32 °30' N off the southeastern 

United States than south of It, and that juveniles occupied warmer 

water in the winter and cooler water III the summer than adults. 

Haemulon pit/mien 

Areas of spawn and larval concentratIOn are not well known 

Although some spawnIng may occur throughout the yea r, most 

Illdlvlduals appear to spawn III spring. Eggs are probably planktOnIC and hatch withlll 20 h at 24.2°C (Saksena and Richards 

1975). Haemulid larvae were collected near shore b) Houde et al. 

(footnote 4) III the eastern Gulf of MeXICO. 

Juvemles may be abundant III shallow water III a vanet) of habitats. 

Courtenay ( j 961) reported that juveniles are particularly abundant over 

grass beds at the edge of sand flats; Spnnger and Woodburn (1960) 

found young on grass flab north of Clearwater, Fla., III summer and 

fall. Tabb and Manning (1961) found Juverules up to 53 mm FL in 

Whitewater Bay, Fla., most commonJ) in clear water over Tha/assia. 

Reid (1954) collected young H. pIli/men at Cedar Key, Fla. , from September to November on moderately deep flats ""Ith mudd) sand bottom and on shallower flats from July to October. 

Juveniles are also found III associatIOn With hard substrates. 

Davis (1967) reported juveniles common among rock) shorelines 

in the Florida Keys, and Beebe and Tee-Van (1928) found young up 

to 100 mm SL on shallow reefs in Port-au-Prince Bay, Haiti. In St. 

Croix, U.S. Virgin Island, juvemles up to 150 mm SL are found in 

heterotypic schools with H. jlal'olinearum on patch reefs and are 

usually associated with the corals Porites porites and Acropora 

palmata; larger juvenile (120-150 mm SL) often form roaming 

schools on reefs and are less closely associated With particular corals than are smaller juveniles (Ogde n and Ehrlich 1977). Billings 

and Munro (footnote 3) observed small schools of unidentified 

j uvenile haemultds throughout the year in Jamaica, but particularly 

in April, May, and August at 2-25 m depths in reef areas, especially 

above heads of Mollfasrrea sp. and Madracis sp. corals, on sand 

flats among Diadema antillarum spines, over Thalassia beds, 

among mangrove roots, and near jetties. The smallest juveniles 

observed were 10-20 mm FL. Young H. plumieri change to adult 

color pattern and body form at about 40 mm SL (Billings and 

Munro footnote 3). 

5Manooch, C. S., III, and C. A. Barans. 1980. Distribution. abundance. age and 

growth of tomtate, Haemulon aurolinearum Cuvier, along the southeastern United 

States. Unpub!. manuscr., 18 p. Southeast Fisheries Cente r Beaufort Laboratory, 

National Marine Fisheries Service, NOAA. Beaufort. NC 28516. 
See 2.3 unu 3.22. 

2.22 Auulh 

/{aCllllllolI 1Il1mlill(,lI/l11/1 

L.,t.e the JuvenJle~. adult H. alimlillcullIlIl have been (lUnd tll 

occupy a wlue vanety o( habitut~ 

1/(l/JIfCll 

AllillUJI recl. 

()1I\hllr~ man In ... dc platform, 

(ira ... , h~(h 

lhn~l'n relicl olo\\n 'hnmp grounli ... 

1.1\ c hottom and ,hl'l feJgl' 'pongL' 

Jnd (orJI hJhlt~I('" 

S"Urt ( 

Smllh el al 14/<1 

Spnnger anu \\o"uhurn 141>0, Ha I 

mil' 1'172 . 'ourtena) anu SJhlmJn 

147X 

Da\l' l'In7. BIlling, anu \lunm 

lIoo,nOle .~). Smllh Ill7h. Hoc,,' 

.. nd Mo",e 1'177 CourtenJ) Jnu 

Sahl man 147X 

Beebe and fce Van I 'I~~ Da\ I 

1'167 Bohlke and ChaplIn 1'I6k 

H liucbmnd 1'I'i4 

Chillendcn and \1cEachran IlJ 71> 

Bearden and McKenlle I'ln. Baran, 

and Burrell 1970. ~IJn""ch Jnd 

Baran, (IonlnlHc 5) 

HU£'IIIUloll olirolill£'lIll11l1 may occupy a number of different 

microhabitats within a fairly restncted geographic area. such a~ reef 

tops. forereef~. backreef gras~ beds. patch reefs. and ak} nnanan 

coral patches In the Flonda Key~ (Davl~ 1967) 

HOCII/uioll llurolilli'OIIlIll occur~ from shallo\\ water to offshore 

reefs. open sandy areas. and ~hel I II\e bottom areas at least as deep 

as 91 In (Manooch and Barans fnotnote 5). though not ver] com 

1110nly deeper than 60 m (Fishery Management Plan lor the 

Snappcr-Gmuper Complex of the South AtlantiC Region') Smith 

(1976) reported that H llurolill£'ll/lllll IS one 0. the most common 

ueep-water reef fish in the ea~tern Gulf of MeXICO at depths> ,0 

m. 

HCU'lIIl1lol1 [lllIlIIien 

Like the JUlenJles. adult H. plllllllen occup~ J \ anet~ of habitats 

long the southeastern Atlantic coast of the United tJtes they 

occllr pnmanl) 01 er Ille-bottom she! f areas ( truhs'lker 190Y: 

Bamns and Burell 1(76). 

Bearden and McKenne (1972) reported H [11111111<'11 fml11 33 ~ tl) 

55 6 m (I g-,0 fathom~) on 11\ e h\)ttolll oft \)uth Can)linJ, anJ 

nden-on and Gehnnger (1965) r'ported them from e plnmtof] 

L'ru,se~ off the ~ollthea~tern ntted tates at 0-37 0 111 (0 20 lath 

om,) m \\ mter. and 37.0 1(53 m (20 100 fatl1\)(ll'o) In fall; the) 

1\ erl' absent It )m ~pnng and ummer ample Hat lIIu/on p/1II1l1 fl 

appeJf", tl) prefer shel f area~ 1\ nh ndgl's Ill' hard sull,trJtc ,ontamIng soft matenab In het\\ l'Cn (Fi,hef] ~ lanagem~ nt Plan f,lr the 

SnJppcr-Grllupcr omple of the \)lIth AtlJnttl' RCf!11 n lootnote 

6) 0 H. 1,llilllla; I\erc taken Jt . I-\R~I'\P (~Ianne Re lJU cC • 

lontt\lnng. \"e'osment and PrcJIl'lIl1nl and-txlttl m lr. II 1 t.ltl\)n. in thelluth -\tlant,· Bight (\\cnn r ct al I 7Q d.h.c.d). 

ca"(lh.m th II 

olf Junng the dJ) 111 JUlllJl . 

2.3 [)cternUllJllts of dl tnbUtlOll ch n'l,: 

HoclI/nll/on aum/m tun 

f dl tnhuu n 
Richards (1980) rcported H. llLlrolilll'llfll11l from IOshore reefs In the 

South Atlantic Bight and classified the speeie, as warm temperate 

tolerant. Olaechca and SausKan (1974) also lounu seasonal uilTcr 

ences in distribution on Campeche BanK uue to changes In hottom 

water temperature. 

Hal'lIIUloll olirolilll'llflllll preler~ f;:lIr1, high sallilit) <tnu clear 

water and appears to avoid estuanne area,. Relu ( 1(54). Tahh <tnu 

Manning (1961). Wang and Raney (1971). anu Ogren anu Brw.ilel 

(1977) reported taKing H. Gliroltllt'llfl/111 only 111 ueepel~ morL ,ailnt.: 

parts of the estuanes they studied Reid (19(\4) colleClcu 'pe~ imcns 

at 17 .5%0 at Cedar Key. but most collections rcpOlted in thc lucrature were from near or abovc 30oA)o 

Food availabihty also affect> ulStnhulllln. Ranuall (1%3) !nunu 

H GlIrolilll'(lflll1l to be much more abundanl on artificial palch rccls 

than on fringing reefs III St. John. L Virgin Islanus. anu thcll' 

rized that the difference may haw heen UUL tll fllod a\ailahilil). 

Indi\luuals undertake nocturnal migr.tllnns .1\\ a) from their u.t)-

time habltab tL forage on nearby grass anu ',II1U lIals (R.tndall 

1963; Da\ IS 1967. Smuh and T) ler 1972: Bililngs and ~ll1nrll 1001-

note 3. Hastll1gs et al. 1976) 

Eggs and larvae are probabl) pelagl\.· (BlillOg' and ~Iunrll ,,")1-

note 3) and their distribution IS Influenced h) CUlTenh 011 Cam 

peche BanK most spawl1lng aCII\it) taKes place nn the \:,1 lall part 

of the BanK and the) oung are earned tn the \\ e,t h\ currcrll 

(Sauskan and Olaechea 1974) Spa\\ I11ng and keulng MCd Iln 

Campechc Bank are also oL\.upled sea,,)nall) and.ll"c IIlllucn eu i:I) 

currents and food avaiJabiht) (SllKOlo\ a Il.Jb5. au KUll ulld ( )Iac 

chea 1974). 

Distribution is also Influenccd b) sut",trate t) P\.' Cnkarcous bOi 

tom. such as areas of shell. limc,tlll1e. or (Ither bnl\..en rellcl. arc 

preferred In the Gulf of Mexico. calearellus bllttomlliten IS Inhah 

ited by the Pill]... shnmp commumt) On br,l\\ n shnmp ground, In 

the northwestern Gulf. H. ollrolilll'£lllllll is prim;tr1I) u"pclalcd 

with small patches of broKen relict (Chittenden and 1\ kE:J\.·hl:tn 

1976). Tht: grt:ater abundance of H. 1Il1rulillCafll/Il 1)11 hllre In the 

northern Gulf of Mexico may be due to the ,can:lt! nfbrllKen-relict 

bottom close to ~hore 

Haelllllloll pll/l/lien 

There I little information on factors inllucnclng di~tribullon 

changes in H. plumier; Temperature undoubtedl) limit the range 

of this warm-water species. M iller and Richards ( 1980) reported H. 

plumieri from intermediate depth reefs in the South Atlantic Bight 

and classified the species as SUbtropical tolerant Seasonal it) exhibited by H. plllmieri m shallo\\. water in the northern Gulf of ~1e.\ico 

may be due to temperature. thiS specic~ general I) bemg absent from 

shallow water m winter and earl) spring (Reid 1954: Moe 1963: 

Gnmes 1971: Grimes and Mountain 1971: Wang and Rane) 1971. 

Hastings 1972). Haemulol1 plumier; has been found m water as low 

as l3°C in the northern GulfofMexico (Hastmg 1972). Seasonal 

movements to deeper water could also be due to spawning activity 

(ReId 1954). On Campeche Bank. H. plumieri is mfluenced by 

temperature and may seek deeper water in summer (Lyubimova 

and Kapote 1971); greatest concentrations were found from August 

to December, with larger fish at greater depths. 

Habit and food availability influence local distributions of H. 

piumieri, certain habitats such as hard substrates possibly being 

preferred (Juarez 1975). Like H. aurolineGtlltn, H. plumieri is 

often associated with calcareous substrates and broken relief (Chittenden and McEachran 1976). Diurnal-nocturnal foraging migrations account for diel habitat change (Davis 1967; Randall 1967; 

8 

Billings and Munro footnote 3; Ogden 1\)77; Ogden MH! l:hrtKh 

1(77) 

CUIn:nts prohJhly allell egg ,lfId lanai di\lnhutiofl. though 11111 

is known conceflllng lhls 

See 2.22. 1 1(,.3 :12.3 41. and 1 S I 

2.411)bndi/atlon 

Hyhlld\ hel\~eCn /I (1//1(11111/'(/111111 dill! 1/ HI /IIt/(/llill"'/, bnd /I 

(ll/mlillnl/l/lII dill! 1/ hOl/lllllllt'. h.l\c h ell reported (F. CCfvl£IIn 

pel" LIlIIlIT1Ull. a\ \.Ited in \.juea R. and P":rCl 1!J71). 

" h) hmb, III II /1/1/111/1'11 dr\: krl"" n 

lBIO 'O\lICSA DIIH.HISIOf{Y 

l I KepI' du lion 

l II e lI.lill) 

I hCfe I nu \e Ud} dllnorphl III III bod} hapc Ilrcolor In II tllllU 

IIIIt ,,//111/ or 1/ pll/lIllt II. thou ,I] mdle\ rro\\ larger Ihan female 

(BilllI1g~ :Jnl! \llInn. footn te ) .IU \..an <lnd Ulaechea (}CJ74) 

reponed th,1l n ampcchc [I.mk. klnut prcd nllnated In /I 

Illro[I/I/{/(1/1/I pc 1m n II <; 194 em !lypc 01 lenglh mca urcIllcnt ullspcclficu) m I pn:dul1un.tlcd In pcunlen I - 21 m 

(l11ed urcment un pc lfi d) dnd .Ill pcllmen 0 er~' - m \\oer 

mal (l-ig:) 

r here I no C\ IlJcnlc lf e, re\ cr.al or hermuphrodlll rn III cllher 

p\: II! 

<; ~ II 

100 

90 

80 

70 

60 

... 

Z 50 ... 

V 

QI: 

... 40 A. 

30 

20 

10 

0 

6 8 10 12 14 16 18 20 22 24 26 

LENGTH (em) 

Figure 5.-Se, composition (in percent) of Haemuloll aurolinealliin of increasing bod) length (Col) on Campeche BanI... 1964-71. "ature indhiduals appear 

to the right of the hea,) line. and immature indhiduah to tbe left. JU\ eniles are 

indicated b) un patterned areas. (From Sau,kan and Olaechea 197.l. fig. 7.1 

3.12 Maturity 

Haemulon aurolineatum transforms from juvenile to adult color 
tern and body form at ahout 5 4 (nl SL (Courtenay 1961), 

ugh sexual matunty docs not occur unlll fish arc about tWice that 

gth. Beehe and Tee-Van (1928) reported that an I I 0 cm SL 

ale and two 8.0 cm L malt.:s from Haiti appean.:d to he iO 

wn1l1g conditJOn. The smallest npe female caught In Jamaica by 

ling~ and Munro (footnote 3) was 13 .0 cm FL and 34 g and the 

allest ripe male 14.7 em FL and 55 g. Huemulol1 aurolit/ealUIIl 

Campeehe Bank mature when about 1 yr old, and all of the 

mercial catch on the Bank is ~exually mature (Sokolova 1965) 

skan and Olaechea (1974) found that H. aura/ifleolum on amhe Bank from 7.5 to 13.4 cm (mea~urement unspecified) were 

to 3 yr old and mostly immature , and specimens 13 .5 19,4 Clll 

easurement unspecified) were 3-5 yr old and about half were 

ture . Specimens 19.5-25 4 em (measurement unspCLllled) were 

rold orolderanu all were mature. Females mature fiN, atll.5-

.4 em, mainly I 1.5- 13.4 em, and males mature at 11.5-15,4 CIll, 

inly 13 .5- 15.4 cm (measurement un~pecified) (Sau~kan and 

cchea 1974). At < 9,4 cm (measurement unspecified) and I yr 

, external sexing i~ Impossihle (Sauskan and Olaechea 1974). 

(lelllllloll plumieri matures during the third year of life on Camhe Bank, at < 18 em (measurement unspecifi ed) and 0.30-0.35 

body we ight (Lyubimova and Kapote 1971). Billings and Munro 

otnote 3) found male fully mature at 24.0-25.9 cm FL, with a 

an size at maturity of 20 cm FL. Females were fully mature at 

.0-27.9 cm FL, with a mean size at maturity of 22 em FL. The 

all est ripe male found was 14.5 cm FL and 60 g, and the sma ll est 

e female was 14.3 Clll FL and 59 g (Table 4). 

Tahle 4.- Pcrcenlage, of ripe or 'penl fj,h and compari,on of 

lenglh-frcquenc) dblribulion, of male and female J/oell/uloll plumier; in Jamaican \\akp.., (From Billin~' and J\lunro ~ee te,t footnote 3, lable 3.6.1 .) 

L~ng.lh Male, FCI11i.JIc ... 

(F l. .em) 1/ c; rir~ or 'pent 1/ C:C nrc or ... rent 

12.n 1.1 I) 12 '(0) 27 0 

14 .0 15 <.j 85 4 17 1 4 

In n 17 <) 153 X 1(>1 4 

180- 19 I) 124 22 116 17 

2n 0 2 1 I) 100 27 117 34 

22 0 23 <) 66 2X 123 -II 

2-1 0 25 4 67 55 1-10 56 

26 0·27 <) 50 18 180 82 

2H.0 24 <) , X' - 12 IXX 77 

10.0 11 4 11-1 45 III (11 

32013 .9 30 47 12 (75) 

1-1 035 .Q (31) ( 100) 

3(> (1 17 Q ("0) 2 ( 100) 

3~ 0 N 9 0 0 

1l)I,,1 X8t'\ 1.11<) 

Pl.'H..:L' l1IJt!l'''' In r;lIl lllh~ ... ~, Jr~ hJ'I,.'J \lll "'~l1nrl~, (11 f(,\\I,.'r Ih.lIl ~() 

(,,11 

Nl) rccl)rd, of mating In H . 1II1rOhIlL'(1/111I1 l)J' H. "llll/l1( ri \\ Cn:' 

und o Billing, and 1unr\) (fl)otnlllc 3) ,tmcd Ihal PJJnng probabl~ 

)Cs not lak.e placc . Pairs of H. l,llill1l( n OCCaSll)Oa\l\ tacc and push 

llCh olhcl \\ Ilh ()pen mouth,. but \\ hcther thl IS court hip nr I'm-

'nalll~ I, lwt I.:no\\ n (Bo'hlkc and haplm 1<.)0) pa\\ nrng aggITatl )n, mJ~ l ccur (Billings and \1unro 1~)l)tol)tL 3) 

~ 'C 3.51 

3. 14 Ferttillalliln 

f-crtllil.tlinn 1\ proh. hi) eternal" Ilh nd perm 10 Ii 

cratcd'imuitaneou I) (BiIllOg dnd \lunro I tnllt 

3 15 Gonad 

Hacmulid gonad, arc paired, elongated 

do~al wall of Ihe hod) C<I\Il) Idlalcnt to til 

o\anes are rounded and lr..rn,lucent, ,md th teste a 

cross section and opa4ue (BIIlmgs and \ll1nro ootn lie t lllJtu 

nty thc ovane ... arc p1l1k-hrov. nand thl' Ie Ie \\.h1l 

Mean percentage of hoJy \\l'lght of O\j.\O s nf \en IITlC1lS f fI 

lIurolilleatwll (mean hod) v.elght l) I g) m<:a! un."d h) Hillin' and Munm 

(footnote 3) w~ 407% . &!be and 1~<: ,11\ 192 ) found the I c of a 

12.5 cm SL male to be 1.5 cm long and (U ern \\ Id 

GonosomatJc Indices of ripc Jdult II plllllll< n fmm 

Bank were found to bL 7.0-9 3 for females .ll1d [) 7 I 

lOpe h 

for mal 

(Lyubimova and Kapote 1971) Bdlll1gs Jnd \lunn) (footnot 

proVIded the followll1g ll1fOrm,!lIOn for femak II plll/men lrom 

Jamaica: 

Bodv Ii'eighl 

(g) 

158-250 

251 388 

Sec 4.3 1. 

3.16Spawning 

MCl/1I Alcon 

hod\ In'on 

I\'cighl 1i(,I1~hl 

(g) (g) 

209 % 

326 I () 20 

HOCll1ul(l1/ Wllvllll( tll/lll/ 

\1eoll 

p,rt maRl 

Spawning appea~ tn lake pl;Jcl (ncr IlIULh of thl: e.l[ \ Ilh 

spav.nlng peab occurring. The tllnc 01 sp'.I\\OIng 1'1..1 \an () 

gmphically. 

Manooch alld B;Jrans (fontn()le 5) reported thJt II (/uro/11l at" 

prohahl) spa\\ n, pnmanl) 111 Ilud pnng 111 th 

Bight Runnlng-ri pc males and p.lll1alh PC'lt f ma 

111 Apnl 1979: a l'ol1SldcrJhlc deL'fc.I\c In (",II) \\elght f !I! \ 

Apnl ~ampling. JU\~niles (3 1 em I i.) laken 111 

catches probabl) \\ L'fl' 'pa\\ ned thc pre\ I u 

and \\<: t 

r \\ mn I\' 
WINTER S~RING 

20 20 

N N 

21(' W 

21(' 

920 900 aao W 

SUMMII AUTUMN 

22 2 

N L-__ ~ __ ~ __ ~ ______ ~~ __ ~ 

D 200 -soo SO-l00 <so o 

SPECIMINS ~IR HOUR TRAWLING 

Fij!ure 6.-A\Cnl~< abundanc~ of 'flU" nlll~ iodi, idual, "I JlatlnultJlI aurtJ/lllealum In 1'11>-1 ~~ on ( .. mil ch .. Ban~ h\ 

,c:a"lO. (trum "au "an .Ind ()I.dllhl'a I""''', fie s .) 

in . pring. and another on the central BanI.. in 'pring. um ncr. and 

fall. 

~lunro et al (1973) collected nre fem.lles In 1.Jnuar~. Apnl. 

May, July. and August In JamaIca and obsen ~d Juveniles thr,tughout the year. B.llings and Munro (footnote 3) abo repl)rted pa\1 flIng year-round in Jamaica. \1 ith a peal.. from January thn)ugh pril 

Least spal\ ning actlY It) occurred September through December 

Peak spa~ning occurred at temperatures ,ncr ~8")C. \\1th liltk 

spawning activit) at IOYler temperatures (Billings and :-'lunro 1001-

note 3). 

Haemu/oll auro/il/earulIl reportedly \pa\\ ns from January to ~la) 

and in Jul) and August tn Puerto Rico (Erdman 1977); ) ear-round 

in Brazil (Cenigon 1966): and throughout the year in Venezuela. 

with a peak in Jul) (Cenigon 1966). 

HaeIllU/OIl p/wllieri 

Spawning appears to occur over much of the year. with a peal.. in 

spring. Smith (1968) found ripe fish near Miami. Fla .. in late 

March and early April. but not in February. On the west coast of 

Florida, Reid (1954) reported that the presence of juveniles inshore 

in fall indicated spring spawning and perhaps an accompanying 

migration to deeper water. Moe's (1966) studies on the west coast of 

Florida indicated that large schools may form for spawning . Part) 

boat catches examined by Moe (1966) had large numbers of ripe H. 

plumieri in May. some ripe and some spent fish in June , and no ripe 

individuals in July. Later tagging studies in the eastern Gulf of 

10 

\1,'1(110 (\ ~ 1f)7~) hOVh:d thaI ra\\nmg a[:![:!regallnn ()(curred 

near home reef II J!h I.ule 'l1IgrJII tn On C ampLLhL BanI... H ph/-

II/lcn rJ~n In \pnland~la) F'crmannand I r,h(l902) found 

pal' nlng H p/UIlIIl n 011 Puerto RIlo.n chool on hoal and (l\ <:r 

Ii ~~ hOI!\)111 1'1 \ugu land ·cplcmbc:r.buIErdmJn(19 .... 6 IQ~-) 

, I'll! 'PC H. p/z(//uen an Puerto RILo from F-ehruary 10 \pnl. ,md 

i1 ,p cmber and. 'n\ embcr. \\llh a pcak in , larch .. lunro CI al 

( 1 P3) fnund lImc 'pa\\ nlDg aCli, il) ~ eJr-round lD Jamal~a . \\ lth a 

ma unum an \la rch and pnl and a mlDan1Um lD Octobcr (Fig . ) 

Thc~ noted Ihal pal' ning on off. hare banI.. VI a probabl) Ie, easnnal. Billings and )'Iunnl (ll)otn,tle 3) tound the greale,t pmpor100 

80 

.. 

~ 60 v 

: 

.. 40 

20 

0 

M II. M 

1970 1971 

MONTH 

W RIPE ~ ACTIVE D INACTIVE 

Figure 7.-Seasonal ,ariation, in Ihe proporlion. of ripe. acti,e. and inactile 

gonads of Haemuloll plumieri in Jamaica (0-15 01). (From ,\lunro.1 al. 1973. 

fig. -1a.) 
tion of ripe H. plumieri in Jamaica from January through April. 

with a secondary peak in October and November. Minimum activity occurred in June, July, and December (Table 5). Haemulon plumieri appears to shed its eggs in batches over a period of time 

(Billings and Munro footnote 3). 

Tabl e S.-Monthly percentages of inactive 

(f), acti ve (A), ripe (R), and spent (S) 

go nads in samples of ma ture Haemu/o n 

p/umieri in J ama ican wa ters. (From Billings a nd Munro see text footnote 3, table 

3.4.1.) 

Percentage 

Month 11 A R S 

Januar) 157 38 18 38 6 

Februar) 176 31 29 34 8 

March 120 28 38 25 9 

Apnl 275 7 8 47 38 

Ma) 222 33 35 21 \I 

June 163 55 34 4 7 

Jul) 155 52 32 \I 5 

August 147 51 27 19 3 

September 174 72 13 5 10 

October 199 56 \I 22 \I 

No\ember 262 39 00 22 17 

December 217 58 26 9 .. Total 2.267 

3.17 Spav.n 

Although no specific information was found on the eggs of H. 

aurolilleatlll11. BIllings and Munro (footnote 3) provided general 

information on haemulid eggs. Pnor to being spawned. the eggs are 

,pherical. opaque. negatively buoyant in water, and about 0.5 mm 

In diameter. Ripe ovarie\ of most grunb examined had two egg 

size-classes-large ones ready to be spawned and smaller ones 

probably to be shed the following season. At ovulatlon, which 

probably take\ place shortly before spawning. the eggs increase in 

size and become transparent. 

Ripe ovarian eggs of H. plumieri are about 0.50 mm in diameter 

(Billings and Munro footnote 3) and the spawned egg 0.90-0.97 

mm in diameter (Saksena and Richards 1975). The eggs are 

un culptured, contain a single oil globule 0.22-0.24 mm in diameter. and have a narrow (0.03 mm) perivitelline space and a transparent chorion (Saksena and Richards 1975). 

3.2 Preadult phase 

3.21 Embryonic phase 

Although there is no information available regarding H. aurolineatum. the eggs are probably pelagic. Hatching time is 

unknown. 

Eggs of H. plumieri are probably pelagic and hatch in the laboratory in < 20 h at 24.2°C (Saksena and Richards 1975). 

3.22 Larvae and adolescent phase 

Haemulon aurolineatum 

There is little information on larvae of H. aurolineatum. 

Manooch and Barans (footnote 5) reported that identification of H. 

aurolineatum larvae was not possible at that time. Length of planktonic life of the larvae is not known, but may be short. Munro et al . 

11 

(1973) did not find haemulid larvae in open-ocean plankton samples taken near Jamaica, though Guitart Manday (197 1) reported 

haemulid larvae (5 .0-7.0 mm TL) in plankton hauls taken between 

o and 50 m, at night. 

Juveniles are often found in shallow water, though distinguishing 

species is diffic ult in small individuals. The smallest juvenile 

grunts observed by Billings and Mu nro (footnote 3) in Jamaica 

were 1-2 cm (measurement unspecified), with small schools of 

juveniles present year-rou nd, but especially in April, May, and 

August. Young school on nursery grounds or on reefs near adult 

schools and over Thalassia beds where they congregate at edges of 

sand patches (Billings and Munro footnote 3). 

Courtenay (196 1) described and illustrated juveniles from 18.5 

to 53.9 mm SL. Head and caudal colorations are of prime importance in separating juvenile Haemulon species (Fig. 8); Courtenay's description of H. aurolineatum juvenile pigmentation 

follows: caudal spot and lateral stripe connected by scattered, dark 

melanophores in young individuals, later becoming separated; as 

growth proceeds, the oval spot is slightly constricted over the caudal base, spreading onto the caudal. 

Juvenile H. aurolineatwn change to the adult body form and color pattern at lengths over 5.4 cm SL. Thmsforrnation also involves a change in 

feeding strategy from diurnal plankton feeding to noctumal benthic foraging in most grunts (Billings and Munro footnote 3). 

Haemulon plumieri 

Saksena and Richards (1975) reared H. plumieri in the laboratory 

and described larval development. At hatching, larvae were 2.7-

2.8 mm SL with a large ellipsoid yolk sac and white pigment scattered over the body and yolk sac. Some yolk and the oil globule 

remained 24 h after hatching, but were almost completely absorbed 

after 48 h, when larvae were actively feeding. The following 

description of larval development is from Saksena and Richards 

(1975) (Figs. 9-12). 

Morphology 

Body depth at the pectoral fin symphysis 9.4 % SL just after yolk 

absorption. 20% SL in 5.8 mm larvae,and 32.9% SL in 23.4 mm 

juveniles. Growth in the laboratory was most rapid during the first 

30 d after hatching but slowly declined afterwards. Larvae grew 

from about 2.7 mm SL at hatching to 10.0 mm SL at 20 d, and 12.6 

mm SL at 30 d, with an average daily growth rate of 0.32 mm/d. 

Standard length was about 96% TL for newly hatched larvae, but 

82 % TL for larvae 6.0-8.0 mm SL, and 80% TL for larvae larger 

than 12.0 mm SL. The larval gut was relatively short. A detailed 

table of morphometric data was presented by Saksena and Richards 

(1975) for larvae of various ages. 

Pigmentation 

Newly hatched larvae had several small melanophores from the 

snout to above the eyes, a few large stellate melanophores on the 

anterior edge of the yolk sac, several others along the gut and on the 

dorsal and ventral edges of the myomeres in the trunk region, and 

two small melanophores on the ventral edge and one on the dorsal 

edge of the notochord near its tip. The eyes began to develop pigment 1 d after hatching and were completely pigmented by the following day. 

At 3.7 mm SL, melanophores became larger and more numerous, though pigment was absent from the head region. A few large 

stellate melanophores were on the dorsal edge of the gut and were 
4.4 Mortality and morbidity ..... . ... . ..... . ..... . . ...... ................................................. 29 

4.5 Dynamics of population ............... . ......... . ............................... . .................... 29 

4.6 The population in the community and the ecosystem. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ..... . 3 J 

5 Exploitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ....................... . 31 

5.1 Fishing equipment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .......... .......... . ................... 31 

5.2 Fishing areas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ................... 3 I 

5.3 Fishing seasons ................................................................................ . 32 

5.4 Fishing operations and results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ..... ........... . ............. 32 

5.41 Effort and intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ............. 32 

5.42 Selectivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ........... ........... . ....... 32 

5.43 Catches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ..... . ................... 32 

6 Protection and management . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ................................................ 34 

6.1 Regulatory measures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ........... ....................... . .. 34 

*7 Pond fish culture 

Acknowledgment ..................................... ............... . ... . .......................... 34 

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .......... . ............. . 35 

*No information available. 

IV 
Synopsis of Biological Data on the Grunts 

Haemulon aurolineatum and H. plumieri 

(Pisces: HaemuJidae)l 

GEORGE H. DARC~ 

ABSTRACT 

Information on the biology and fishery resources of two common species of western Atlantic grunts, 

Haemulon aurolinealum and H. plumieri, is compiled, reviewed, and analyzed in the FAO species synopsis 

style. 

INTRODUCTION 

Grunts are often given little attention in discussions of western 

Atlantic marine fisheries because of their small size. Yet some species, 

including the tomtate, Haemliion aurolineatllm, and the white grunt, 

H. plumieri, are among the most abundant fishes on reefs, live-bottom 

shelf areas, and in a variety of inshore habitats in the Caribbean Sea, the 

Gulf of Mexico, and along the sou thea tern Atlantic coast of the United 

States. Due to their nocturnal foraging migrations, they are important 

energy importers to reef communities and are probably major prey of 

many larger species, such as snappers and groupers. Grunts are among 

the fishes most caught by recreational and subsistence fi hermen in this 

geographical area. They are quality food fish and are commercially 

important in areas such as Cam~che Bank. Exploratory trawling in the 

northeastern Gulf of Mexico uggest that potentIally exploitable populations of grunts may exist there. This paper summarizes information 

on these species. 

1 IDENTITY 

1.1 Nomenclature 

1.11 Valid name 

Haemulon aurolineatum Cuvier, in Cuvier and Valenciennes, 1829 

(Fig. 1). 

Tomtate, Haemulon aurolineatum Cuvier, in Cuvier and Valenciennes, 1829:237-238, type localities: Brazil, San Domingo. The 

name comes from the Greek haima (blood) and oulon (the gums), 

referring to the red mouth interior, and the Latin aurum (gold) and 

Iineatus (marked with lines), referring to the yellow body stripes of 

adults. 

Haemulon plumieri (Lacepede, 1802) (Fig. 2). 

White grunt, Haemulon plumieri (Lacepede 1802:480), type 

locality : America. The name comes from the Greek haima (blood) 

and oulon (the gums), referring to the red mouth interior. Named 

after Father Plumier, a French naturalist who sent drawings of this 

species to Europe from Martinique (Jordan and Fesler 1893). 

IContribution No. 82-30M of the Southeast Fisheries Center. 

2Southeast Fisheries Center, National Marine Fisheries Service, NOAA, 75 Virginia Beach Drive, Miami , FL 33 149. 

1.12 Objective synonymy 

The following synonymies are based on Courtenay (1961): 

Haemulon aurolineatum Cuvier 

Haemulon chrysopterum Cuvier, 1829 (misidentification, in part) 

Haemulon chrysopteron. Cuvier in Cuvier and Valenciennes, 1829 

(in part) 

Haemulon aurolineatum Cuvier in Cuvier and Valenciennes, 1829 

Haemulon chrysopteron. DeKay, 1842 (misidentification, in part) 

Haemulon jeniguano Poey, 1860 

Bathystomajeniguarno. Scudder, 1863 

Bathystoma chrysopterum. Scudder; 1863 

Haemulum aurolineatum. Cope, 1871 

Diabasis aurolineatus. Goode and Bean. 1883 

Diabasis chrysopterus. Bean, 1883 

Diabasisjeniguano. Bean, 1883 

Haemulon rimator Jordan and Swain, 1884 

Bathystoma aurolineatum. Jordan and Evermann, 1896 

Haemulon (Bathystoma) aurolineatum. Metzelaar, 1919 

Bathystoma aurolineatum aurolineatum Ginsburg, 1948 

Bathystoma aurolineatum rimator Ginsburg, 1948 

Bathystoma aurolineatum angustum Ginsburg, 1948 

Haemulon plumieri (Lacepede) 

Guabi coarca brasiliensibus Marcgrave, 1648 

Perca marina capite striato Catesby, 1743 

Percaformosa Linnaeus, 1766 (in part) 

Labrus plumierii Lacepede, 1802 

Haemulonformosum. Cuvier, 1829 (in part) 

Haemulon arcuatum Valenciennes in Cuvier and Valenciennes, 

1833 

Labrus plumieri. Cuvier, 1834 

Haemulonformosum. DeKay, 1843 (in part) 

Haemulon parae. Castelnau, 1855 (possibly misidentification) 

Haemulon arara Poey, 1860 

Haemulon subarcuatum Poey, 1861 

Haemylumformosum. Scudder, 1863 

Haemylum arara . Scudder, 1863 

Haemulumformosum. Cope, 1871 

Diabasisformosus. Goode and Bean, 1883 

Diabasis plumieri. Bean, 1883 

Haemulon plumieri. Jordan, 1884 

Haemulon plumierii. Meek and Hildebrand, 1925 
A 

B 

c 

D 

"7"'--:;:-""'--~--"'-:- - - - __ _ 

.'. ------------

"., :1 ; .. : ..... ','.' . 

.. .. " 

... . " .... .. ; 

••• " • •• ', ,',',', ', •••• & ,',' •• - - '- -

f~1i~ --' 

Figure S.-Head and caudal regions of juvenile Haemu/oll auro/illealum . (From Courtena) 1961 , figs. l3b-d, 14a.) 

12 
A 

B 

c 

D 

Figure 9.-Larvae of Haemulon plumieri: A. newly hatched, 2.7 mm SL; B. 1 d after hatching, 3.2 mm SL; C. 4 d after hatching, 3.7 mm SL; D. 6 d after hatching, 4.6 mOl 

S1.. (From Saksena and Richards 1975, figs. 1-4.) 

13 
A 

B 

c 

--- " 

-"01 

.~ ~ 

~ 

.... :. ..:. .... ;". 

Figure 10.-Larve of Haemuloll plumieri: A. lateral and ventral vie\\s of larva 9 d after hatching, 5.8 mm SL; B_ lateral and 

ventral vie\\s of larva 12 d after hatching, 8.2 mm SL; C. 13 d after hatching, 9.1 mm SL. (From Saksena and Richards 1975, 

figs. 5-7.) 

14 
.' 

.............. ' 

Figure I I.-Lanae of Haemuloll plullllen: \ . doc,aI.lateral. and Hntral ,ie," uflana I, d after hat hID .9 m 

hatching. I L5 mill Sl.. (From a\..,cna and Rilhard I ~':. [j . .) 

15 

1 B 

.... " 

d ft r 
A 

c 

" ..... .~ ~~: ~ . . ~ .. ", '" .. . \ ' .. . .... .~- ~ . ' ... 

#- •• " 

. .. : ~ 

~. 

.' 

Figure 12.-Juveniles or Haemulon plumieri: A. 40 darter hatching , 13.6 mm SL; B. 102 d after hatching, 23A mm SL; C. 107 d after hatching. 38.5 mm SL. (From 

Saksena and Richards 1975. figs. 10·12.) 

16 
almost continuous with a series of melanophores arranged along the 

ventral edge of the myomeres. A small melanophore was present on 

the ventral edge of the notochord near its tip. Several large stellate 

melanophores were scattered on the ventral edge of the gut and a 

large melanophore was present on the anterior edge of the anus. 

Pigmentation changed little from 3.7 to 8.2 mm SL, but melanophores became more numerous between 8.2 and 9 .1 mm SL, with 

concentrations of pigment developing on the snout, premaxillaries, 

and lower jaw, and a midlateral stripe of melanophores ending in a 

dark spot on the caudal peduncle and fin . A series of large stellate 

melanophores was present posterior to the anus on the ventral edge 

of the body and at the base of the second dorsal fin . 

At 9.8-1:2.5 mm SL, additional stellate melanophores formed on 

the dorsal part of the body and completed a dorsal body stripe . The 

midlateral stripe darkened and the caudal spot grew larger. 

At 13.6 mm SL, the pigmentation had increased in the form of 

scattered melanophores over the entire body surface, including the 

fins. The dorsal and midlateral body stripes and the caudal spot 

were more pronounced , and a third body stripe developed between 

the first two stripes, extending from the eye to posterior of the first 

dorsal fin. 

Between 23.4 and 38.5 mm SL, pigment was less intense and no 

definite stripes were present. Except for the caudal spot there was 

no definite body pattern. 

Fins 

Newly hatched larvae had a prominent finfold which persisted, 

in part, until 5.7 mm SL. Fin ray development was completed 

between 9 .5 and 10.0 mm SL. Fan-shaped pectoral fin without 

rays developed I d after hatching, at 3.2 mm SL. Rayed fi ns developed in the sequence: caudal, anal and second dorsal, first dorsal 

and pelvics, and pectorals. The full compl ement of principal caudal 

rays (9 + 8) was present at 7. 8 mm SL, with secondary rays starting 

to develop at 7.9 mm SL. The anal fin began to form at 5.7 mm SL, 

with the first fin elements arising at 5 .8 mm SL. Although the full 

complement of 12 anal fin elements (II, 10) was present at 6.9 mm 

SL, the normal adult arrangement (III, 9) was nut present until 38.5 

mm SL. The third anal spine J eveloped first as a segmented ray. 

Elements of the second dorsal fin began to develop at 5.8 mm 

SL, reaching their full complement (I, 15) at 7.9 mm SL. Pelvic 

fins appeared as buds at 7.9 mm SL, with the spine developing at 

8.3 mm SL, and all elements (1. 5) fo rmed by 9.5 mm SL. Spine 

development in the first dorsal fin began at 7.9 mm SL and reached 

the full complement (XI) by 9.5 mm SL. The first pectoral rays 

were discernibl e at 7.9 mm SL, with the full complement (16 or 17) 

reached by 9.8 mm SL. Pectoral fin development was dependent 

more on age than length of the larvae. 

Saksena and Richards (1 975) also desc ribed osteological development of the c ranium, fin supports, and vertebral column, and 

squamation . 

Courtenay (1961 ) desc ribed and figured j uven ile pigmentation of 

H. p/ulllieri (Fig. 13). Above about 40 mm the young have essentially the adult color pattern . Unlike mo t Haelllu/on species, H. 

p/Lllnieri juvenil es lack lateral stripes. 

See 1.21. 

3.3 Adult phase 

3. 31 Longevity 

Courtenay (1961 ) reported that matu re adult H. auro/inearllm 

average 11 .5- 17 .5 cm SL. the largest specimen he examined being 

J 7 

18. 5 cm SL. Much larger adult~ ha\e mce b CII report d Sok 

lova (1965) fo und that H. auro/illcalum reach 30 em (m~a uremcm 

unspecified) and li ve more than 5 )ir on :.tmpclhc SilOk. Sau k.Jn 

and Olaechea (! 974) fou nd that H. allm/incarlllll rna) n:~ h 7 n Id 

on Campeche Bank. The largest specimen cnllcct~ h)' Billing and 

Munro (footnote 3) in Jamaica was 19.5 em SL. a )mptotll lork 

length estimated for Jamaican H aurolillcollll1l \\..Is 2", () m rh 

largest indi vidual sampled by Manooch and Bamns ( J 974) 1fT th 

southeastern United States was 28.9 cm TL and 9) r okl Hlkk 

brand and Schroeder (1928) reported J maxImum \\ eIght nf ahout 

0 .45 kg (lib) for H. auro/inealul1l in the Chesapcake Ba) regillfl. 

and Konchina (1977) gave a maximum weight of about () .5 kg 

Haemulon plumieri reaches about 45.0 cm and 2 5 kg (Konehma 

1977), though most specimens are much smaller. The large t 

reported taken on Campeche Bank (Kapote 1971) was a -' I 5 ~ III 

male that was more than 6 yr old. Manooch (1976) reported that 

males off the Carolina coast of the United States reach 12 ) r, and 

females 9 yr, and calculated a theoretical maximum length (L ) of 

64.0 cm TL. The largest H. p/umieri taken m Jamaica b] Billings 

and Mun ro (footnote 3) was 40.6 cm FL, the calculated as) mptotle 

fo rk length for Jamaican specimens being 42.0 cm. 

See 3.43 and4. 13. 

3.32 Hardiness 

Haemulon aurolineatum, though basically a WaIll1-water spoxles, IS 

more tolerant of low temperatures and ranges farther north on the ca.,t 

coast of the United States than any other haemulid, except possibl) H. 

plumieri (Billings and Munro footnote 3). Haemulofl aurolilleatwn and 

H. plumieri were the only grunts to remain in shallow water oft Flonda 

in the cold winter of 1957 (Courtenay 1961), though H. aUmlifll'llIIlJII 

has been reported killed during cold waves at temperatures belnv, abnul 

14°C (Galloway 1941). Manooch and BardllS (footnote 5) n:ported 

them absent at temperatures below about 10°C and rare bclov, l3 eC 

off the southeastern United States. Young may be ~omewhat mon: tol 

erant of temperature and alinity extremes than adult~ (Billing and 

Munro footnote 3), though Manooch and Barans (footnote 5) noted that 

juveniles were found at more moderate temperatures than adulh off the 

coast of the southeastern United State. Haemu/ofl aurolincolllll1 Wd 

one of the few grunts observed by Bulings and Munro (footnote 3) III 

polluted Kingston Harbor, Jamaica. 

Haemu/on plumieri has been collected m water as cnld a~ 13 C 

(Hastings 1972) and as warm as 35°C (Wang Jnd Rane)' I 1}71) 

Cl)ld deaths of H. plumieri were reported at SaOlhd blJnd. Fla 

(Storey 1937); Key West, at a water temperature of 14°C (GJllo 

way 1941); and Indian River lagoon, Fla ,at a v, atc:r temperature of 

11.1 °C (Gilmore et aI. 1978). 

Haemu/oll p/umieri is falrl) tolerant of s..lllnJt~. being lound from 

relatively low salinities (180/00) to 0\ er 37°00 (Roe ler 19 0) 

Juveniles may have greater tolerance 01 allllll:V nJ tl:mpcrdtun; 

than adult~ (Billings and Munro footnote 1) 

Although Springer and WOl)dburn (11}60) It ted If plllllll r 

one of the species killed b) a red lIde off Pmella ount r I 

1957, Smith (1976) reported thJt It \\ a one of the fi he In hall \\ 

water in the eastern Gulf of I\1exlco that WIth t ud th red lldc f 

1971. Smith ( 1976) speculated that the u~.:e of If pi In It r dur 

ing and after the red tide ma) haH: bLLn due to It n n pc If l h b 

tat needs and relatl\ e lack of. !net lle to the ub trate 

Billing~ and I\1unro (footnote 3) fLlUnd that O\t:f f If P un I 

entenng tmps in JamaIca died \\ Ithln 2 V, k of emenn the lm 

See 223 
• 

A 

B 

........... , .. . .... ...... . 

c --

D 

Figure B .-Details of head and caudal base pigmentation in young Haemuloll plumieri: A. 19.8 mm SL; B. 23.2 mm SL; C. 35.6 mm SL; D. 

42.5 mm SL. (From Court enay 1961 , figs. 12b-d, l3a.) 
3.33 Competitors 

The most important food competitors of grunts are porgies', goatfishes, wrasses, hogfishes, and mojarras (Randall 1967; Billings 

and Munro footnote 3). Interspecific competition for food is probably not great due to the variety of food items consumed and because 

each species favors a particular food type (Billings and Munro footnote 3). Some competition may exist for schooling sites on reefs, 

though competition for both food and space resources may be 

reduced through diurnal-nocturnal migrations, and habitat differences between juveniles and adults. Although small grunts are usually not territorial, large individuals may vigorously defend 

territories up to 2 m in diameter, intraspecifically (Motchek and 

Silva Lee 1975). 

Haemulon plumieri often associates with other species and may 

school heterotypically. Davis (1967) found H. plumieri with H. 

album, H. melanurum, and H. parrai near hard substrates and with 

H. aurolineatum over grass patches near Alligator Reef. Ogden and 

Ehrlich (1977) reported young H. plumieri schooling with H. 

jlavolineatum and Mulloidichthys martinicus in the Virgin Islands. 

See 3.51. 

3.34 Predators 

Haemulon allrolineatum is preyed upon by a wide variety of 

predators. Randall (1967) listed H. aurolineatum from the stomachs of several other reef species: Gymnothorax moringa , Aulostomus maculatus, Lutjanis analis, L. cyanopterus, L. jOCll, 

Ophichthus ophis, Epinephelus striatus, and Seriola dumerili. 

Haemulon aurolineatul1l has been found in stomachs of Scomberomorus maculatus off Ceara, Brazil, (Ferreira de Menezes 1970) 

and in adult Pagrus pagrus off the Carolinas (Manooch 1977). In 

Jamaica, grunts are preyed upon by serranids, lutjanids, and 

carangids (Billings and Munro footnote 3) and in Cuba by serranids 

and sphyraenids (Valdes Munoz and Silva Lee 1977). 

Hael1lulon aurolineatum may be important' n diets of other commercially valuable fishes. Springer and Woodburn (1960) reported 

that a specimen of Epinephelus morio taken on a reef off Tampa 

Bay, regurgitated hundreds of small H. aurolineatul1l. Packer (pers. 

commun. as cited in Manooch and Barans footnote 5) found young 

specimens approximately 33 mm long (TL) in stomachs of black 

sea basses, Cenrropristis striata, off North Carolina near ridges on 

the shelf. 

Predators of H. plumieri include groupers (Epinephelus spp.) and 

lizardfish (Synodus spp.) (Ogden and Ehrlich 1977); Spanish 

mackerel, Scomberomorus maculatus (Ferreira de Menezes 1970); 

dog snapper, Lutjanus jocu (Randall 1967); and sharks (Billings 

and Munro footnote 3). Other suitably large piscivores probably 

also prey on H. plumieri. 

No information regarding predators on eggs or larvae is available. 

3.35 Parasites, diseases, injuries, and abnormalities 

Trematodes have been found in the caecum and intestine of H. 

aurolineatum: Genolopa ampullacea from Gulf of Mexico specimens (Manter 1954) and Hamacreadium osciTans, Lasiotocus 

longicaecum, and L. longovatus from Jamaican specimens (Nahhas 

and Cable 1964). 

Overstreet (pers. commun. as cited in Smith 1968) reported that 

H. plumieri in Biscayne Bay, Fla., contained monogean, digean, 

acanthocephalan, nematode, and cestode parasites. Billings and 

Munro (footnote 3) found cestode larvae (c. f. OTObOThrium dipsacum) in bladders of H. plumieri from Jamaica, as well as nematodes 

in gut mesenteries and ovaries. 

Billings and Munro (footnote 3) mentioned haemulid injuries 

which probably were the result of predatory attacks. Gudger (1933) 

reported a pug-headed specimen of H. plumieri. 

3.36 Chemical composition 

Korzhova (1965) analyzed the composition of H. aurolineatum 

from the Gulf of Mexico and Caribbean Sea and found that 34.5 % 

of the total fish weight was skin and flesh and 41.5 % of the total 

weight was head. Flesh contained 20.6% protein and had a caloric 

content of 97 caloriesll 00 g wet weight. Composition of the whole 

fish was 72.4% water, 17.2% protein, 4.5% fat, and 4.9% ash. 

Edwards (1973) found the content of homogenized H. aurolineatum from Venezuela to be 67.51 % protein, 22.73% ash, 9.36 % 

lipids, and 0.59% carbohydrates. Total caloric content was 4.762 

kcal/g dry weight. 

Detailed studies on protein digestion, absorption, and peptidase 

activity in H. plumieri were conducted by Smith (1967, 1968, 

1970). Earlier studies by Pierce (1936) found digestion time in H. 

plumieri to be 25-33 h. Composition of H. plumieri is 77.3 % water, 

19.4% protein, 0.7% lipids, and 1.6% ash (Gutierrez B. 1976). 

Cequea R. and Perez (1971) studied plasma proteins and hemoglobins of H. aurolineatum and H. plumieri, using electrophoresis. 

3.4 Nutrition and growth 

3.41 Feeding 

Haemulon aurolineatum and H. plumieri are primarly benthic 

feeders, although their larval and early juvenile stages may feed 

higher in the water column . 

Foraging of H. aurolineatum is usually on sand or grass flats near 

reefs, primarily at night (Randall 1963 ; Davis 1967; Collette and 

Talbot 1972; Smith and Tyler 1972; Ogden and Ehrlich 1977; Parrish and Zimmerman 1977). Hastings et al. (1976) reported similar 

nocturnal foraging near offshore platforms in the Gulf of Mexico 

and noted the significance of this behavior in importing biomass to 

reef or platform communities. Sauskan and Olaechea (1974) 

reported that feeding of H. aurolineatum on Campeche Bank was 

most intense in winter and spring and low during spawning. 

Haemulon plumieri also feeds primarily at night, migrating off 

reefs to forage in seagrass beds and over alcyonarian patches (Randal11963, 1967; Davis 1967; Motchek and Silva Lee 1975; Ogden 

1977; Ogden and Ehrlich 1977). Hastings et aI. (1976) found that 

H. plumieri feeds on attached material on pilings around offshore 

platforms in the northern Gulf of Mexico. Very small juveniles pick 

plankton from the water column during the day (Ogden and Ehrlich 

1977). 

See 3.51. 

3.42 Food 

Haemulon aurolineatum 

Haemulon aurolineatum is a generalized carnivore, preying primarily on benthic invertebrates of moderate to very small size; 

small juveniles feed primarily on zooplankton (Randall 1967). 

Beebe and Tee-Van (1928) reported that H. aurolineatum is omnivorous, their specimens from Port-au-Prince Bay containing sand, 
mud and bottom detritus, algae, worms, mollusks, copepods, isopods, shrimps, small crabs, and plankton, but Randall (1967) felt 

that the algal and detrital material they reported was probably 

ingested incidentally along with invertebrates. Longley and Hildebrand (1941) noted that H. aurolineatum is less intimately associated with coral habitats than some other grunts and found 1,000 

planktonic copepods in the stomach of a single 50 mm SL specimen. 

Davis (1967) found alpheid shrimps, stomatopods, ostracods, 

arnphipods, isopods, hermit crabs, copepods, fish, polychaetes, 

gastropods, and sand in H. aurolineatum collected in the Florida 

Keys. Randall (1967) examined guts from Puerto Rico and Virglll 

Islands specimen (Table 6). Edwards (1973) reported polychaetes, 

amphipods, and mollusks as the main foods of H. auro/ineatum In 

Venezuela. Sauskan and Olaechea (1974) reported the diet of H. 

aurolineatum on Campeche Bank as about 70 % cru taceans (squIllids, brachyurans, penaeids), 18 % polychaetes, 7% blue-green 

algae and 5 % small fish. Valdes Munoz and Silva Lee (1977) 

found H. aurolineatum feeding on polychaetes, unidentified eggs, 

amphipods, and mollusks on artificial reefs off the outhwest coast 

of Cuba. 

Table 6.-Stomach con tents of Haemu/oll auro/mealum from Puerto Rico and 

the L.S. Yirgin Islands. (From Randall 1967.) 

Food 

Shnmps and shrimp lanar 

Polychaetes (Chlocia sp.) 

Unidentified crustaceans 

Untdentified eggs 

Hermit crabs and Ian ae 

Crabs and crab larvae 

Amphlpods (Ampelisco sp .. £/asmopIlI sp £lIrnlht"lls sp. 

Megamphoplls sp., PhOliI sp.) 

Cope pods (Unciillil/a w/gariI) 

Gastropods (A/milia <lUberrUlIi1. CaeculII pllh"hel/lIl11. Rt'lu\(/ sp I 

Pelecypods (So/elllm oceiciCll{(Jlis) 

Barnacle larvae 

Scaphopods (Cadll/us oem. Delllo/Illm sp ) 

Isopods 

V()lum~ /~l 

33 I> 

11 () 

8 H 

n.::! 

44 

) 7 

1 4 

2 5 

2 I 

10 

0.9 

04 

0.2 

About 44 % of the specimens from Puerto Rico examined by 

Parrish and Zimmerman (1977) contamed crabs, primarily the xanthid sandflat crab Micropanope lobifrons, but also Portunus spin icarpus, Portunus sp., and Pinnixa. Also frequently abundant in the 

guts were shrimp, such as the carideans Leptoche/a serratorbita , 

Sicyonia, and Latreutes, and alpheids. The squillid Pseudosquil/a 

ciliata; ascidians (mostly Microcosmus helleri); the bivalve mollusks Musculus latera/is, Tellina consobrina, and Pitar sp.; arnphipods (mostly Chevalia aviculae); polychaetes; tanaids; and isopods 

were also found . 

Haemu/on piumieri 

Haemulon plumieri is also a generalized carnivore, adults feeding particularly on benthic invertebrates (Manooch 1976). Beebe 

and Tee-Van (1928) reported H. plumieri collected in Haiti contained echinoderms, polychaetes, mollusks, shrimp, crabs, and fish 

(including Diodon hystrix). Haemulon plumieri from Alligator 

Reef contained polychaetes, majid crabs, alpheid shrimps, isopods, 

fish, and sand (Davis 1967). Parrish and Zimmerman (1977) found 

crabs and amphipods in specimens of H. plumieri from Puerto 

Rico. Randall (1967) and Valdes Munoz and Silva Lee (1977) presented detailed accounts of food items of H. piumieri in Puerto 

Rico and the Virgin Islands, and Cuba, respectively (Tables 7,8). 

20 

Table 7.- Stomach contenl'> of lIoemulon plumieri from Puerto Rico and the 

U.S. Virgin hland., . (from Randall 1967.) 

I'ood 

lrahs (Mllhrul 'p.) 

Pol yc haete s 

.... "( hlnoid ~ (DlluJemll wlfll/ufum, Fucidari\ Inhu/mdn , 

Volume (% ) 

260 

14 .5 

'palang(lId) 12 4 

SlpunculI,h (.4 ,['u/ol/phofl 'P) 8 3 

Ga'lrnpolh (4, l1Ia('(/ anlll/arl/l11 Siromh'/l gigo<) (, 9 

Shnmps (alphe ld ,) ~ H 

Ophlurolds (Opillolhrlt) 5 7 

Unu.1entlfIeolru la lcans 5 3 

FIShes 3 3 

HcmllhnrJ.llC' 3 J 

l nlocnililed anImal nMlter 3 (J 

Hololhunan I fill 0"" p'''I/Jojll_" ..... " ..... , ______________ 2_7_ 

Table S.-Food of Jlaemulon p/umieri in Cuba. If rom \ald(:, \1uo1o, and iha 

let 1977. tables 8.9.) 

nl~ t: h iH.'{C' 

Cru,taf,,:\.;,tn'S {("anua (I tr.J('tltid. dmph'roJa. tantJl(t.1CCa 

pnrtunIo,. pa ~unJ lnpcp(xh .• ,opoJ mYSlJ. 

CU014JLCanlio nchaliaLcans) 

TUnlcah:, 

f:ch.n"Janh (hlllcHhunan OphlUrcllU) 

\lclllu,b (scaphopc)(h. chllons I urcilld ilmpets) 

F"h h1phlchth) IUS) 

L nlocntlilcu 

Cru:-.ta~e;Jns 

PoI) chactt:' 

:\cmcrtt..'Jn, 

'1"l1u,~, 

FI.. h.n .. .lJerm 

l nlo~ntlilco 

From natural reef. Cayo Ole\?0 Pc!rel 

Volume ($1 ) 

2S 7 

5 

.8 

35 

19 

I I 

03 

l6 

I) , 

O'i 

() , 

425 

Carr and Adams (1973) found that juvenile (21-40 mm SL) from 

Cry stal River were mainly plankttvorou . Copepods were the main 

food item in peclmens 21-35 mm SL. followed b, a tran~itlOn to 

mysids and postlarval shnmp (60% of tomach contents of 36-40 

mm fish) . Sand grains and detritu were also found in the stomachs 

examined. ReId (1954) reported similar results from juveniles collected at Cedar Key. 

3.43 Growth rate 

Haemu/on aurolineatum 

Several workers have published information on growth of H. 

aurolinearulll. most indicating that the, grow fairly rapidly before 

maturity and quite slowly after maturity. Sokolova (1965) backcalculated sizes at age from ~cales and reported rapid growth in the 

first 2 yr of life of Campeche Bank fish. Sauskan and Olaechea 

(1974) also found that H. auro/ineatum on Campeche Bank grew 

more rapidly prior to maturity than after. Olaechea and Hernandez 

(1976) calculated a von Bertalanffy relationship for Campeche 

Bank H. aurolineatum, with considerably different parameters. 

Manooch and Barans (footnote 5) aged specimens from the South 

Atlantic Bight, using otoliths and scales, and found that the first 

annulus of the otolith usually formed in late winter or early spring. 

Growth increments of H. aurolineatum aged by scales were: 
Age increment (yr) Length increase TL = 1. 7489 SRO.9572 (r= 0.93, n= 103). 

I-II 

II-Ill 

Ill-IV 

IV-V 

(mm, TL) 

46.5 

21.1 

17.0 

14.5 

Haemulon aurolineatum grows faster than most other previously 

studied reef fishes from the South Atlantic Bight (Manooch and 

Barans 1979). Size-at-age data based on the work of several investigators appear in Table 9, and von Bertalanffy growth parameters 

are listed in Table 10. 

where total lengths (TL) and scale radii (SR) are related by the function: 

Length-weight relationships of H. aurolineatum were derived by 

several workers (Table 11). Other growth data were provided by 

Table 9.-Size-at-age data for Haemulon aurolinealum. 

Mea!\urement Age (y r) 

and 2 3 4 5 6 7 8 9 ~our('e 

\lean lenglh (em) 

Sokoll)\ a 1965 ' 8.94 1240 15 .2 1 17 .22 18.94 20.20 , 

Sauskan and 

Olaechea 1974J 10.7 14 7 172 18.0 18.5 19.3 20.0 

Manooch and 

Bamn;, (IC\I 

foolnolc 5)" 13.5-1 18 19 20.30 22.00 23.45 2557 26.58 2770 28.67 

Manooeh and 

Bamn, (le\1 

footnote 5)'\ 1030 10.93 1993 21.60 2304 2-162 25.80 26.62 28.05 

1>lanooeh and 

Bam", (ICX! 

footnote 5)0 12 2-1 1594 1892 21.31 23.22 2476 25.99 26.98 27.78 

\Ican "eighl (g) 

Sau,kan and 

Olacchca 1974' 46 ( lJ2 3 1257 130.0 1-1-16 156.0 166.8 

Sau,kan and 

Olaechca 1974' 33 6 ~O.8 116 .8 1383 150.5 1573 160.1 

ICampcche Bank. 1964. len~lh, back-calculaled from scalc,. Iype of lenglh measuremenl unspecified. 

~ Not gJ\cn 

'Campechc Bank . observed lenglhs. I) pe of lenglh measuremcnl unspecified 

"South AllanliC Blghl . observed IOlal lengths. 

'Soulh AllanliC Bighl . back-calculated total lengths 

.Soulh Allanlic Blghl: [ncorellcal lola I lengths based on von Benalanffy equa li on: 

I.; 310(l-exp[- 0220 17(1+ 128)1) 

7Campeche Bank. obsened ",eights 

'Campechc Bank: Iheoretical weighls based on \'on Benalanffy equal ion 

W; 1.645 .5( I-exp[ - 0664(1+ 0.34))). 

Table IO.-Von Bertalanff} growth equation parameters for Haemulon aurolinealum and H. plumieri. Standard equation for length at age is: I, (I

e kI, '0'), and for weight at age: w,= (I- e kl' '0' lb. 

Billings and Munro (footnote 3) from Jamaica: SL = 0.88 FL+0.3, 

where SL is standard length and FL is fork length, both in centimeters; and d = 0.3001- 0 .20, where I is fork length and d is body 

depth, both in centimeters. SpeCll!~ 

and 

~ource Localll) 

H GUI"(I/iIlCOIUIH 

Sau,kan and Campeche 

Olaechea Bank 

1974 

Olaechca and Campcchc 

He rmindez Bank 

1976 

Manooch and Soulh AllanBaran, 1979 

li e Bighl 

H. p/umieri 

Billing, and Jama ica 

Munro (text 

iootnOle 3) 

Manooc h 1976 Carol inas 

'Nol give n. 

Parameter 

~ W~ to 

(cm) (g) (y r) 

_I 164 5 - 034 

27. 13 - 1.75 11 

3 1. 0 - 1.28 

42.0 

64 .0 - 1.007 

K 

0.664 

0. 1839 

0 .22017 

0 .34 

0.1084 

b 

3 

21 

Ha emulon plumieri 

Several studies have been conducted to evaluate growth of H. 

plumieri. Moe (1966, 1967) provided growth data based on tagrecapture studies on the west coast of Florida (Table 12). Length-atage data were collected by Kapote (1971 ) and Manooch (1976) for 

H. plumieri from Campeche Bank and the Carolina coast, respectively (Table 13). Growth rate appears to be higher along the coast 

of the southeastern United States than on Campeche Bank . Billings 

and Munro (footnote 3) studying Jamaican populations, found that 

offshore populations grew faster than inshore populations. Growth 

rates in Jamaica were higher than those reported by Moe (1966, 

1967) and Manooch (1976); von Bertalanffy growth equation 

parameters are presented in Table 10. Growth curves prepared by 

Billings and Munro (footnote 3) and Manooch (1976) appear in 

Figures 14 and 15 . 
Table ll .-Length-we ight parameters for Hoem uloll ourolineatum and II. plumieri. The length-weight relation,hip 

is defined as: L = aWl>, where L is length in centimeters and Wis body weight in gram~. 

Species Loca lity Rc,tnetlon, 

H. auro/inealUm South Atlantic Bight 

Jamaica Specimens 7 2 11 .6 em FL 

Jamaica pecimens l21-ISOemFL 

Venezuela 

H plLl/nieri N. and S. Carolina Male, and females 

Nand S. Carolina Male, only 

Nand S. Carolina Female, only 

Jamaica 

IUsed rotallenglh. wet body weight r= 0.996." = 70 

2Used fork length, wet body weight 

'Used standard length. dry body weight 

"Used total length, wet body weight; r= 097 

Table l2.-Growth of tagged-recaptured Haemulon 

plumieri in the eastern Gulf of I\l e\ico. (From 1\loe 

1966, table 7; 1967.) 

TL at tagging TL at recapture Gro"th rate 

(mm) (mm) (mm.mo) 

225 234 36 

262 338 20 

264 280 19 

280 286 24 

287 307 1.4 

Length-weight relationship parameters of H. plumieri are presented in Table 11. Females weigh somewhat less than males of a 

comparable length (Manooeh 1976). Billings and Munro (footnote 

3) calculated other body relationships of H. plumieri: 

SL=0.92 FL-0.5, where SL is standard length and FL is fork 

length; 

TL= 1.15 FL, where TL is total length; 

body depth (em) = 0.366 FL (em) - 0.50; and 

scale radius (mm) = 0.366 FL (em) + 1.35. 

L.,_ 40 

1 30 X 

... 

" ~ ~ 

'" 2 .. 

2 

10 

Parameter 

a b Source 

00099 30905 M"nooch and Baran, 

(text footnote 5)1 

00094 3 25 Billing' "nd Munro 

(texl footnote 3)2 

00298 279 Billings dnd Munro 

(text footnote 3)2 

0.0021 34396 Edwards 1971' 

o 01 ~O 30229 Manooch 1976-' 

0.0135 30503 Manooch 1976-' 

0.0153 30214 Mdnooch 1976" 

0.0238 2.93 BillIngs and Munro 

(text footnote 3)2 

0+2 t+4 t+6 

AGE (yro) 

13S3 W 

1000 

soo 

~ ... 200 % 

~ 

100 i 

so 

,.,8 

Manooeh (1976) related scale radius (SR) to total length as follows: 

TL=0.5180 (SR) I 18'4 (r= 0.93). 

Figure 1~ .-Gro" th c un e for Hoem ulon plumien' in Jamaica estimated using 

the gro" th eq uatio n : I H+ I)=Lw(l-k) + kl, derh ed from \ on BertaJanIT). "here 

Lro= 42 cm. k= 0.71. a nd (= I )r. The instant a neoll5 gro\\th r a te (K) is 0.34. 

(From Billings and Munro see text footnote 3, fig . 3.7.) 

Table 13.-Size-at-age data for HaelTluloll plumieri. All lengths are in centimeters. 

Source 2 3 4 

Kapote 1971 1 , 23.8 26.5 

Kapote 1971' 23.6 25.8 

Kapote 1971 4 23.6 26.0 

Manooch 19765 17.7 22.7 26.7 29.6 

Manooeh 19766 20.9 26.3 30.5 

Manooeh 19767 9.7 18.5 24.4 28.3 

ICampeche Bank; males only; total lengths. 

'Not given. 

3Campeche Bank; females on ly; total lengths. 

Age (yr) 

5 6 7 

29.1 30.7 

28.3 

28.0 30.7 

32.9 35.6 39.0 

32.8 35.8 39. 1 

3 1.4 34. 1 36.7 

4Campeche Bank; males and fema les combined ; total lengths. 

8 

41.2 

40.0 

38.9 

5North and South Carolina; empirical to tal lengths based on sca le readings. 

6North and South Ca rolina; empirical total length$ based on otolith readings. 

7North and South Carolina; calculated total lengths based on scale readings. 

22 

9 10 II 

43.0 45.0 46.8 

42.8 44.9 45 .6 

41.4 43.9 46.6 

12 

48.2 

48.1 

49 .2 
-

E 

E 

-

x 

~ 

C!) 

Z 

UoI 

..... 

X 

'" """ ..... 

~ 

~ 

0 

~ 

600 

SOO 0·····0 Observed 

.-. Calculated 

/1---6. Theoretical 

AGE 

Figure IS.-Comparison of absolute. back-calculated. and theoretical gro\l th runes for Haemuloll plumieri from the southeastern Atlantic coast of the United States, 

obtained from scale readings. (From Manooch 1976, fig. 6.) 

3.5 Behavior 

3.5 I Migrations and local movements 

Haell/ulon aurolineGlllni 

Movements of H. allrolineatum are of three types: seasonal 

inshore-offshore migrations, size-dependent habitat changes, and 

nocturnal foraging migrations. Seasonal inshore-offshore migrations have been noted on Campeche Bank by several authors. Individual apparently gather in fall on ea tern Campeche Bank to 

spawn (Sokolova 1965). Lyubimova and Kapote (1971) similarly 

reported H. aurolineatum most dense on the eastern and central 

Bank in fall and winter. Offshore migrations in spring-summer and 

inshore migrations in fall-winter were mentioned by Olaechea and 

Sauskan (1974) and Sauskan and Olaechea (1974) (Fig. 16). Billings and Munro (footnote 3) noted migrations of young grunts from 

nursery areas to adult grounds in Jamaica. Stone et al. (1979) 

observed that size plays a role in movement of subadult H. aurolilleGlum in reef areas of southern Florida. Different size-groups 

inhabit different patch reefs, and apparently move away when they 

reach a certain size. Manooch and Barans (footnote 5) noted a 

minor offshore migration of adults and juveniles in the South Atlantic Bight in winter. Haemulon aurolineatum appears to avoid water 

shallower than about 20 m during winter north of Florida (Manooch 

and Barans footnote 5). 

Nocturnal foraging migrations of grunts, including H. 

aurolineatum, are well known. Such migrations have been noted in 

the Virgin Islands (Randall 1963; Collette and Talbot 1972; Smith 

23 

and Tyler 1972; Ogden and Ehrlich 1977), Puerto Rico (Parrish and 

Zimmerman 1977), Jamaica (Billings and Munro footnote 3), the 

Florida Keys (Davis 1967), and on offshore platforms in the northern Gul f of Mexico (Hastings et al. 1976). These migrations usually begin at dusk, with the dense diurnal schools becoming less 

organized and finally dispersing over grass and sand flats. Fish 

return to their diurnal resting schools at dawn . Such migrations 

may reduce predation on the adults, increase avaiJable foraging 

areas, and play an important role in energy input to reef communi210 

YUCATAN 

PENINSULA Ij 

200 N 

920 910 900 890 880 870 W 

Figure 16.-Main zones of concentration of Haemulon aurolillealum on Campeche Bank and probable seasonal movements. (From Olaechea and Sauskan 

1974, fig. 2.) 
ties. No information exists on foraging migrations of grunts in 

deeper water. 

Haemulon plul11ieri 

Although some short-range migrations of H. plumieri may occur, 

most evidence indicates that movements are slight and confincd 

primanly to foraging . Tagging studies by Moe (1966) on the west 

coast of Florida showed that only 2 of 39 recaptures moved more 

than 8.0 km (5 mil from the tagging site, and only 26% moved 

more than 4.8 km (3 mil. Moe (1966) felt that these were probably 

casual wanderings ratherthan true migrations, and that individuals 

spent most of their lives in the same general area. The greatest 

movement probably occurred during or before spawning, when 

large schools formed. Beaumariage (1969) reported that three 

tagged H. plumieri were recovered 4 yr later at the tagging site. 

Springer and McErlean (1962) found little migration of tagged H. 

plumieri on patch reefs off Upper Matecumbe Key, Fla.; even during a hurricane most of them maintained home reefs. McLean and 

Herrnkind (1971) reported two dense schools of grunts, primanly 

H. plumieri, migrating along the shore of North Bimll11, Bahamas. 

over open sand. These schools of 4,500-10,000 IndIviduals ma) 

have been moving to avoid turbid water created by a storm. 

Juveniles may change habitats or reef resting sites dunng theIr 

growth, not selecting adult territories for some time (Ogden 1977; 

McFarland 1980). Billings and Munro (footnote 3) reported that 

juveniles may migrate away from reefs at Port Royal, JamUlcll, at 

about 25 cm FL, but their migration routes are unJ...nov, n. 

Most documented movements of H. plul1Iien are dlurnalnocturnal foraging migrations. Ogden (1977) and Ogden and 

Ehrlich (1977) studied these migrations extensively m St. CroIX. 

using tagging and diver observation. Heterotypic grunt schools, 

including H. plumieri, moved off reefs to feed beginning about 10 

min after sunset and returned to the reef ~0-30 min before sunrise. 

Largest fish left the reef first and returned last. Schoob remaIned 

together until they were off the reef, possibly to reduce predatIon. 

Migration routes distributed fish evenly over nocturnal foraging 

grounds, such as grass beds. These routes remained constant over 

the short term, and more or less so over the long term. 

3.52 Schooling 

Haemulon aurolineatum and H. plumieri, like most grunts, are 

schooling fishes. Both adults and juveniles form schools on reefs or 

over other hard substrate during the day. These schools are usually 

composed of one size-class but may be heterotypic with other 

similar-sized species. Daytime resting schools may be quite dense 

and occur in a variety of habitats. Billings and Munro (footnote 3) 

reported adult schools among corals on Jamaican reefs, with juvenile schools over Thalassia beds, near edges of sand patches, or on 

reefs near adults. The density of schools may depend on light intensity (Billings and Munro footnote 3) and schooling may defend 

against diurnal predation (Davis 1967). Conspecific attraction has 

been noted in some grunts (e.g., H. plumieri) and may account for 

large trap catches (Billings and Munro footnote 3). 

Haemulon aurolineatum has been observed in mixed schools 

with H. striatum and Emmelichthyops atlanticus on deep reef 

ledges of Alligator Reef with 200 to 300 mm SL individuals of Lutjanus griseus and H. parmi off Windley Key, Fla ., (Davis 1967) 

and with H. striatum in Biscayne National Monument, Fla. (Stone 

et aJ. 1979). Juveniles of H. aurolineatum school with juvenil e H. 

24 

jlavolineUlum in St. John (Sm ith and lYle r 1972). Ogden ( 1977) and 

Ogden and Ehrl ich ( 1977) reported heterotypic resttng schools o f 

grunts, includ ing H. aum/inca/um and H. plumicri, on patch reefs in 

St. Croix. Starck and Davis (1966) found H. aurolillealum in small 

resti ng schoob among the coral Acropora cervicornis III the Florida 

Keys. 

Hacll1l1/on p/ulllieri has been observed school Illg with H. 

SCiuru.l, H. jlavolinealUm, ACanlhurtl.1 chirur/?u.I, Mulloidichlhy.1 

lIIarlillicus, and other species (Col lette and Talbot 1972; Motehek 

and Sliva Lee 1975; Ogden and EhrlIch 1977) Schools arc common in reef areas dunng the day, partIcularly near patch reefs 

(Bohlke and Chaplin 1968; Ogden 1977; Ogden and EhrlIch 1977). 

Collette and Talbot (1972) reported H. plilmieri common in small 

resting schools of 5-20 IndIVIduals on Virgin I51and reefs. Ogden 

(J 977) found that small juvenile H. plilmien ~chool with Juvenile 

H. flam/incatum on patch reef~ at St CroiX, while larger juveniles 

(15-25 cm. measurement unspecIfIed) roam the reefs and adjacent 

Oats. HeterotypIc schools of young grunts arc often assocIated with 

the corals Po flies porites and Acropora pa/mala on St. Croix patch 

reefs (Ogden and EhrlIch 1977). v,hereJs larger 5ubadult (12-15 

cm, measuremcnt unspecified) In schools of up to .... everal hundred 

indl\ Iduals are no longer assocl.Jtcu v,\th any particular coral formatIons. Schools remam Intact dunng the dJ: and break dov.n at 

nIght v, hen the fish Icave the rcef to forage. 

Conspecific attraction tends to drav, H p/umleri into fish traps. 

though ,>ome arc "nov,n to eSLape (Billings and Munro footnote 3). 

Billings and Munro (footnote 3) founu that If more than four H. pluI11leri v,ere in a JamaIcan fish trap. catches thereafter increased conSIde rably . 

4 POPCLATIOl\ 

4. I Structure 

4. I I Sex ratio 

Sex rallO of H. ([lIrullllelllulII IS llose to I: I (So]..olo\·a 1965). 

Billing. anu Munro (footnote 3) reported 530/, females anu 47"'r 

males from fi h-trap catches at Port Royal . 

Sex ratios of H. pllll11ieri from trap catche in JamaIca generally 

favor females (Billings and Munro footnote 3) (Table 14). Trap 

catches from Port Royal reefs vary around the 50% female level. 

with no apparent seasonal pattern (Fig. 17). 

See 3.11. 

Table 14.-5e, composition (in percent) of Haemulon plumieri 

from Jamaican trap catches. (From Billings and Munro see te,t 

footnote 3, table 4.1.) 

Local It) 

Sex Pon Royal reefs Pedro Cays Pedro Bank 

Females 57 59 69 

Males 43 41 31 

4.12 Age composition 

Sokol ova (1965), Sauskan and OIaechea ( 1974), and Olaechea 

and Hernandez (1976) studied age composition of H. auro/in eatum 

on Campeche Bank (Tables 15 , 16). Only fish 3 y r old and older 

ente r the commercial fishery on Campeche Bank , with 4-yr-olds 

dominating catches. 
III 

.. 

... 

c 

:e 

.. 

... 

~ 

100 .-----------__ --------------------------------, 

so ------------~-~-=-~ -- --- ------

~--r_--r__,--~--_r_r_.--_.--,_--,_--._~ 

J -F M -A M -J J -A 5-0 N-D J-F M -A M-J J-A 5-0 N-D 

1970 19 7 1 

MONTHS AND YEAR 

Figure 17.-Variations of sex ratio in bimonthl) trap catches of Haemulon plumieri from Port Royal, Jamaica, reefs over a 2-) r period. (From Billings and 

Munro see te,t footnote 3. fig. 4.1.) 

Table IS.-Age composition (in percent) of 

Haemulon aurolillealum on Campeche Bank in 

1964. (From Sokol ova 1965, table 6.) 

Age (yr) 

TIme of catch 3+ 4 + 5+ 

January. February 9.0 680 230 

March-May 19.6 652 152 

June-Augu,t 18 I 638 18. 1 

September-December 23.-1 620 14 .6 

Total 18 .7 64.4 16.9 

Table 16.-Age composition (in percent) of Haemuloll aurolillealum on Campeche Bank, and total fishing effort in thousands of hours of trawling of an 

SRT-M-type vessel. (From Olaechea and Hernandez 1976, table 1.) 

Age group' Total 

Year 2 3 4 5 6 effon 

1969 020 770 2850 30.50 26 10 ,30 0.70 18.689 

1970 0. 10 6.30 2600 34.40 2760 8.50 I 10 16. 125 

197 1 440 24.20 29.90 2970 10 .20 I 60 20431 

1972 o 10 9.20 28.90 30.30 2500 5.80 0.70 12680 

1973 140 9.47 26.47 38.34 25.44 7.9 1 1.64 14 . 187 

1974 078 12.47 30. 15 28.53 22.59 4 .77 0.81 I 7.109 

I E,lImated effon 

Manooch (1976) reported that older H. plumieri from off the 

Carolinas were always males. Males up to 12 yr old were collected, 

but females reached only 9 yr. 

See 3.43 and 4.4. 

4.13 Size compo ition 

Experimental trawling for H. aurolineallllll on Campeche Bank 

(Sokol ova 1965) produced a frequency mode of 17-19 cm (measu rement unspecified) . This changed only slightly with season and 

was imilar to the size composition of commercial catches. except 

that the mode of commercial catches was 19-20 cm (measurement 

unspecified) (Fig. 18). Length and weight distnbutions for three 

year-classes on Campeche Bank are shown in Figure 19. Weight of 

H. aurolinearum in commercial catches varied from 60 to 190 g, 

with 30% of the fish weighing 110-130 g (Fig. 20). Wenner et al. 

(l979a. b) developed length-frequency plots based on MARMAP 

trawl surveys off the South Atlantic Bight in 1973 and 1974 (Fig. 

2 1). 

25 

!! 

.. 

v 

Z 

.. 

'" '" ;:) 

v 

v 

0 

3 0 

2 8 

26 

2 4 

22 

20 

18 

16 

14 

12 

10 

8 

6 

4 

2 

0 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 

LENGTH (em) 

MAY 

AUGUST 

NOVEM8ER 

Figure 18.-Size composition of Haemuloll aurolillealum from Campeche Bank 

in 1964, caught b. experimental tra,\ling . (From Sokolo.a 1965. fig . 7.) 

40 

30 

~ ~ 

V 

Z 20 ... 

:I 

0 

... 

III: 

... 

10 

0 

12 13 14 15 16 17 18 19 20 21 22 23 24 

LENGTH (em) 

30 

l 5. 

~ 20 V 

Z 

... 

:I 

0 

... 

at: 

... 10 

LL~~~~--~--~--~~--~~~~~ 

50 60 70 80 90 100 110 120 130140150 160 170 180 190 200 

WEIGHT(g) 

Figure 19.-Length and "eight distributions of three )ear-classes of Haemulon 

aurolillealum on Campeche Bank in 1964. Ages are expre,sed in )ears. (From 

Sokolo,a 1965 data, tables 8, 9.) 
~ III 

u 

Z 

III 

III: 

III: 

::l 

U 

u 

0 

18 

16 

14 

12 

10 

' 8 

6 

4 

2 

o 60 70 80 90 100 110 120 130 140 150 '160170 180 190 

WEIGHT(g) 

Figure 20.-Weight of Haemu/oll aurolilleafllm in commercial catches from 

Campeche Bank in 196-'. (From Sokolo\a 1965, fig. 8.) 

22 

20 ., 

18 I , , I 

I 

I 

16 I 

I 

I 

~ I 

14 I 

I III I 

u 12 I 

Z I 

I III I 

III: 10 I 

III: I 

::l I 

I 

U 8 I 

U I 

I 0 6 I 

I 

I 

I 

4 I 

; 

I 

2 I 

I , 

, 0 , 

S 10 lS 20 2S 

FORK LENGTH (em) 

Figure 21.-Length-frequenq distribution of Haemu/oll aurolitleafum caught 

in the South Atlantic Bight during fall 1973 (solid line), and sprin g 1974 (broken line), b} MARl\IAP tra,,1 inHstigations. (From Wenner et al. 1979a, fig. 

SF; 1979b, fig. 16F.) 

Billings and Munro (footnote 3) provided length-frequency distributions of H. plumieri from Jamaica and compared these data 

from different areas and for different gear types (Figs. ::!::!-::!4). 

See 3.43 and 4.4. 

4.14 Subpopulations 

See 1.23. 

4.2 Abundance and density 

Haemulol1 aurolineatum 

Haemulon aurolineatum has frequently been reported as an 

abundant or dominant species where it occurs. Beebe and Tee-Van 

(1928) reported H. aurolineatllm very common in Port-au-Prince 

Bay; Hildebrand (1954) found them extremely abundant over shell 

bottom on pink shrimp grounds off Campeche, Mexico; Cummins 

26 

40 

20 

20 

VI 10 Z 

'" ;a; 

8 

... 

VI 20 

0 

Z 

10 

20 

10 

15 20 25 30 

FOR K LENGTH (em) 

Mosquito Coy. 

S.rranilio aonk 

July 1971 

".12' 

( , P.dro Bank 

F.b. 1970 

Novldod and 

SII",., Sanks 

Sept. 1969 

n: JI 

35 40 

Fi!(ure 22.-Len!(th-frequenC) di,tribution, of Haemulon plumieri from trap 

catche, (probaille me,h ,ife " . 13 cm ma,;mul11 dimen,ion) from four areas 

near Jamaica. Peak" marked" itll arro"" are probable )ear c1asse~. (From 

Billing" and \lunro see te-t footnote 3, fig. 3.3.) 

40 

.. 30 

u 

~ ~ 

~ II< 

... 20 

:; 

" ~ 

~ u 

~ 10 ... 

lS 20 

"\\, 

· · · · . . , 

, 

· · · 

2S 30 

FORK LENGTH (em) 

. 

. , 

35 40 

Figure 23.-Length-frequenc} distributions of line catches (broken line) and 

trap catches (solid line) of Haemulon plumieri from Jamaica stations" here 

both gea rs "ere u,ed. (From Billings and ~Iunro see te,t footnote 3, fig. " .2.) 

et al. (1962) caught H. aurolineatllm in potential commercial quantities off the southeastern United States on the Silver Bay; and Parrish and Zimmerman (1977) reported that H. allrolineatum 

dominated grunt catches in Puerto Rico. 

Randall (1963) found that juvenile H. alirolinealUI11 were the 

dominant inhabitants of artificial patch reefs in St. John. Five hundred seventy-seven individuals, weighing a total of 7.38 kg, occupied a reef of about 125 m~, with a habitat area of about 50 m". 

Ogden (1977) and Ogden and Ehrlich (1977) found up to 1,400 

individuals on patch reefs in St. Croix. Stone et al. (1979) found 

that juvenile H. aurolinearulIl were by far the dominant inhabitants 

of patch reefs in Biscayne Nation!!l Monument, with over 800 individuals on a single patch reef. Hastings et al. (1976) and Smith et 

al. (1979) reported them common on artificial reefs and platforms 

in the northeastern Gulf of Mexico. 
H -.N __ _ 

0.5 

025 

> 

5 

.. 

• 

0 i 0.1 

0 I 

! 02 
the abundance of H. plumieri on the shelf; party boat catches on the 

Carolina continental shelf include large numbers of H. plumieri 

(Sekavec and Huntsman 1972): 

Offshore 

In hore 

Percenrage of toral 

catch 

No. 

12 % 

43% 

WI . 

6% 

42% 

Haell1ulon plumieri is second only to black sea bass in inshore 

catches on the Carolina shelf. 

On Campeche Bank, H. plumieri is caught in trawls at depths of 

23-36 m, but is not a major component of the catch (Sokolova 

1965). Sauskan and Olaechea (1974) and Olaechca and Sauskan 

(1974) reported that the mean total annual abundance of H. plumieri on Campeche Bank is about 10,000 t (compared with 200,000 

t for H. ourolineatum), with a mean density of 0.00 18 t/ha (4.04 t/ 

10 nmi~) in the sampled area. Lyubimova and Kapote (1971) found 

greatest densities of H. pllllnieri on the central and eastern parts of 

Campeche Bank in fall and winter. 

Relative abundance of grunt species may vary considerably from 

reef to reef in certain areas (Billings and Munro footnote 3). On 

Pedro Bank reefs in Jamaica, H. plulIlieri and H. jlOl'ollllealulIl are 

numerically dominant (Billings and Munro footnote 3). 

4.3 Natality and recruitment 

4.31 Reproduction rates 

Mean fecundity of 13 lipe female H. ourolineatil/ll from Jamaica 

(mean length 15.4 cm FL. mean weight 69 g) \.\as 30.000 eggs/ 

female (435 eggs/g). The greatest observed fecundity was 83,000 

eggs in a 13 .1 cm FL, 135 g female (615 eggs/g) whose ovary 

weight was 7.5 g (Billings and Munro footnote 3). umber of 

spawns per year is not known. 

Billings and Munro (footnote 3) calculated a fecundity-weight 

relationship for H. pill/Ilieri from Jamaica of e = 607 II' - 93,000, 

where e is fecundity and II' is body weight in grams (Fig. 26). Mean 

calculated fecundity for H. plumieri from Port Royal reefs was 

about 42,000 eggs/female per spawning season (average fish 

weight 220 g). For unexploited parts of Pedro Bank, Jamaica, 

150,000 eggs were found per female per spawning season (average 

fish weight 400 g). 

4.32 Factors affecting reproduction 

Little is known about factors affecting reproduction in H. 

aurolineatum and H. plumieri, though Munro et al. (1973) suggested that spawning of grunts in Jamaica is initiated when water 

temperature drops below 28 °C. Peak spawning in Jamaica occurs 

during periods of low water temperature. 

4.33 Recruitment 

Haemulon aurolineatum 

Sokolova (1965) reported that lack of fluctuations in catches or 

age distributions of H. aurolineatum from Campeche Bank (1962-

65) indicated a steady recruitment to the commercial populations 

there. Olaechea and Hernandez (1976) calculated a yield per recruit 

curve (Fig. 27) and an age of first recruit on Campeche Bank of 

28 

200 

150 

I 

" 100 .. 

l1li 

l1li 

• 

Ii 

.!. 50 ~ 

... 

Q 

z 

:;, 

u 

.. 

. . 

. . 

: O~rT"TT'-~'-rT'-rTonrr~-r~-rTJ-r~-r~-rrT~ 

100 300 400 500 600 

WEIGHT (9) 

-50 

~OO-L __________________________________________ j 

Figure 26.-Relation~hip of fecundit) to ,Hight in Haemulon plumieri from 

Jamaica. Slope of the line i, 607 egg' g and the y-intercept - 93,000 eggs. 

(From Billings and \1unro see text footnote 3, fig. 3.1.) 

ISO 

:§ 100 

... 

:; 

u '" ... 

'" 

'" so ... 

~ 

Q 

... 

... 

~ 

0.1 0 .2 0.3 0.4 O.S O.b 0 .7 0.8 0 .9 1.0 I.S 2.0 2 .S 3.0 3.S 4 .0 

FISHING MORTALITY (F) 

Figure 27.-Yield per recruit curve of Haemulon aurolilleatum on Campeche 

Bank for different values of fishing mortalit) (F). (From Olaechea and Hernandez 1976, fig. 1.) 

3.32 yr. Recruitment of H. aurolinearum to fish traps on Pedro 

Bank was studied by Billings and Munro (footnote 3), who calculated that the minimum recruitment size to the traps is 13.5 cm FL. 

the mean recruitment size is 15.9 cm FL, and the size at full rec ruitment is 16.5 cm FL. The minimum length of H. aurolinearul11 

retained by 4.13 cm mesh traps is 14.4 cm FL. 

Haemulon plumieri 

Recruitment of H. plumieri to patch reefs in St. Croix may vary 

throughout the year (Ogden and Ehrlich 1977). At Port Royal H. 

plwnieri is fully recruited to the fishery at 15 .5 em FL, with a mean 

recruitment length of 14.9 cm (Billings and Munro footnote 3). At 

unexploited parts of Pedro Bank, mean recruitment length is 26.2 

cm FL. At Port Royal reefs, main recruitment takes place from 

May to October. 

See 4.5 . 
4.4 Mortality and morbidity 

Haemulol1 aurolineatum 

Natural mortality of H. auroli'}eatum is caused by predation, disease, and senility (Billings and Munro footnote 3). A mortality rate 

calculated from Pedro Bank trap catches is given in Tabl e 17. 

Sauskan and Olaechea (1974) calculated a mean total annual morTable l7.- Mortality rate of Haemulon aurolineatum from 

the ' relative abundances of successive length groups captured in traps (4.13 cm mesh) on Pedro Bank, Jamaica. 

Because Pedro Bank is essentially unexploited, the total 

mortality rate (Z) is equal to the natural rate (M). The 

instantaneous annual growth rate (K) is not known. 

Method is that of Beverton and Holt (1956). (From Billings 

and Munro see text footnote 3, table 4.7.) 

Length grou p 

(FL,cm) 

12 .0- 12.9 

13.0- 13 .9 

14.0- 14.9 

15.0-15.9 

16.0-16.9 

17 .0- 17 .9 

18.0-18.9 

19 .0- 19.9 

20.0-21. 9 

~= 23 cm 

K= ? 

% Frequency 

1.7 

5.5 

28.9 

43.0 

17.9 

2.6 

0.4 

!s (length at fu ll recru itment) = 16.0 cm 

Relative frequency 

0.04 

0. 13 

0.67 

1.00 

0.42 

0.06 

0.01 

I (mean length above full recruitment) = 16.9 cm 

Z= K(L~-/) =? 

(7 -Ie) 

ZIK= L~-I =6.78 (= MIK) 

/-1 

tality rate for 5- 7 yr old H. aurolineatum on Campeche Bank of 

Z = 1.65 . Olaechea and Hermfndez (1976) calculated a total annual 

mortality rate of Z=0.6766 for H. aurolineatum on Campeche 

Bank and derived the relationship: 

2= M+ qf= 0.1332+ 0 .0356 (j) 

where 2 is total mortality, M is the instantaneous rate of natural 

mortality, q is the catchability coefficient, and fis fishing effort in 

thousands of hours of trawling of an SRT-M-type vessel. This 

equation appears as 2= 0 . 1332 - 0.0356 if) in Olaechea and 

Sauskan's paper, with the mean value of 2 given as - 0.6766. This 

may have been an error; the equation and 2 value presented above 

seem more reasonable. Although not enough information is given 

to evaluate their mortality estimates, Olaechea and Sauskan's figure 

fo r natural mortality seems remarkably low and suggests heavy 

fishing pressure. 

Manooch and Barans (1979), worlGng with fish aged by scales and 

otoliths, stated that H. aurolineatum does not live as long and has a 

higher natural mortality rate than most other reef fishes previously 

studied in the South Atlantic Bight. The mean total annual mortality 

rate from catch curves for 1972-78 was 59 % (2= 0.887) based on fish 

age IV and older (Manooch and Barans footnote 5). 

29 

Haemulon plumieri 

Mean total annual mortality rates of H. plumieri in Jamaica were calculated by Billings and Munro (footnote 3) (Table 18). Manooch 

(1976) reported that H. plumieri was fully recruited to the hook-andline fishery off the Carolinas by age VII, with a mean total annual mortality of37-51 % (Table 19, Fig. 28). 

4.5 Dynamics of population 

Haemulon aurolineatum 

Manooch and Barans (1979) derived a yield per recruit model for H. 

aurolineatum entering the hook-and-line and trawl fisheries off the 

southeastern United States that indicated the maximum yield per recruit 

at natural mortality (M) of 0.3 is 50 g with fishing mortality (F) as low 

as 0.8 and recruitment age of2.75 yr. Eighty percent (40 g) of the maximum potential yield per recruit is available when F= 0.25 and recruitment age is 1.5-2.5 yr. Sixty-seven percent of the yield per recruit is 

available at F= 0.2, M= 0.5, and recruitment age 0-1 yr (Manooch 

and Barans footnote 5). 

Olaechea and Hernandez (1976) calculated that maximum yield per 

recruit of H. aurolineatum on Campeche Bank is approximately 110 g 

Table IS.- Derivation of a nnual mortality rates from the relative abundances 

of successive length groups of Haemuloll plumieri beyond full recruitment to 

traps (4.13 cm mesh) at Port Royal, Jamaica, reefs, and at unexploited parts of 

Pedro Bank (for =42 cm, K=0.345). Because Pedro Bank is essenlia ll) 

unexploited , fishing mortality (F) is 0, and total annual morIa lit} rate (Z) is 

equal to the natural mortality rate (M). (From Billings and Munro see texl foolnote 3, table 4.3.) 

Length Unexploited Pedro Bank Port Royal Rcef\ 

group Relative 

(FL.cm) % Frequency frequency % Frequency 

12 .0- 12 .9 

13 .0-13.9 

14 .0- 14 .9 

15 .0- 15.9 

16.0- 16.9 

17 .0- 17 .9 

18.0-18.9 

19.0-19.9 

20.0-20.9 

210-21.9 

22.0-22.9 

23.0-23.9 

24.0-24.9 

25.0-25.9 

26.0-26.9 

27.0-27.9 

28.0-28.9 

29.0-29.9 

30.0-30.9 

31.0-31.9 

32.0-32.9 

33.0-33.9 

34.0-34.9 

35.0-35.9 

36.0-36.9 

37.0-37.9 

0.27 

0.57 

0.33 

1.07 

1.1 3 

4.20 

9.00 

11.80 

16.43 

15 .30 

14.83 

12.60 

8.03 

2.40 

0.83 

0.37 

0.07 

0.27 

0.20 

Ie (length at full recruitment) 

Mean FL 

ZIK 

Z 

F 

£(= F/Z) 

0.02 

0.03 

0.02 

0.07 

0.07 

0.26 

0.55 

0.72 

1.00 

0.93 

0.90 

0.77 

0.49 

0.15 

0.05 

0.02 

+ 

0.02 

0.01 

0.08 

2.83 

7.82 

11 .78 

1665 

11.27 

II.D:! 

8.97 

808 

8.18 

6.42 

5.17 

3.80 

1.92 

085 

007 

0.07 

0 

0 

0 

0 

0 

0 

0 

0 

0 

27.0 em 

29.45 cm 

5.12(= MK) 

1.77( = M) 

o 

o 

RclatJlc 

Irequenc) 

0.01 

0.24 

0.66 

1.00 

0.99 

0.96 

0.94 

0.76 

0.69 

0.69 

0.54 

0.44 

0.32 

O.ln 

0.07 

0.01 

001 

0 

0 

0 

0 

0 

0 

0 

0 

0 

AdJu,tcd 

Ircqucnc) 

145 

77 

63 

1.2 

o , 

01 

+ 

+ 

(I 

o 

o 

o. 

o 

o 

II 

o 

o 

210cm 

220 em 

200 

690 

5 13 

074 
Table 19.-Total annual mortality estimates for Haemulon plumieri from North Carolina and South Ca rolina , 

1972 to 1975 . (From Manooch 1976, table 3.) 

Method 

No. of Robson Beverton 

Sample size age and and Mean 

Year Area(s) (VII-oldest) classes Chapman Jackson Heinke Holt mortalltyl 

1972 all 1, 126 7 0.52 0.46 0.46 0.55 0.51 

1973 all 954 7 0.49 0.42 0,42 0.53 048 

1974 all 398 7 0.48 0.40 0.40 0.53 0.47 

1975 all 399 7 0.44 0.36 0.36 0.46 0,42 

1972-75 all 2,877 7 0.49 0.43 043 052 048 

1974-75 Cape Lookout 2237 8 0.38 029 0.29 0,43 0.37 

1974-75 Cape Fear 368 7 0,48 0.40 0.40 0,47 045 

1974-75 Cape Romain 332 7 0.42 033 0.33 0.42 0.39 

IMean of Robson and Chapman, Jackson, and Beverton and Holl method values. Heinke estimate was not used In 

Obtaining mean mortalities Since method was almoslldcntical to Jackson 

2Sample size was age V-XII. 

8 

>-

v 7 

Z 

..... 6 :J 

() 5 ..... 

DI: 

w. 4 

.,(J) 3 

0 2 .... 1 

0 

II III IV V VI VII VIII IX X XI XII XIII 

AGE 

Figure 28,-Catch curves for 5,135 Hael1llllon plumieri sampled from the headboat fishery off North and South Carolina, 1972 to 1975, where b, the slope of 

the right descending limb of the curve, equals the instantaneous rate of mortality, (From Manooch 1976, fig, 7,) 

Figure 29.-Dependency of potential yield per recruit (in grams) 

and of potential annual yie ld for R = 0.48 X 10' individua ls on coefficient of fishing mortality (F), "here M= 1.65,1,= 2.88, 1m ", = 7, 

10= -0,34 yr, W~= 164.5 g. Average weight refers to the predicted 

mean weight of individuals caught at each F. (From Sauskan and 

Olaechea 1974, fi~. 10,) 

II> 

... 

:I 

u '" ... 

'" 

'" ... 

a.. 

Q 

.... 

... 

... 

110 

100 

90 

80 

70 

60 

50 

40 

30 

20 

10 

at F= 0.7. Sauskan and Olaechea (1974) constructed a yield-perrecruit curve that differs considerably from that of Olaechea and 

Hermindez, leveling off at higher values of fishing mortality (Fig. 29). 

Haemulon plumieri 

Billings and Munro (footnote 3) estimated recruitment indices for H. 

plumieri from Jamaica are shown in Table 20. 

Huntsman and Manooch (1978) determined that the minimum size 

limit that would maximize yield per recruit of H. plumieri is 31.4 cm 

TL, based on growth data of Manooch (1976) . 

Table 20.-Estimated recruitment indices for Ha emulon plumieri from 

Jamaica, (From Billings and Munro see text footnote 3.) 

No. per Monalit) Recrullment 

I, 1.000 lrap-da), rate Inde\ 

Localion (om) FL Of Z R' 

E. Pedro Ban!.. 26.2 262 1.77 463 

S. Pedro Ban!.. 26.2 481 1.77 85 1 

W. Pedro Ban!.. 26.2 83 1.77 147 

Port Royal reefs 14.9 265-278 6.9 1.828-1. 918 

------ 50 

40 ------- ~ ----- "' ,... Potential Ave . C 

Annual W •. 30 

Yieldh09

g_) _(gL 

37 118.7 :~ 

Yi ~ 

Recruit 

3.0 76.59 

3 .5 80.33 39 118.2 20 4 .0 83 .44 40 117. 8 

10 

2 3 4 5 10 15 

FISHING MORTALITY (F) 

30 
~.() '1 hc pOpUlall!ln In Ih\: ~(lI1lJ1lUnll) ,IIlJ Ihe e " ) lem 

I/OI'll/II/Oil (lum/il/t',1/1II1/ ,IIlJ 1i.1'/IIIIl/( fI, hl:\:au .: 01 thl:lr ahunalln: in vOlrious OJr':OlS, prohOlhl) ~.:n.: as lI11p0r!Olnt 10rd!!l' Ilsh~s 

or IJrgcr carnl\(lics. B.:cause 01 Iheir acccptJrlCC 01 a \\ Idc \,met) 

)1 hJhltJh. th.:) maj he 1ll':ll1bers 01 s.:\.:r,i1 Jillcr.:nt COllllllUnltlC • 

uch as r.:d. rocky .. hor.:. sand or grass flat. or Ii, e-hotto111 olhhorc 

'(lITlIl1Unilies 

A"oclJtcs of II. (/liro/il/('(1/1I11l lrom s<Jnd bottoll1 tAR lAP tJ 

linn .. in th.: Soulh Atlantic Bight ar.: listed In T<Jhlc :! I Indices of 

~il11il<Jrity. hm\c\.:r. IIldlcate that the .. e spec I': ... wllh the posslhle 

'xccption of Cel/lw{lrLl!1I ,l/rW(lI and Rhtll/1/Jo{l/i(('I tlllroruht'lLI. 

re not do,.:ly <Jss()ci<Jted with Ii llllro/il/"tl/lII/1. l\1aIl) of th.: .. pe-

·Ie .. lIsted In Table 21 Illay actuall) he more commonl) asso('lated 

Wllh hard hottolll cOllllllunIties (Miller"). 

able 21.-Sirnilaril) indice, for 'pecie, found ."ocialed "ilh f1a emulon auroillfalulII durin): 1ARMAP inH'II):alio", in Ihe Soulh Atlanlic Bighl, 1974-75. 

and bOll om ,Ialion", 1\1elhod i" in>er,c anal},i, specie, clu,tcr. Canberranetric correlalion (Boe,ch 1971). (From Wenner ct at. 1979 a. b. c. d,) 

Ca/umu.\ Icuco.\lon 

J ll'miplCroflofU \ n(lWlcu/a 

Pural,d"hn £lOIIO(U\ 

Pnol/u/us _\(,/111114 _' 

I'Srhf1crOlJl'_' lIIocUlafU,\ 

uunciu hnHi/tt'fl.\H 

CCllfrnpri,\ li\ \lr/ellil 

Orlhopri,\lIs ("hrnoplera 

J LI:'O(Jon rhomhou/c\ 

( hllt'fOd,plt.'ru \ faht" 

,..lolllorrI1ll.\ am t'rln.lII LI \ 

Aful/II.\ c.wrufII,\ 

RLI('lrn'('1IlrOIl ('u/wc/um 

Da\HlIi.\ {('nlrOut" 

ClllluUlYU( 1"/4.\ .\("ho("pji 

f<IIII 'I( ,\ l,ul('c'oillflJ.\ 

Rhr'mhc'plilt'S (wroruhc"II\ 

Olfll'llIJwm omo\ligmum 

,'t("llotCJ11W.\ an4leat/1S 

(J,./lIdIOllht'tllll 

PI"/O,,"w\ t'lJt(l/tllll.\ 

I'rttlfJOfU.\ "plrnCl\ 

"('("fltlt'lld hrd"I't ,'-"\ 

(\,/flP'('lIdJimhrzd/f1 

HI 111111\ oc'( ""'/1\ P"CIIIOIU,\ rtl.\t 11\ 

.'l'htICTOld,.\ \PC 1lI:1t fl 

'lit 1t1f1\1,lnJlI\ ,1rgollfU,\ 

Spring 

1974 

0.2 

o 

()2 

0.2 

0,2 

o ' 

0.2 

0.2 

0,2 

0,2 

0,2 

0.2 

() 2 

() 2 

Summer 

1974 

u 

() , 

II 

0 

0 

Fall 

1<)74 

Il OS 

() 05 

{I , 

II I 

{lOS 

rlnter~ 

carl) 'pring I 'I7~ 

01 

{I I 

{I~ 

(II 

0 

n 

Il 

II 

01 

01 

() \ 

Rllling~ .1l1J ... lunn) ll\"",\!l1(ltl' -') Jla!!r.lJl1llled thL' mle of grunt m the 

,halk)" \\ .ttl'f Cant.bl'an ec()~) stl'1ll (i-tg. 30). Thl' \ .Inc!) 1)1 habitat 

1 .... :CUpll'll b) the~l' .Ix'\.'il'~ anJ tlll' diu rnal-nl ..... ·\U rnal h.lhllat "hange 

pl.l) an IlllptlI"Wnt t\lk In bringll1g nutn~rll into l\.'Cf l , ) ll:m (BI\Iillg~ .1l1d ... tUrin) Inotn, te 3), E tl'n~t' ~ sand ,tnd gra~ fl.lt .If"(' rill 

Ii r pt\\\ Idll1g l(lod. \\ htle Ll)ral 1\.'Ct" or lIth 'r h~rtI ub tr.ll<.: prO\ Id 

dturnal ,hl'itcr. AbunJa,I1(,l' lIt Iu 'l1l11hd 1 pmb.lbl) pml ru naI t Lh 

allll)lInt ,It 'llffi111I1JIlll,! f~ :Jlllg pnllnJ (Bllltn~ ,md ~ lulU\) fIno! 

-toe 'urna Habltot 

era" and Sand JI." 

Phytoplanl.ton 

O.t'ltu~ 

~ HAIMUUOS 

Pr.dator, 

-9 Snapper., 

Group '" 

Barrocuda 

p, 4.'0" 

• g Jo, .... 

80rfocloldo 

.,~un· .111 . Iliagram III illu Ir It tb pOSlilon "r Ih I; 

nah and an:i.l\ of loral ~r(l\\th. (I nun 111111111.: 

lig . ~ X.I 

G 

of •••• 

"'0'. , 

1) Hal'l11ultds are arnl\<lrc of the tlllrd )r f1Urth I p I 

bt:l 

Sec 333 .Illd 3. 

5 EXPLOIT\TIO. 

1/(/('//111/011 (/l/ro/1I1 arum 1 L 

nIt , 

t r:.t \\ Is and haul 

catch 1)1 rcLn...JlIonal 

,mall boat IlL'.ar hon.: 

fit/( /lUI/Oil 1'/111111< 11 

L'uth a t m l 'nitro 

'c t H {I/1II1I1t n In HaitI 

tom tr.ml 

- 2 fl hm f"('~ 
Haemulon plumieri is fished throughout its range, both commercially and recreationally (Manooch 1976). Lyubimova and Kapote 

(1971) reported that H. plumieri is one of the most valuable fishes 

caught on Campeche Bank and produces the best catches in fairly shallow areas (20-30 m) on the central and eastern Bank. Moe (1963) 

reported that H. plumieri are commonly caught by recreational fishermen on the upper west coast of Florida, often with black sea bass and 

black' grouper, and are the basic catch of party boats in Brevard, Monroe, Sarasota, Pinellas, Levy, Taylor, and Wakulla Counties, Fla. 

Haemulon plumieri is abundant in recreational catches along the southeastern Atlantic coast of the United States, especially north of Cape 

Canaveral (Fishery Management Plan for the Snapper-Grouper Complex of the South Atlantic Region footnote 6). 

Haemulon plumieri also forms large portions of trap catches in the 

West Indies and is the species of greatest abundance (10.74% of the 

total catch) caught in traps on Port Royal reefs (Billings and Munro 

footnote 3). 

See 5.43. 

5.3 Fishing seasons 

On Campeche Bank, H. aurolineatum commercial catches are highest in fall when density is high due to spawning, but occur year-round 

(Sokolova 1965). Olaechea and Sauskan (1974), however, reported 

that H. aurolineatum is most concentrated on Campeche Bank in winter and spring at 20-60 m, especially on the eastern and central Bank. In 

Jamaica, Billings and Munro (footnote 3) reported that grunt abundance was at a minimum between September and November Recreational catches in the northern Gulf of Mexico appear to be greatest in 

winter; St. Andrew Bay anglers surveyed by Sutherland (1977) caught 

H. aurolineatum only from October to March. 

Haemulon plumieri is fished throughout the year, though abundance 

may vary somewhat locally and seasonally. Lyubin10va and Kapote 

(1971) reported best catches on Can1peche Bank in fall and winter; 

Olaechea and Sauskan (1974) reported H. plumieri most common in 

winter catches on Campeche Bank. Juarez (1975) found H. plumieri on 

eastern Campeche Bank only in spring. Anderson and Gehringer 

(1965) collected H. plumieri off the southeastern United States only in 

winter and fall, dUling exploratory cruises. 

See 2.22 and 2.23. 

5.4 Fishing operations and results 

5.41 Effort and intensity 

Catch per effort of H. aurolineatum by recreational anglers in St. 

Andrew Bay was reported by Sutherland (1977) as 0.03 fish/angling 

hour using squid bait and 0.16 fish/angling hour using live shrin1p bait. 

Manooch and Barans' (footnote 5) studies of H. aurolineatum in the 

South Atlantic Bight inclicated that most (67-80%) of the total yield 

could be attained at low values of F (0.2-0.25), whereas three times the 

effort would be required to catch the remaining 20-30%. From an economic and biological viewpoint, Manooch and Barans (footnote 5) recommended that effort in the fishery should be kept low. 

Catch of grunts per unit of effort in the trap and handline fisheries of 

Puerto Rico appears in Table 22; grunt catches in Puerto Rico include 

H. plwnieri, H. aurolineatum, H. jlavolineatum, and H. sciurus (Fishery Management Plan for Shallow Water Reef Fish, Puerto Rico and 

U.S. Virgin Islands9). 

See 4.12. 

32 

Ta ble 22.-Landings per fish trap and per 

handline for grunts, Puerto Rico, 1971-75. 

(Data from Fisher y Management Plan for 

Shallow Water Reef Fish , Puerto Rico and 

the U.S. Virgin Islands text footnote 9, 

table 4-2.) 

Pounds per un it of effort 

Year Trap Handline 

1975 76 4 

1974 74 0 

1973 48 I 

1972 44 3 

1971 36 9 

5.42 Selectivity 

Selectivity of H. aurolineatum catches is probably limited to hookand mesh-size considerations. Billings and Munro (footnote 3) provided some information on fish trap mesh-size selectivity in Jan1aica. 

See 4.13,4.33, and Figure 23. 

5.43 Catches 

Catch statistics for H. aurolineatum and H. plumieri are scanty, 

because separate statistics are not usually reported for this species 

(Courtenay and Sahlman 1978). Grunt landings on the South ~tlantic coast of the United States (primarily H. plumieri and H. 

aurolineatum) appear in Table 23. In the Gulf of Mexico, 

commercial grunt catches averaged 89,600 kg (240,000 lb) from 

Table 23.-Landings of grunts on the South Atla ntic coast of t he United States, 1971-75. (Da ta from 

Fishery Statistics of the United States, 197 1-75, 

U.S. Department of Comme rce, NOAA, NMFS, 

Statistical Digests 65-69.) 

Landings Value 

Year (1.000 kg) (1.000Ib) ($1.000) 

1975 13.6 30 5 

197 .. 130.3 287 57 

1973 31.8 70 15 

1972 19 .5 43 4 

1971 13.6 30 3 

1972 to 1976, with a value of $45,000 (Environmental Impact 

Statement and Fishery Management Plan fo r Reef Fish Resources 

of the Gulf of Mexico'O). Grunt landings in Puerto Rico (H. plumieri, H. aurolineatum, H. jlavolineatum, and H. sciurus) are presented in Tables 24 and 25. 

9Fishery Management Plan for Shallow Water Reef Fish. Puerto Rico and the U. S. 

Virgin Islands. Caribbean Fishery Management Council. San Juan. Puerto Rico. 

Draft. July 1979. 

IOEnvironmental Impact Statement and Fishery Management Plan for Reef F;ish 

Resources of the Gulf of Mexico. Gulf of Mexico Fishery Management Council. 

Tampa, Fla . Draft, February 1980. 
Jahl~ 24. - Quanlll) and ,alul: 1)/ I:ruIII\ landed In PUl:rl<o R,lIl. 

1'I71 7~. \alue, on parcnlht\c\ .. r< ptrccnla~c, or IIllall",hoor~ "'h 

landln~' flf PunIH Riw. (l)ala froom Stulu, ,,/ • i,hl:roe' "r Puertfl 

RicfI. Puc rIO) Ric" Departmenl "f \Aricullurc , .. , appe .. " in I i,h· 

en \Ianaj:unenl Plan f"r Shallo\\ \\altr Red t;'h . Puerlo Ritu 

arod l.S \ irAin "land, Inl '''"In,,\e 'I. lablt' )·3,3--'.) 

YI.'i.lr 

1<)75 

1'1 74 

147, 

1'172 

1<)71 

1<)71 75 

.·\\cragc 

I.anlitn)!, \aluc' 

I I UOO ~g) (I non Ih) 

J 16.~ h<)7 (17) 244 (15) (177 

271 'i 59R (19) IX6 (I'i) () foX 

I X5 2 -lOX (14) 12" ( 12) o fo7 

161 .2 355 fl7) 9'1 (141 001 

187.5 413 (121 I ()\ (101 0.54 

222 5 490 !'il 007 

Table 25.- 0i,tribulion of calch of grunl' b) 

re~ion of Puerlo Rico , J 975. (Oala from Stalu, 

of Fi,herie\ in Puerto Rico . Puerlo Ri co 

Departmenl of Agricullure. a, appea" in Fi,h· 

er) I\ lanagemenl Plan for Shallo\\ \\3I er Reef 

Fi'h . Puerto Rico 3nd l .S. \ irgin 1,land, lexI 

fool note 9. lable 3-5.) 

Landong' Pcn . .'L'nt 

RCgH.ln (1.000 kg) (I.(JOO Ib) IOlal 

North I 8 4 

Soulh 985 217 31 

1ea,1 395 X7 12 

We'l 176.6 389 56 

TOlal 316.4 1:>97 100 

Ih 

035 

032 

() 31 

02X 

025 

ruo 

fla mIl/on (lUr( /111 cllUl 

imul11 annual har\c~t 01 7.f~1()40. 

I i 7i 1'1 

lathom~) oi'lhc IOI.l1 \\Clghl 01 LlllnmCrl.l,lI fJ h lr.I\\, din ur 

off Soulh Camlina \\crc grunt (If. 1Ilim//IIulllllll ,md If I 

with an a\Crdgc \\cight off) 15 k' (0 ~2Ib) (ITtn~ 0 () 0 

0.1-0.510). 

HU(,IIIII/Oll 1II1roltll('li[lIfII cntcr the rc rc.ltlonal fl h I) tim. u 

out Ih rangc. Hcadh\l.lh tWill orth CarollO t) IX ( .w.! 

caughl .In a\Crdgc 01 23.2 t III II (/l/ro/IIIWf/l1Il 10 Iq76 nd I J 

(R. L Dl\l1n pc~. commun a Lltcd In \1anoo h lid B.lr n h!l\[ 

notc 5). Hi/clIluloll (Jumlin('l/II/111 \\.1 th InO I onlln< nl ldU ht 

grunt by these bn.lh. 

Catchc nt H. p/llllliell 111.1\ be 1 • .111/\ 111gb 1111.)( <:.1 

and Tcc \al1 ( 1428) rcpnrkd that /I JllulII' /( \\ rc 

~ccn In l11.lr\..eh in Port u-Pnn(c \Iol(hc\.. and "',1 

r 'pnrtcd Ii. ,,11I1Il/!'r1 and II \, 1L1I1/\ 

CUO.ln Ull11lllCrllal c.ltchc 

I\torc \jU,II1t1tJtI\ c J.H,I ar(' .1\ II bk for I.rtaln 

and Gehringer (1405) rI'port<:d gnml I.lndm' 

CanJ\cml .lrea (l.lhle 2(1) \lthou 'h the IndIO 

Table 16.-Recrealional and commercia l ro,her) calch., or grunt- in Ihe ( "I"' ( .lOla' l-ral, II" .. all,l (I rOm \ 'ldH n 

and Gehri ngtr t <)65.1 

RL't,,;r a 10 11 c lllh '196' 

Spnnt! SU(J'lrcr Fat 

Ll..h .. 'J\I\)n ,,, k~ I~ I~ b 

Clr~ ..lna\ ~rJl ... ,1uthL'rn 

hrh..lgc, and C~llhe\\ J~'" ~"( 10(, 6'4 4' _0 " 6 , 

Cap!! C .JnJ\ CTJI, ,tluthcrn 

l)L'Can [ncr.-. 2'\1 ,~ II~ 

P,lrt CanJ\ erJl, ,"~II.k " ,- It> <Il 2, 40 6-14 I .. IQ , 

PI..\rt Cana\ cr..tl. \,)uhidc XI:> cO ~.< 41~ 1 

t\lpl..' Calla\ cr..tl. n(uth('TIl 

hank, 4. Q LI 1~' 

Clre Cana' er.J1 o\)nhcrn 

hndgL·' 0"'2 I" HI:> 4c4 Q4 ~O , 4' 

CJ.~ Cana\ c!'ra!. nl'rthl:ro uri 41:> 1 

Carl..' Cana, erJI. nl.'rthcrn hoat 

( l 

1%0 
ken down to species, H. plumieri was probably included . Huntsman (pers. commun . as cited in Manooch 1976) reported that over 

475,000 H. p/ul11ieri, weighing about 329 t and comprisi ng 22 % of 

all fish caught in North and South Carolina recreational catches 

from headboats, were caught from 1972 to 1975. Ulrich et a l. 

(1977) found that small catches of H. p/ul11ieri were made in the 

handline fishery directed at snappers and groupers off South CaroI ina: Hael11u/oll p/umieri catches were 90.8 kg (200 Ib) in 1972 and 

272.4 kg (600 Ib) in 1973, with no catch reported in 1970, 1971, or 

1974. Average ex-vessel price for H. p/lIl11ieri taken by these vessels was $0.55/kg ($0.25/lb) in 1976. 

Mean catch rates of H. p/lIl1lieri caught in unbaitcd traps (4.13 

cm mesh) at Port Royal were 265 fish in 1970 and 278 in 1971, 

expressed as corrected numerical catch from 1,000 traps, each 

soaked for I d (Billings and Munro footnote 3). Seasonality, catch 

and ingress, and conspecific attraction affects on trap catches arc 

examined in Figures 31-33. 

0 .7 

.. -;: 0.6 

... ~ 0.5 ... I '" ... '" !! ... ~ 

~ ; ~ :; 

-

0.4 

0.3 

0.2 

0.1 

OLJ-L-L~~~-L-L~~~-L-L~~~-L-L~~~ 

J FMAMJ JASONOJ FMAMJ J ASONOJ F 

1970 1971 

MONTH 

Figure 31.-Variations in month I) mean availabilit) of Haemuloll plumieri 

from trap catches at Port Ro) ai, Jamaica, reefs. (From Billings and l\lunro see 

te,t footnote 3, fig. 4.7.) 

6 PROTECTION AND MANAGEMENT 

6.1 Regulatory measures 

Haelllu/on allro/ineatlllll and H. p/lIl1lieri are subject to regulation under the Fishery Management Plan for the Snapper-Grouper 

Compex of the South Atlantic Region (footnote 6) and the Fishery 

Management Plan for Shallow Water Reef Fish, Puerto Rico and 

the U.S. Virgin Islands (footnote 9). Neither species is covered by 

management measures in the Environmental Impact Statement and 

Fishery Management Plan for Reef Fish Resources of the Gul f of 

Mexico (footnote 10). 

ACKNOWLEDGMENTS 

I thank Donald M. Allen, Joan A. Browder, and Albert C. 

Jones of the National Marine Fisheries Service, Southeast Fisheries 

Center; George C. Miller and Barbara 1. Palko, NMFS, SEFC, 

Miami Laboratory: Elmer 1. Gutherz, NMFS, SEFC, Mississippi 

Laboratories; Gene R. Huntsman, NMFS, SEFC, Beaufort Laboratory; and Eugene Nakamura, NMFS, SEFC, Panama City Laboratory, for critically reviewing the manuscript. 

34 

3 

Dc:( 

~ 

... 

~ 2 IU 

Dei) 

IU 

:z:: 

en 

.... 

d 

Z 

Z 1 

c:( 

IU 

~ 

YAHOO REEF 

APR-NOY 1970 

NASSAU 

/ _----.J 

REEF 

/ / 

/ 

FEB-NOY 1970 

DAYS SOAKED 

Figure 32.- lean cumulathe trap catch and ingres~ of Haemulon plumieri into 

trap' on "iassau and Yahoo Reefs, Port Ro} aI, Jamaica. Solid lines represent 

cumulathe ingress, broken lines cumulatiH catch. The difference bet"een the 

t"o lines for each reef represents e~capement or death in the traps. (From Billings and Munro see text footnote 3, fig. 3.19.) 

... 

Z 1.0 III 

:,;: 

III 

III: 

V 

Z- -% 

>--

...I~ '" C(~ 

00 0.5 zO 

IIIZ 

:1-

() 

III 

III '" 

:I 

'" 

o 1 2 3 4 5 6 7 

NO. OF FISH IN TRAP 

Figure 33.-Effect of conspecific attraction on trap catches of Haemuloll plumieri. If there are more than four individuals of H. plumieri in a trap, the s\,bsequent daily increment of the species increases sharpl). (From Billings and 

Munro see text footnote 3, fig. 3.18.) 
LITERATURE CITED 

\ \ DFRSOI\. \\ \\ 

146, FI,hc, IJ"'~n dUring ... hnmplrJ\\ IlIlg ;llong the ,t,)ulh ALlantlc UJa ... l 01 

Ih~ L nllcd IJI,". 1 431 ') L S F"h \\ IIdl 'icn. SpCl SCI Rcp F"h 

)"0 nO p 

\DERSO\; \I \\ .andJ W GI· HRI\;Gl·R 

1465 8h110gll .. JI-".III,III,,:31 (l.'n"'u", ~)I the ... pC-.... IC ... entenng !", ... hene", In the 

CapcCanJlcr"I.Ir"J' L" F"hWlldl "en . "pce Sll Rep. F"h 51-1.74 

p 

\RV)\ A 

145~ -\ prdlmlll.Jr~ fl'\ h.'\\ III thl' \\I,.',(ern "'llh \tlanill.' lI ... h..:-, l)1 thL' genu ... 

Hd'l/Iul,'/1 BlIli \br SCI Gull C.lflhh ~ 414 -1,7 

-\RA\S.C \ and\ (, BLRRILL JR 

4-'h Prclall~ n.ln Ilndlne, l)r lr.J\\ ling lIn th .... dlnltm::nl,J1 ,hl'lll ff thL' '~lu(h 

C.J,ll'rn l nlkd -tJIC', dunng- h\ur 'l'J',Hh III..)"'~ 11..)"",,) S C \LJI Rl.',\lUr 

Cenl klh Rcr I' In r 

\I dnd \1 0 1.1..:£ III 

hr2 \n n\ (,!'Iu,:oJtHln 01 the- ulhhl1fl..: dl'lll\,'r JI I "h fI .. ",lUfu." tIl Slluth C.lrn 

IInJ S C \IJr Rc,,,ur enl Tldl RI'p ~ 14 P 

1- -\L \1 RI-\Gf::. D " 

Illh9 Rdurn, from Ih 14 b" S ... hl II to.agglng prngr.11ll In\.. I d n~ ~ lUIllUIJtlh: 

• .mal~ 'I III pn.:\ 10U\ rc,ull Fla [)Io?r Jt Rt:,nur \IJI Rl." L .. lh kdl 

Srr 'iQ , r 

BH:BF \\ dnd J THe -\ 

IY~ fhe..' I ,he" )' Pllrt JU Pran\..(' RJ\ H.tlll "llh ,I 'Ultllll.lr\ tIl the ~Ihl\\ 1\ 

~l.K of n'Urine II h ,lithe I la.,d (If HJIII ;JnJ SJnt~) I)llfl' n I, Illnl\\gll.a 

\ , 10 1-~7'1 

'Q3, r:II.::1dhooJ...ofrhl.: hllft::11 he (1 Bc.:r'1wJJ l. P PutnJIll'I\\lIl .... 

33- r 

B~\ FRTO R J H .nd" J Hill I 

1'1'1) A rnl 

popul." n 

,"mplmg 

I!()FSCH. D 1-

" )f f11CthLld t Ir I.: IImatmg morlJI t\ rale n c'pl,)ltc 1 I h 

\\lIh p\.'\..,-.l1 rCfl.n.:nll. I,) ,oun.:e ,l, h j, III I,.all.'h 

R"pr r \ R,un C. ~ IN r p~ I, \I r 1-IIlI I h7 

1'1 ~ \rrlKdllon of nUIllCII al.IJ "'lall"n In l'",I"~I,.1 Inll'sllg .• II"n, 01 

.. alcrpollull<," FP-\ IJ{lIll r 0\, Ilol Rc S" 114 P 

BuHLKE J r "nd l C (j (H \PI I 

I 'It, f I h", n<the I!Jh~ " "nd ad Jl nl !rl1Plldl \\"l., I Inn Punl 

Co \\ nne" .. xI Pd ~"I P 

BRA D,I;I). J " 

I'IM C,(n dr.' de nomc do p<'I\CS f".'rlugu 

hlad Pe ld 414 J I 4(, 

BRI l>FR JR 

1';14 f 1I.:ld buvl rrwr n.: 'l!rIohc II I t; \11 nOlL lU I Irom L II r.ldor In 

Ie J (J P Putna !rio ~llO '1" P 

J ( DA" \ 

C 'l UP\ In' "".1' <.I t'll.J III thl. 

['dn Arr II h S"l 1112511 

-' ( he \\ ... I' In\\<.:1 (I.lrJLn K""d r'd 

Vt.c II.: rn (Inb~c.:.iJll Ph [) 1 hl I' 

t 0: ~-l r 

1471 \lflallun IfltrJ c Inlt.:r p4..l.I1II.;J lh .. hl.;'l1lll'ltlhln;J \- prollln;JS lkl 

rlJ nlu ~L:na L r~clc, dd nl:f( lIa l1IullIlI In "pJIl I'ngl 

~ 'r I [1,,1 In I ()le ,'r l nlV One Ole 1111: 7'1 ~ 

( f ~\ I(,(J \1 r 

Illhh I" pt.:'-.\: mOlr .l,J \t.:n\:/ud.1 \111 II l-unJ IS .. JIL (Icn,- ~,Ilur 

'&.IraI.;J' \10r1o 'r 11 44Y ')~ 1 

(HIIIl-'I'[)f \If JK ."ndJU:-1d\(IiK\N 

IY7h (Of! po Ilion l:lOlllg\. dn<J J)'nalllll' II dt.:nll.;r\alll!t.!a lOl1ll11unttic\ on 

Ihl.' ntlrlh\l.t;stern (Jull 01 \!1c.:XII,.O ulntancllIal hl'lI. v.lth 'Imilar '}1l0P'I'" 

for Ihe enllre (;ull kx A&M L nl\ Sca G,anl Coli lAM SC, 76 2(JX. 

10-1 P 

C!)I LETt r I! Hand r: H TALBOl 

JY7~ Adl"ll) pattern ... 01 (oral rt,;cI I"he\ \l,-Ith cmrha .... i ... on noc.:turnal 

diurn,,1 ehangC()\Cf /11 B B Collelle and S A Earle (edllo,s). Resulls ollhe 

Tckllic Program ecology 01 LOrdi reel ",he,. p 98 124 Bull Los Ang 

OJ Mus N., H", &, 14 

COURTENAY W. R JR 

1961 WCSlern AilanliC fi,he, of Ihe genu, HU(,Jllu/o" (Pomada'Yldae) sys 

Ie malic "alus and Juvenile p'gmenlal,on Bull Mar Su Gull Canbb 

II 66 149 

35 

COURTENAY. W R .. JR . and H. F. SAHLMAN. 

1':178 Pomadas) Idae. III W. Fischer (edilor). FAO species identificalion 

sheelS fOI fishery purposes. WeSiern Cenlral Allanlle (Fishing Area 3 1). Vol. 

4. unpaglnaled. FAO. Romc 

CRA IG. A K 

1476 Trapping esperimenls wilh snappe.s III soulh Flonda /11 H. R. Bullis. 

J r and A C Jones (edllors). Coll04UIUlll on snapper·grouper fISher) 

n.: .... oun:l' ... nrthl' \\ ~ .... Il'rn (e!llm] Allantlc Ocean. p ::!::!2-2J6. Fla. Sea Grant 

Rep 17 

CUtl-Itl-1i S. R . JR. J B RIVERS. and P STRUHSAKER. 

1462 Snappcr Ir.m ling esplmallons along Ihe soulhcaSlcrn coasl 01 Ihc 

L1nlled )Iales C(ll11mer Fish Rev 2-1(12) 1-7 

D.\\ IS. \\ P 

14f',7 f..l'olnglLal InICrJdIOI1\. comparall\'C bIOlogy anJ c\'olution,H} trclllb 

01 II1Irleen p<ll11ada', Id t"he, 0' AIII),!cII<lI" Reel. Flonda Kcys Ph D The-

'IS. L nil \I,anll CHal Gahles. Fla. 128 p 

[)co L \\lAn R F D and \\ R OURTFI\AY JR 

Il)74 h\h \l.ule\ <.I, \cen by \canning electmn micro~copy. Fla SCI. 

n 141 I-Ill 

I D\\\RDS R R C 

147.' PltlJuLlwn L'L·lllog:-. oj t\~~) Carinbcan manne CCO\)~tC1ll'. I Ph).'IL'ul 

I'nl II '"11111' 111 "lid lallna hlllanne Co"slal ~lal S" I ,m 3'8 

I RD\l \1\ [) S 

1<150 Recenl II,h 'l'cord, Inllll PUCrio RIC" Bull Ma. Sci GUll Canhb 

h ~ 15 '-Ill 

1474 ~pll1brc, \ulgJrl.':-' Je PCl.'l'''' en Pucrlo RIL'n CtlOlrib Agropcc Pl''''lj 

Sen \1I\ Clnl Ocp Ag"" 6(2) 50 p 

1477 Sp.1I1 !ling pallern, 01 II,h I fllIlI Ihc nonheaslern Canbbean. /11 H 8 

Ste",Jr'. Jr (COIIO!) Coopl.'ratl\c imc\tlgation, or Ihc Callohcan and au.lueenl regll,"' \I Car"ca,. VenelUcla. 12-16 Jul) 1976. p. 145-169 FAO 

hsh Rcp 200 

f·\ I R\l,\, " B \\ .• Ind \I C \IARSH 

1'1112 T he I "hes 01 PoriO R,co. Bull l.S. F"h ollllll 20. PI I. 1900:-19-

,50 

FERRf::IRA Dl' ME E7l'S. M 

1970 Alllncnlacaoda serra Snomberomorus maCl/lalUS (Mllchell). em aguas 

COsleWJs J., c"ado do Ceara Arq Clenc Mar 10: 171 ·176 

FIl-[)L~R R H. \1 J LOBEll "ndC R LUCAS 

14-17 1he I"henc, and IIshcn resources ollhc Canbhcan area U.S. Fish 

\\ !ldl Scn . F"h Lean ~'i<l ~ 10 p. 

G \1 LO\\A\ J 

1941 LClhal cllcu "I Ihe cold" inlcr 01 1939-10 on manne fishes al Key 

\\c',. Flonda Copela 1941 I 18-119 

Gll \tURF, R G L H BULLO K. and F H BERRY 

1<178. Hy pOlhermal mOrlalll) In mannc IlShes of soulh-cenlral Florida Janu -

dr) 14T 1\<>rlhl''''' Gull SCI ~ 77 ll7 

GIr-;SBl'RC< I 

144X The 'rCllt.:' oj' B£llli\'\1011l0 (PI ... ce\. Hal'lllUlllnidac) Z{)(ll{lgica 

(\ \ ~:; 1'iI I 'it> 

f.RI\IIS.C B 

IY71 Thl'rll1al .H..IJllinn \tUJll' .... 01 the Cry\tal RiYl'r ~tcam electric \taI"" [IJ Dcp al Resour Mar. Re' Lab. Prof Pap Sor 11.53 r 

(,\<I\lIS ( B andJ A M()L1N1AIN 

1<)71 [lltTI ollhl'rm.ti dfluc..:-nt upon Illannc r"hc~ ncar the Cry .... tal Ri\cr 

"e.lln dec 1"11 "allon FI" Dep Nal Re,ouf \lor. Res Lab. Prol Pap 

Sc. 17. h-l P 

(,L 1)( ~R I \\ 

Iq~, \ put! h~aJcdgrunl Hal'lliu/ollphlll1il'1'I Am r..1u\ N~)\it 607.op 

(,llll"RJ "ANDA). n J 

1971 Un nuevo ~1 ... tt:ll1a para arlllar rl'uc", de 1l.'IIt>plancton. 111 S)-mpo~IUIl1 on 

Im .... sllgi.Jtlfln' and rl' ... ourl.'C~ of the Caribbean Sea and adJ4ll.'cnt region ..... p 

-14945'1 f-.\(). LIN fSCO. Pans. 

GLJITILRRl:1 B. P 

147h Comro:-.iuon arpnnimaJa Lic algullo\ PCl'C\ de Venezuela Fund 

1."SalkC,enc Nal 16(104) 165·174. 

HASlINGS. R W. 

1972. The origin and scasonal,ly 01 Ihe fish fauna on a new Jelly In Ihe nOrlheaslern GulloI' Me\lco. Ph D. Thes". Fla Slale Un" . Tallahassee 555 p 

HASTiNGS. R w.. L. H OGREN. and M . T MABRY. 

1976 Ohservallons on Ihe fISh fauna assoclaled wilh offshore platforms in the 

norlheaslcrn Gulf 01 MC.\lco FISh. Bull. U.S. 74 387-402 

HERCULANO SOARES. L .. and H De HOLANDA LIMA. 

1966 SObrc a biomelria da xira. Haell/II/ol1 aurolillealllll7 Cuvier. da COSla do 

e"ado do Ceara Bol. Ins!. BioI. Mar. Univ. Fed. Rio Grande do Norte 

3:39·52 
1967 . SObre a biomelria da xim, Haelllll/oll allro/illealLilll Cuvler, da CO'la do 

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HILDEBRAND, H. H 

1954 . A sludy of Ihe fauna of Ihe brown ,h rimp (Pellaells a: lecu; Ives) 

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1955 . A slud) oflhe fauna oflhe pln~ shnmp (Pellaell.1 dllora/'ll/ll Bur~enroad) 

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,. 232. 

HILDEBRAND, S. F., and W. C. SCHROEDER 

1928 . Fishes ofChesapea~e Bay, U.S. Bur. FISh. Bull. 43(parl 1),366 p . 

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1977 . Fis hes o f Ihe Gulf of M eXIco, Texas, LOUISiana , and adJacenl 

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HUNTSMAN , GR. and C S. MANOOCH III 

1978 M,nllllum Sl7e 1111111, for reef fishe, Proc Annu ConI' Soulheasl 

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1893 A rCYle\\ of Ihe sparo ld fishe, 01 Ame n ca ano Europe Rep. Comm 

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JORDAN , D Sand J. SWAIN 

1884 A rev le\lo of Ihe spec ies oflhe gen us Hoelllllioll Proe U. S NaIl Mu s. 

7:28 1-3 17 

J UAREZ, M . 

1975 . DISunbucltin cuamilaliva y a lgu nas aspeclOs cuaillall vos del IcIlOplanclOndel BancodeCampeche. Re\ In \e,1 In sl NaL Pesea 1(1)'~7-71 

KAPOTE , A M 

1971 On the age 01 the main commercial bottom I \,.,ht.!~ of Campcche BanI-.. 

lin Russi III A S Bogoano\ (eollor), SO\cIs~o- ublns~i)e r)b. issku 

IS", lei-Cuban fisher) In\esllgallonsl, (\,0 3 p. 77-81 ~IOSL'O" 

KONCHINA . Yu \'. 

1976 . The S) slemalics anu d"lribul lon of Ihe grum famil) (Pomadas) Idaci 

lIn Russ I VOpnlS) I~hll ol 16 '97 1-9 8 (Transl J lchlh)ol 16883-9001 

1977 Some uala on Ihe blolog) 01 grums (Famlll Pomadas) Idael lin 

Russ. I VoproS) I~hllol 17 '6~ 1-633 (Transl J khlh) 01 17 548-558.) 

KORZHOVA, Yu A 

1965 Tcchmcal and chemical chartlctcti. ... tlc~ {)j ... omc u)lllmcn:wl fi .... h and 

ill\enebrale.s 01 Ihe Gulf 01 MeSIC" anu Ihe Canbhean Sea III A S 

Bogdan(" (cdllor). SO\ leI-Cuban lI,her) Il1\esligall(lnS, p. 3n-340. (Engl 

lransl b) Israel Prog. Sci Transl , Jerusalem. 1969) 

LIVINGSTON, R J 

1976 Diurnal and .... ca~onal nuctuation~ l)f org •. mi~nh in a north Florlua ~~tuar) Estuarine Coaslal Mar SCI 4:373 .jOO 

LONGLF), W H , and S F HILD EBRAND 

19-11 S}~(elll,.ltIC catalogue 01 the fl~he~ of Tortuga~. Flonua. \\ Ith oth~n alions on color, habil'. and local dislnbullon Carnegie Insl. Wash. Publ 

535 1-33 1 

LYUBIMOVA, T G , and A. M . KAPOTE 

1971 Biological characteri~tlc~ of ~omc ~pecle~ of commercial bottom fI",he~ 

from Ihe Bank 01 Campechc. lin Russ] III A S Bogdano\ (eullOr), 

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MANOOCH, C. S, Ill. 

1976 Age. grovvlh and mOrlalil) of Ih~ "h,le gru nl. Hliell/lllol1 pllli/lleri 

Lacepede (PISces: Pomadas) idae). from North Carolina and Soulh Carolina. 

Proc Annu. ConI'. SOUlheasl. Assoe. FISh Wilul Age nCies 30:58-70 

1977 Food~ 01 the red porg). Pagni., pogrll.' Linnaeu~ Pi~ ce~: Sparidae). 

from North Carolina ano Soulh Carolina Bull Mar SCI ~7 776-787 

~lANOOCH. C S, III. and C A. BARANS 

1979. Gro\\lh and Yleld/recruil o f IOmlale. Hoell/lllol1 ollm/il/ellllllll. all 

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MANTER, H W 

1954 . Tremaloda oi'l heGulf of Mexico U,S. F"h Wildl . Serv., Fish . Bull . 

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McFARLAND, W. N 

1980 . Observallons on recruilmenl In haemulio fishes. Proc. Gulf Caribb. 

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McLEAN, R., and W. F. HERRNKIND. 

197 1 Compacl sc hooling during a mass mo\e me nl by g runls. Copela 

1971 :328-330. 

MILLER. G. C .. and W. J. RICHARDS. 

1980 . Reef fish habi!al. faun a l assemblages ano faclOrs oelermlning dislribu -

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Annu. Meel. , p. 114- 130. 

36 

MOE, M . A., JR. 

1963 . A survey of offshore IlShlng In Florida Fla Board Conserv. Ma 

Res Lab, Prof Pap Ser. 4, I 17 P 

1966 Tagging fishe, In Flonda o(hhore walers Fla Board Conserv Ma 

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1967 Prolonged surV ival and mlgrallon 01 three lagged reef fishes In Ihe Gul 

of Me"co. Trans Am FISh Soc 96228-229. 

1972 Movemem and mlgrallon of soulh Flonda fishes. Fla . Dcp Nal 

Rcsour Mar Res. Lab . Tech Ser 69,25 p. 

MOTCHEK , AD, ano A F SILVA Lee. 

1975 Conducla socwl oel genero Haell/lllol1 Acad Cienc Cuba Ser Bioi 

~7 I 10 

MUNRO, J L. , V C GAUT, R THOMPSON , and PH REESON 

1973 The spawning ,ea,ons 0 1 Canbbean reef fishes J Fish BIOI 5 :69-84 

NAHHAS, F. M , and R M CABLE 

1964 Dlgenelic and aspldogaslnd IremalOdes from manne fi'hes of Curaca 

anu Jamaica Tulane Slud Zool I I 169-228 

OGDE J C 

1977 Bchavlou r ano eeolog) of schooling groups of Juvenile grums (Porn. 

ua ,y ,dae) . III H B. Slewarl, Jr (eunor). Cooperallve Invesllgallon, 01 Ih 

Canbbcan and adJacem regions II Caracas. Venezuela, 12- 16Jul) 1976. 

323-3 \2 FAO F"h Rep . 200 

OGDEN, J C , and P R EHRLICH 

1977 The behaVior 01 helerolYPIC resl Ing school, ol)u \ enlle grunls (Pomada 

'J,uae) Ma r Bioi !Ber! ),\2 27~ 280 

OGRE L. H .. anu H A BRUSHfoR 

1977 The u"lnbullon and abunuance olI fishes "ughl" IIh a tra,,1 111 Ihe SI 

\nure\\ Ba) s)Slem, FloriJ" 1\.or!hc,,-sl Gulf SCI I 83-105, 

OLAECHFA, A . anu C HER 'A 'DE? 

1976 E\aluau(ln ue la' pllhlaCllml:" Jc jenlgua no H£1emlilon aliro/l1lealUI1J 

pel pluma Clllalllll.' 1I{lt/O.HI.' en cl Banl:o de CampeLhe In t j\;ac Pc ... c 

Cuna Cenl. In \esl Pe'q Res. Inv e,1 ~ 1'i7-16U 

Ol.Af:CHEA, A .. anu \ I 'iALShAr\ 

I Y7.\ Cartas ue pesca uel Banco ue Campeche, 197~ Resumenes Invesl 

In sl Nac Pesca Cem In vcsI Pe sq I 176-llll 

PARRISH J D. anu R J ZIMMER IA 

1977 Ullli/alhm b) I"hes 01 space anu I,")o resources on an olt\hore Puerl 

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PIERCE. E L.,.'R 

IY36. Rales ofd'gesllon in Ihe yello" lall (OCYUrIIl c/rrnurIIs) and Ihe "h'le 

grun l (Ho('//wlolll'llIIlIiail. Cnpela 1936 I~J I~4 

RA NDALL J E. 

IY63 An analy"s 01 Ihe f"h populallons of arllfietal and natural reef, 111 Ih 

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IY67 Foou hablls 01 neef fishes or Ihe \\"'1 Indies Slud Trap. Oceanogr 

(M iamI) 5665-847 

1968. Caribbea n reef fishes T.F.H PUbI.Ine.,NeptuneCIl),NJ ,318p. 

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1954 An eco logical SIUU) of Ihe Gul I' llf Me\lco fishes, In Ihe \ IClnll) a 

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1980 . A IISI of common and sc,enllf,c names of fi,hes from Ihe Unlled Slales 

and Canada. 41h ed Am. FISh Soc . Spec. Publ . I~, 174 p. 

ROESSLER. M A 

1970 Checklisl of fishes In BUIIOn\\ood Canal. E\erglade, Nallonal ParL 

Flonda. and ob~e r\"atlon:-. on th e "ea~onal occurrence and life hi~rorie" (it 

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ROITHMAYR. C. 1\1 . 

1965 Indu slrial bOllamfish fisher) of Ihe northern Gulf of MexiCO, 1959-

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1975 . Desc nplion of egg' and lanae of laboratory -reared "hile grunl. 

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SAUSKAN, Y.I., and A . OLAECHEA 

1974 . BIOlogy and populalion abundance of Ihe IOmlale (Haelllllioll 

aurolilleollllll Cuvier and Valenciennes, 1830, Pamadas) idae. Perciformes) 

on Campeche Bank. lin Russ.] IliA. S. Bogdano\ (edilor). So\'els~oKubins~iye ryb . issled lSov iel-Cuban fishery invesligalions]. No.4. p. 1:15-

150 . Mo,cow. 

SEKAVEC, G . B., and G. R . HUNTSMAN . 

1972 . Reef fishing on Ihe Carolina conllnemal shelf. Proc . 151h Annu . Inl. 

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,q':! pacC' re:-.ource ~hanng In a coral reef rl~h communl(~ In B. 8. Collette 

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S\ lITH. G B. 

I'Po Ecolog) and d"tnbullon e,f ea~tcrn Gull ell ~\e"co red I"he, Fla 

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S\IlTH. G B. H. ~I \LSTI' . \ BORTOl\,E. R. \\' HASTI'JG . and L H 

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19"5 F"h" of the Flonda \llddle Ground <11th commenh 1m ecolog} and 

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S\lITH. G B. D. \ HE:"SLE) , and H H \t,\THE\\ S 

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S\ IlTH R L 

'40' orne a"ped.., I,.)f pn)tcln dlge\IIon In the \,hllc- grunt. Haollu/oll plumltri Copela 1967 -lo -~-l , 

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IQ"'O PeplidJ'c actl\ It~ 10 Hu( mu/"" riuJ1IIt.'ri (PI'CC... PornadJ-

,) Idacl. Copela 1<l'0"40 741 

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1965. DI\tnbutlf.ln and bH,,)k1glcal char .. h.'tcri,tlC' \,)1 the main ((lmlllcfuJI ri~h 

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1960 An ecologICal ,tud) of thc I"he, 01 thc Tampa Ba) area Fla . DcI' . 

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1966 ,'\'ght hablh nll"he 01 Alligator Reci, Flelrlda khth,e,1 Aquarium 

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STO 'E R B. H L. PRATT, R 0 PARKER, JR and G E DA\ IS 

1979 A compan,on e)1 II h population, on an arllllcwi and natuml reel In the 

FlondaKc), "'iar FI h Rc\ 011(9) ' 1-11 

37 

STOREY, M . 

1937 . The relation between normal range and mortal it) of fishes due to cold at 

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STRUHSAKER, p 

1969. Demcr;al fish resou rces composition , distribution , and commercial 

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1977 Catch and catch rate, of lishes caug ht by anglers 10 the St. Andre" Ba) 

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TABB.D C. and R B MANNING 

1961 A checkli,t of the nora and rauna of northern Flonda Ba) and adjacent 

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1904 Preliminary explomtor) '"hlng on the Flonda west coa'L Fla. Board 

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1'i77 Statu, reror! on the commercial ,napper-grouper li,herie, off South 

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\ALDES ~ILNO Z. F . and A F SILVA LEE 

1977 AltmentaclOn de lo~ plxe~ de JrreLlfe~ artlflClalc~ en la plataforma ~uroccldenlJl de uba. lin Span .. Engl ,umm J Acad Cienc. Cuba Inf CientTecn 24. cl r 

\\ANG, J C S, and E CRANEY 

1971 D"tributlOn and fluctuation, In the fish fauna or the Charlotte Harbo r 

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1979a Re,ulh of MARMAP otter trawl 100eSllgatlOn, In the South AtlantiC 

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1979b. Re,ults 01 MARMAP otter 1m" I Insestlgation, In the South AtlantiC 

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1979c Re,ult, 01 MARMAP otter trawl investigations In the South AtlantiC 

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1979d Rc,ults of MARMAP otter trJwl ine'eotigations In the South Atlantic 

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4-1.59 P