NOAA Technical Report NMFS 44

FAO Fisheries Synopsis No. 148

Synopsis of Biological Data

on the Porgies, Calamus arctifrons

and C. proridens (Pisces: Sparidae)

George H. Darcy

September 1986

u.s. DEPARTMENT OF COMMERCE

Malcolm Baldrige, Secretary

National Oceanic and Atmospheric Administration

Anthony J. Calia, 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 NMFS 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.

II
CONTENTS

Introduction

1 Identity

1.1 Nomenclature 1

1.11 Valid name 1

1.12 Objective synonymy 1

1.2 Taxonomy

1.21 Affinities 1

1.22 Taxonomic status 3

1.23 Subspecies 3

1.24 Standard common names,

vernacular names 3

1.3 Morphology

1.31 External morphology 3

*1.32 Cytomorphology

*1.33 Protein specificity

2 Distribution

2.1 Total area 4

2.2 Differential distribution 5

2.21 Spawn, larvae, and juveniles 5

2.22 Adults 5

2.3 Determinants of distribution changes 5

2.4 Hybridization 7

3 Bionomics and life history

3. 1 Reproduction 7

3.11 Sexuality 7

3.12 Maturity 8

3.13 Mating 8

3.14 Fertilization 8

3.15 Gonads 8

3.16 Spawning 8

3.17 Spawn 9

3.2 Preadult phase 9

3.21 Embryonic phase 9

3.22 Larvae and adolescent phase 9

3.3 Adult phase 9

3.31 Longevity 9

3.32 Hardiness 9

3.33 Competitors 9

3.34 Predators 9

3.35 Parasites, diseases, injuries,

and abnormalities 9

*3.36 Chemical composition

'No information available.

III

3.4 Nutrition and growth 9

3.41 Feeding 9

3.42 Food 10

3.43 Growth rate 10

3.44 Metabolism 10

3.5 Behavior 10

3.51 Migrations and

local movements 10

3.52 Schooling 11

*3.53 Responses to stimuli

4 Population

4. 1 Structure 11

4.11 Sex ratio 11

*4.12 Age composition

4.13 Size composition 11

4.14 Subpopulations 14

4.2 Abundance and density 14

4.3 Natality and recruitment 16

4.31 Reproduction rates 16

4.32 Factors affecting reproduction 16

4.33 Recruitment 16

*4.4 Mortality and morbidity 16

*4.5 Dynamics of population 16

4.6 The population in the community

and the ecosystem 16

5 Exploitation

5.1 Fishing equipment 16

5.2 Fishing areas 16

5.3 Fishing seasons 17

5.4 Fishing operations and results 17

*5.41 Effort and intensity

*5.42 Selectivity

5.43 Catches 17

6 Protection and management

6.1 Regulatory measures 17

*7 Pond fish culture

Acknowledgments 17

Citations 18

Synopsis of Biological

Data on the Porgies,

Calamus arctifrons

and C. proridens

(Pisces: Sparidae)

GEORGE H. DARCY

Southeast Fisheries Center

National Marine Fisheries Service, NOAA

75 Virginia Beach Drive

Miami, FL 33149-1099

ABSTRACT

Information on the biology and fIshery resources of two common species of western

Atlantic porgies, Calamus areli/rons and C. proridens, is compiled, reviewed, and

analyzed in the FAO species synopsis style.

INTRODUCTION _

Porgies are small to moderately large spiny-rayed fishes of the family Sparidae. They inhabit temperate and tropical marine and

estuarine waters from shore to depths of over 100 m. They are

ecologically similar to snappers (Lutjanidae) and grunts (Haemulidae). Most are excellent food-fishes and are of considerable commercial importance (Randall and Vergara R. 1978). Many are also

important to recreational fishermen.

The genus Calamus consists of about 13 species, all of tropical

and subtropical American waters. Two species occur in the eastern

Pacific, and 11 occur in the western Atlantic (Randall and Caldwell

1966). Eight species are known from coastal waters of the United

States (Robins et aI. 1980).

The grass porgy, Calamus arctifrons, and the Iittlehead porgy,

C. proridens, are two common porgies of the western Atlantic

Ocean. The grass porgy is essentially restricted to the northeastern

Gulf of Mexico and is fairly abundant in shallow water off the northwest coast of Florida. The Iittlehead porgy occurs on the east coast

of Florida, in the northeastern Gulf of Mexico, in the Greater

Antilles, and on Campeche Bank where it is one of the most important commercial fishes. This paper summarizes information on

these species.

1 IDENTITY

1.1 Nomenclature

1.11 Valid name

Grass porgy, Calamus arctifrons Goode and Bean, 1882 (Fig. 1):

425, type locality: Pensacola, Florida. The name comes from the

Latin calamus (a reed or quill), referring to the quill-like interhaemal

bone (Randall and Caldwell 1966), and arctus (narrow) and frons

(forehead), referring to the dorsal profile of the head.

Littlehead porgy, Calamus proridens Jordan and Gilbert, 1884 (Fig.

2): 150, type locality: Key West, Florida. The name comes from

the Latin calamus (see above) and prora (prow) and dens (tooth),

referring to the projecting canine teeth (Jordan and Fesler 1893).

1.12 Objective synonymy

Calamus arctifrons Goode and Bean, 1882

Calamus proridens Jordan and Gilbert, 1884

1.2 Taxonomy

1.21 Affinities

Suprageneric

Phylum Chordata

Class Osteichthyes

Superorder Acanthopterygii

Order Perciformes

Suborder Percoidei

Family Sparidae
Figure I.-Adult Calamus aretifrons. (From Jordan and Evermann 1896,

fig. 550)

Generic

Systematics of the genus Calamus have been badly confused in the

past (Randall and Caldwell 1966). Reviews of the genus have

appeared in Valenciennes in Cuvier and Valenciennes (1830),

Guichenot in de la Sagra (1843), Guichenot (1868), Poey (1874),

Jordan and Gilbert (1882), Jordon and Fesler (1893), Jordan and

Evermann (1898), Evermann and Marsh (1902), Longley and Hildebrand (1941), and Randall and Caldwell (1966). Species are often

difficult to differentiate, and are especially difficult to identify when

small. According to Randall and Caldwell (1966), 13 species of

Calamus exist. All are American, with two species from the eastern

Pacific, and 11 from the western Atlantic.

The genus is characterized as follows: Second interhaemal bone

enlarged and hollowed anteriorly, receiving posterior portion of

swimbladder; dorsal fin preceded by a small antrorse (forwardprojecting) spine, often hidden by scales; temporal crests thin and

high, joining lateral crests, which form part ofthe orbit above middle

of eye, both crests coalescing with supraoccipital in narrow anterior part of interorbital; body deep, the depth contained 1.8-2.5

times in standard length (SL), and compressed, the width about 6

in SL; head large, its length 2.8-3.6 in SL; dorsal profile of head

from moderately to very steep; eye elevated, the suborbital depth

5.4-12.5 in SL (often greater in adults than in juveniles); mouth

terminal, not large; anterior teeth in jaws conical, those on sides

molariform, in two full rows in lower jaw and three rows in upper

jaw (teeth in middle row small); a bony tubercle proj~cting

downward from lower anterior portion of maxillaries; prefrontal

with a bony tubercle above posterior nostril, this tubercle much

more conspicuous in some species than in others; posterior nostril

long and slit-like; preopercle and opercle entire, the margins smooth;

dorsal fin rays XII (rarely XI or XIII), 11 or 12 (rarely 13); anal

fin rays III, 10 or 11 (rarely 9); pectoral fin rays 14-16 (rarely 13

or 17); pelvic fin rays 1,5; 15 branched caudal fin rays; body completely covered with weakly-ctenoid scales; head with scales on

opercle, preopercle, interopercle, and extending anteriorly from

nape into interorbital region; no scales on dorsal and anal fins except basally on last two rays; scales extend about two-thirds of way

onto caudal fin and occur basally on paired fins, dorsal fin slightly

notched between spinous and soft portions; dorsal fin relatively low,

the longest spine (third or fourth) contained 5.6-9.6 in SL; pectoral fin length 2.1-3.6 in SL; caudal fin forked; gill rakers short,

9-14 on first gill arch; vertebrae 10+ 14; three predorsal bones;

Figure 2.-Adult Calamus proridens. (From Jordan and E"ermann 1896,

fig. 547)

dorsal fin support count 0-0-0-2 (Randall and Caldwell 1966). The

skull was discussed by Jordan and Gilbert (1882), and illustrated

by Gregory (1933).

Specific

The most important characters for separating species of Calamus

are: Number of lateral-line scales; total number of pectoral fin rays;

number of dorsal and anal fin rays; dentition; eye diameter; suborbital depth; length of pectoral fins; length of the longest dorsal

spine (usually the third spine); body depth; color; and steepness

of profile of the head (Randall and Caldwell 1966).

Calamus arctifrons-The following species diagnosis is from Randall and Caldwell (1966): Dorsal fin rays XI to XIII (usually XII),

11 to 13 (usually 12); anal fin rays III, 10 or 11 (usually 10); pectoral fin rays usually 16 (sometimes 15); lateral-line scales 43-49

to base of caudal fin; total gill rakers in first gill arch 10- 12 (usually 10); canine teeth in front of jaws about equal in size; no small

molariform teeth medial to the three rows at sides of upper jaw;

pectoral fins short, their length 3.0-3.6 in SL; eye small; suborbital depth moderately great, 6.6-7.8 in SL; body depth 2.0-2.5

in SL (depth relatively less in adults than juveniles); longest dorsal

spine 6.4-8.9 in SL; dorsal profile of head usually smoothly convex, the portion below eye forming an angle of about 50-57° to

the horizontal (taken from tip of snout above upper lip to mid-base

of caudal fin); maxillary tubercle with a free semicircular margin;

prefrontal tubercle not well developed.

Calamus aretifrons is very similar to C. campechanus, and probably evolved from a common stock in the not too distant past

(Randall and Caldwell 1966). It is distinguished from C.

campechanus by its smaller suborbital depth, higher number of pectoral fin rays, lower number of gill rakers, and lower dorsal profile of the head below level of the eye (Randall and Caldwell 1966).

All specimens of C. campechanus examined by Randall and

Caldwell had irregular horizontal lines on the unscaled portion of

the cheeks, whereas only a few specimens of C. arctifrons had these

lines. The two species are separated geographically, with C. campechanus restricted to Campeche Bank. Environmental conditions

such as tubidity, temperature, and salinity apparently exclude both

species from the western Gulf of Mexico (Randall and Caldwell

1966).
Calamus proridens-The following species diagnosis is from Randall and Caldwell (1966): Dorsal fin rays XII or XIII (usually XII),

II or 12 (usually 12); anal fin rays III, 10 or II (usually 10); pectoral fin rays 13-15 (usually 14); lateral-line scales 52-57 (usually

54-56); total gill rakers in first gill arch 11-13 (usually 11); fourth

canine tooth from symphysis of upper jaw enlarged and strongly

outcurved in adults (enlargement begins at about 100 mm SL and

is apparent at sizes at least as small as 148 mm SL); a group of

small, molariform teeth medial and toward the front of the three

rows at sides of upper jaw, forming a partial fourth row at front

half of sides of jaw; pectoral fins long, 2.5-3.0 in SL; eye small;

suborbital depth relatively great; body depth 1.95-2.20 in SL;

longest dorsal fin spine 6.1-8.7 in SL (in specimens>100 mm SL);

prefrontal tubercle above posterior nostril not well developed; largest

molariform tooth usually smaller than fleshy width of maxillary

tubercle; dorsal profile of head steep and relatively straight, portion below eye of adults forming an angle of 58-65 0 from horizontal and first third above eye an angle of 43-69 0

•

Calamus proridens is closely related to C. pennatula, but differs

in having a steeper profIle of the upper part of the head, greater

average body depth, greater average length of the longest dorsal

fin spine (third or fourth) and slightly higher lateral-line scale counts

(Randall and Caldwell 1966). Large specimens of C. proridens

develop a large, bulging forehead not found in C. pennatula.

1.22 Taxonomic status

Both C. arctifrons and C. proridens are considered morphospecies.

Calamus arctifrons is very closely related to C. campechanus, but

they are considered separate species (Randall and Caldwell 1966;

Randall and Vergara R. 1978).

See 1.21.

1.23 Subspecies

No subspecies of C. arctifrons or C. proridens are recognized.

1.24 Standard common names, vernacular names

Calamus arctifrons-The accepted common name for C. arctifrons

in the United States is grass porgy (Robins et al. 1980), and standard FAO common names are: English, grass porgy; French,

daubenet cendre; Spanish, pluma negra (Randall and Vergara R.

1978). Other names appearing in the literature are shad porgy (Jordan and Fesler 1893; Evermann and Marsh 1902; Beebe and TeeVan 1928), pez de pluma (Duarte-Bello 1959) and porgy (Springer

and Woodburn 1960). According to Moe (1963), C. arctifrons, C.

proridens, and C. calamus are caught on the snapper grounds off

the west coast of Florida, and the names bank porgy, Key West

porgy, grass porgy, and white snapper are applied to all three

species, although Key West porgy usually refers to C. nodosus.

Calamus proridens-The accepted common name of C. proridens

in the United States is littlehead porgy (Robins et al. 1980), and

standard FAO common names are: English, littlehead porgy;

French, daubenet titete; Spanish, plumajoroba (Randall and Vergara

R. 1978). Other names or variations appearing in the literature

include little-head porgy (Jordan and Fesler 1893; Beebe and TeeVan 1928), pez de pluma (Jordan and Fesler 1893; Beebe and TeeVan 1928; Duarte-Bello 1959), bajonado (Campeche Bank)

(Sokolova 1965), crocro and tete-feuilles (Trinidad) (Zaneveld 1983)

3

and bajonado azul (Campeche Bank) (Juarez 1975). The names bank

porgy, Key West porgy, grass porgy, and white snapper are

sometimes applied indiscriminately to C. proridens, C. arctifrons,

and C. calamus from the snapper grounds off the west coast of

Florida (Moe 1963).

1.3 Morphology

1.31 External morphology

Calamus arctifrons-The following morphological description is

adapted from Evermann and Marsh (1902) and Randall and Vergara

R. (1978): Head length 3.3 in SL; body oval, compressed and deep,

its depth 2.0-2.5 in SL; eye width 3.4 in SL; snout length 1.9 in

SL; maxillary length 2.4 in SL; interorbital distance 3.5 in SL;

preorbital distance 2.8 in SL; suborbital space rather deep, 6.6-7.8

in SL; upper head profile usually smoothly convex and moderately

steep; jaws anteriorly with about 8-10 canine-like teeth of about

equal size; jaws with two rows ofmolar-like teeth laterally in lower

jaw and three rows in upper jaw, without an irregular series inside

and toward the front; dorsal fin XII, 12; anal fin III, 10; pectoral

fins relatively short, not reaching to anal fm origin when appressed,

usually with 16 rays; lateral-line scales 43-49; about five rows of

scales on cheek; back not greatly elevated; anterior profJ1e not steep,

a slight angle in front of eye.

Additional meristic data are presented in Table 1. The skull of

Calamus sp. was discussed by Jordan and Gilbert (1882), and

illustrated by Gregory (1933).

Table I.-Meristic data for Calamus areti/rons. Numbers in parentheses are

percentages of the total number ofspecimens examined. (From Randall and

Caldwell 1966)

Character Count (%)

Lateral-line scales 43 44 45 46 47 48 49

(1.2) (8.6) (9.9) (32.1) (29.6) (9.9) (8.6)

Pectoral fin rays IS-IS 15-16 16-16

(11.5) (20.5) (67.9)

Dorsal fin spines XI XII XIII

Dorsal fin rays 12 13 II 12 13 II 12

(1.3) (0) (5.2) (85.7) (1.3) (6.5) (0)

Anal fin spines III

Anal fin rays 9 10 II

(0) (97.4) (2.6)

Gill rakers IO II 12

in first arch (63.8) (27.5) (8.7)

Color of adults in preservative: Light brown; a black spot smaller

than the eye, but larger than the pupil near the front of the lateral

line; five vertical rows of dark brown blotches on body and two

diagonally elongate blotches on caudal peduncle, the last continuing as the upper part of a large V-shaped marking basally on caudal

fin; two diffuse dark blotches mid-dorsally on nape anterior to first

row of spots on body; occasional specimens have irregular horizontal lines on the unscaled portions of cheeks; dorsal fins with

some faint dark blotches in line with rows of blotches on side of

body; lobes of caudal fin traversed by about five blackish bands;

pelvic fins with two faint transverse dark blotches (Randall and

Caldwell 1966).

Color ofjuveniles in preservative: Darker rows of blotches than

in adults, and a dark bar on head running through the eye, its front
Table 2.-Meristic data for Calamus proridens. Numbers in parentheses are

percentages of the total number ofspecimens examined. (From Randall and

Caldwell 1966)

Color in life: Iridescent silvery, with bright bluish tinges on back

and upper sides; diffuse horizontal elongate blue blotch at upper

end of gill opening; blue streak running along lower eye margin

and alternating blue (narrow) and yellow (wide) horizontal lines

across unscaled portion of cheeks; unmarked portion of suborbital

and snout brassy; lips yellowish; inside corners of mouth yellow

(Randall and Caldwell 1966; Randall and Vergara R. 1978).

See 1.21.

edge touching opening of the mouth; a few dark streaks on opercle

and scattered brown dots on body among the dark blotches forming the dark bars; two dark spots dorsally on caudal peduncle; a

row of dark spots on dorsal fin with a second row in spinous portion above and a second row in soft portion below; two rows of

dark spots on anal fin; dark bars on caudal lobes very conspicuous;

pelvic fins with two broad, dark crossbars (Randall and Caldwell

1966).

Description of colors in life: Silvery, olivaceous or bluish above,

the centers of many of the scales pearly and iridescent; paler below;

back and sides with 6-8 dark vertical crossbars, narrower than interspaces, with spots between them; conspicuous black blotch on lateral

line near upper end of gill opening; row of about six salmon-olive

spots along lateral line; snout olive, mottled with bluish and may

be streaked with yellow; interorbital area may have a yellow band;

broad, dark bar from eye straight downward across cheek; an

obscure pearly-blue streak below eye, and two or three similar ones

before eye; preorbital usually bluish, with more or less numerous

longitudinal streaks and dashes of golden yellow; preorbital sometimes pale salmon-yellow, with a few light bluish streaks; cheeks,

preopercle, and opercle pearly, with yellow shades and spots; opercular membrane coppery orange; pectoral fins pale; pelvic fins

bluish white. faintly barred; dorsal fin bluish, with blackish spots

along base of dorsal; anal fin bluish; caudal fin bluish, with small,

dusky salmon spots forming undulating crossbars, posterior margin

slightly dusky (Goode and Bean 1882; Jordan and Gilbert 1884;

Evermann and Marsh 1902; Randall and Vergara R. 1978).

See 1.21.

Character

Lateral-line scales

Pectoral fin rays

Dorsal fin spines

Dorsal fin rays

Anal fin spines

Anal fin rays

Gill rakers

in first arch

Count (%)

52 53 54 55 56 57

(2.7) (5.4) (29.7) (378) (18.9) (5.4)

13-14 13-15 14-14 14-15

(2.7) (0) (83.8) (13.5)

Xl! XIII ----

II 12 11 12

(2.8) (94.4) (2.8) (0)

1I1

10 11

191.9) (8.1)

11 12 13

(75.9) (20.7) (3.4)

Calamus proridens-The following morphological description is

adapted from Randall and Caldwell (1966) and Randall and Vergara R. (1978): Body oblong, compressed, very deep anteriorly

(1.95-2.20 in SL, relatively less in adults than in juveniles); snout

blunt and very steep in adults (less so in juveniles); dorsal profile

of head usually smoothly convex; nape strongly convex, developing into a distinct bump in large specimens; suborbital space rather

deep, about 6.3-9.9 in SL; mouth small (especially in small specimens), the maxilla not reaching to anterior eye margin; eye small;

both jaws anteriorly with canine-like teeth, the fourth from midline on each side enlarged and noticeably outcurved in specimens

>18 cm SL; laterally with molar-like teeth in three rows plus an

irregular series inside and toward the front in upper jaw; dorsal

fin rays XII or XIII (usually XII), 11 or 12 (usually 12); anal fin

rays III. 10 or 11 (usually 10); pectoral fin rays 13-15 (usually 14),

the pectoral fins extending to or beyond anal fin origin when

appressed; lateral-line scales 52-57 (usually 54-56); total gill rakers

in first gill arch 11-13 (usually II).

Additional meristic data are presented in Table 2. The skull of

Calamus sp. was discussed by Jordan and Gilbert (1882), and

illustrated by Gregory (1933).

Color in preservative: Tan to light brown; center of each scale

on upper half of body with a dark brown spot; horizontally elongate

dark brown blotch, broader posteriorly, running over upper end

of gill opening; front end of blotch beginning at or slightly anterior

to dorsal end of free margin of preopercle; supra and suborbital

dark brown lines; three dark brown lines extending anteriorly from

region in front of eye between supra and suborbital lines; unscaled

suborbital region with well-defined dark brown horizontal or slightly

diagonal lines on a bronze background, usually slightly wavy and

sometimes broken into spots or lines: caudal fin about same color

as body; other fins paler, dorsal with some faint brownish blotches

(Randall and Caldwell 1966).

4

2 DISTRIBUTION

2.1 Total area

Calamus arctifrons-Calamus arctifrons occurs in the northeastern

Gulf of Mexico, primarily in shallow water along the west coast

of Florida, but straying on occasion to Louisiana and Texas (Fig.

3) (Randall and Caldwell 1966; Hoese and Moore 1977; Randall

and Vergara R. 1978). The species is apparently absent from the

northwestern Gulf of Mexico (Briggs 1958; Randall and Caldwell

1966), and Campeche Bank. Although there are several published

reports of C. arctifrons from the West Indies (Evermann and Marsh

1902, Puerto Rico; Beebe and Tee-Van 1928, Haiti), Randall and

Caldwell (1966) were not able to find any West Indian specimens

in museum collections; if the species does occur in the Greater

Antilles, it is apparently quite uncommon there. Briggs (1958)

placed the southern limit of the range at I1ha Grande, Brazil, which

is almost certainly incorrect.

Calamus proridens-Calamus proridens occurs along the Atlantic

and Gulf of Mexico coasts of Florida to about Louisiana, on

Campeche Bank, and through the Greater Antilles (Fig. 4) (Briggs

1958; Randall and Caldwell 1966; Dubovitsky 1977a; Randall and

Vergara R. 1978). This species, like C. arctijrons, is apparently

absent from the northwestern Gulf of Mexico (Briggs 1958; Walls

1975). Struhsaker (1969) reported C. proridens as rare, but present, on live bottom in the South Atlantic Bight. Although several

authors (Jordan and Gilbert 1884; Jordan and Fesler 1893; Beebe

and Tee-Van 1928) reported specimens from the West Indies, Randall and Caldwell (1966) speculated that at least some of these

reports may not have been based on natural occurrences, the fish

having been caught at some other location. Calamus proridens is

fished in Cuba (Randall and Vergara R. 1978), and is known to

occur in Hispaniola.
N

50° r---------~Il"""'~~~ll""'"~--.,..--""I

N

50° r---------~:r_~"'T"'":::-_:_--.,...--.,

NORTH AMERICA

ATLANT/€ OCEAN

50 40

AHA TIC CEAN

100 90 80 70 60

o

20

NORTH AMERICA

30

10

40

90 80 70 60 50 40

;.":

100

o

10

20

30

40

Figure 3.-Distribution of Calamus arctifrons. (Based on Randall and Vergara R.

1978)

Figure 4.-Distribution of Calamus proridens. (Based on RandaU and Vergara R.

1978)

2.2 Differential distribution

2.21 Spawn, larvae and juveniles

Calamus aretijrons-Spawning areas of C. arctifrons are unknown.

Unidentified sparid larvae collected by Houde et al. (1979) in the

northeastern Gulf of Mexico probably included C. arctifrons. Most

(>90 %) of the unidentified sparid larvae were collected at stations

<50 m deep, and were most common in winter and spring.

Juveniles are often found in shallow seagrass beds. Most of the

specimens examined by Tabb and Manning (1961) from Florida

Bay Thalassia beds were juveniles of 7-10 cm SL.

See 2.3 and 3.16.

Calamus proridens-Spawning areas of C. proridens along the coast

of the United States are not we!! known. Unidentified sparid larvae collected by Houde et al. (1979) in the northeastern Gulf of

Mexico probably included C. proridens (see above).

On Campeche Bank, Dubovitsky (I977a) reported a steady drift

of pelagic eggs and larvae westward across the bank, eventually

reaching shallow water where the demersal habit is adopted.

Juveniles are found in relatively shallow water. On Campeche

Bank, young stay in shallow, productive waters for the first year

of life (Dubovitsky 1977a). Between ages I and II, there is a gradual

movement offshore toward the northeast and east, where smalJ fish

become recruited to the commercial stock.

See 2.3,3.16, and 3.51.

2.22 Adults

Calamus aretijrons-Adult C. arctifrons occur on shallow seagrass

beds and other vegetated bottom (Evermann and Marsh 1902; Tabb

and Manning 1961; Vick 1964; Randall and CaldwelJ 1966; Walls

1975; Hoese and Moore 1977; Randall and Vergara R. 1978), and

over hard or rock bottoms (Randall and Caldwell 1966; Darcy and

Gutherz3). They occur from near shore to at least 29.6 m (Darcy

and Gutherz3), and are demersal.

See 2.1, 2.3, and 3.51.

Calamus proridens-Adult C. proridens occur in shallow water,

from nearshore to at least 59.3 m on the West Florida Shelf (Darcy

and Gutherz unpubl.), and at least 85 m on Campeche Bank (Dubovitsky 1977a). They are found on or near natural and artificial reefs

(Smith et al. 1979; Miller and Richards 1980), around offshore platforms (Hastings et al. 1976), and on live bottoms of sponges and

corals (Struhsaker 1969; Darcy and Gutherz unpubl.). Smith (1976)

termed C. proridens a secondary reef fish.

See 2.1, 2.3, and 3.51.

2.3 Determinants of distribution changes

Calamus aretijrons-Distribution of C. arctifrons is affected by

temperature, salinity, water clarity, water depth, and habitat. In

the northern part of its range, C. arctifrons appears to be uncommon or absent in shallow water in the coldest months. At Crystal

River, Florida, Reid (1954) collected C. arctifrons from June

through December (except for August); Grimes (1971) collected

specimens March through December 1969, with greatest abundance

in September and July; and Grimes and Mountain (1971) collected

specimens April through December 1970, with greatest abundance

in October and December. In the southern part ofits range, C. arctifrons may remain in shallow water in winter; Tabb and Manning

(1961) found them most common October to April in Florida Bay.

The species occurs in water of at least 7.8-32.2 °C (Crystal River,

Reid 1954), and at salinities of at least 9.7-35.7°/00 (Crystal River,

Reid 1954; Charlotte Harbor, Wang and Raney 1971). Calamus

arctifrons prefers clear water (Tabb and Manning 1961). Turbidity avoidance may be a reason why this species is absent from the

northwestern Gulf of Mexico (Randall and Caldwell 1966).

Calamus arctifrons is stricted to fairly shallow water, perhaps

because benthic vegetation is confined to shallow water, and C.

arctifrons is often associated with vegetation (Evermann and Marsh

1902; Tabb and Manning 1961; Walls 1975; Randall and Vergara

R. 1978). The species has been reported from at least as deep as

29.6 m (Darcy and Gutherz unpubl.), but is usually most common

in shallower water. At Crystal River, Reid (1954) found C. arctifrons on shallow and deep flats and in channels 4.8-5.5 deep. On

5
l..-.,...--""T"""-.....,..----,r--......+--......-.....,.......... 18

Figure 5.-Distribution of Calamus proridens and catch rates on Campeche Bank;

A. January 1970; B. February-March 1971. (From Dubovitsky 1977a, fig. 2)

N

t

88 86

00

• <100

• 100-300

• 300-S00

90

,

\

,

,

,- - 24N "- B t' \ )

jJ ") N /-0 0 1

r

/ o ._ , / ,.-0--,•• __• __ ,-' .-'" --' ....... ' .,~

" --:"" -",~ " • • • 0.0 22 (C • • • Q

• •••• 0 • .,a , • • 0')

,7

,7

'J

'" , (/

" 20

YUCATAN

W 92

A

shallow «30-35 m) water on the central bank in December, small

fish were present in January, and both size groups were present

in March, On the offshore part of the bank, at 35-55 m, small (18-20

em, type of length measurement unspecified) fish dominated from

December to March, their numbers increasing with depth. Juveniles

of 13-16 em appeared in offshore waters of the western and central bank in January, and on the eastern bank in February. From

May through August, size distributions were similar to those of

January. In November, small fish dominated everywhere except

on the shallow central bank, where mixed size groups occurred,

and on the northwestern bank. There was a notable increase in small,

immature fish on the southwestern bank. In general, throughout

the year, large fish were associated with low bottom temperatures

and were most common on the eastern bank where cold-water

upwelling occurred. Large fish penetrated the western bank only

when bottom temperatures there declined. Small fish were usually

associated with higher bottom temperatures and were most common on the western bank and in shallow water, extending eastward

only in times of high bottom temperatures.

Calamus proridens-Distribution of C. proridens is affected by

salinity, temperature, water depth, growth stage of the fish, and

habitat. Although there are no specific references in the literature

to salinities at which C. proridens occurs, it is not found in estuaries

or low salinity area, and probably requires a salinity of at least

30%0.

Distributional changes with depth and season are probably

attributable to temperature changes. Little is known about the

distribution of C. proridens on the Atlantic coast of the United

States. In the Gulf of Mexico, the species occurs from at least II

to 59 m (Darcy and Gutherz unpubl.), with greatest abundance in

19-27 m (in January) and a mean depth occurrence of27.8 m. Abundance was found to vary significantly with depth. Smith (1976)

reported C. proridens from 29-42 m depths on the Florida Middle

Ground.

On Campeche Bank, considerable study has been made ofseasonal

distribution changes of C. proridens. Although Olaechea and

Sauskan (1974) did not feel that water temperature influenced abundance of Calamus spp. on Campeche Bank, most other authors

(Gonzales 1973; Olaechea 1975; Dubovitsky 1977a) did report relationships between bottom temperature and C. proridens abundance.

Gonzales (1973) found good correlation between catch rates of

Calamus spp. and water temperature on the eastern edge ofthe bank,

with fish moving deeper on the slope when shallow waters warmed.

Olaechea and Sauskan (l974) estimated highest abundance of C.

proridens on the bank in spring and summer. Sauskan and Olaechea

(974) reported this species most common in shallow (15-30 m)

water on the central part of the bank, but also common in shallow

and deep (50-120 m) water on the eastern bank, and at moderate

(30-50 m) depths on the western and central bank (see section 4.2).

Dubovitsky (l977a) presented detailed information on seasonal

changes in geographical distribution of C. proridens on Campeche

Bank. In January (Fig. 5A), the bulk of the population was on the

central bank. When bottom temperatures on the bank did not exceed 22°C, C. proridens extended its distribution into deep (65-70

m) water and as far south as lat. 20oN. In February and March

(Fig. 5B), C. proridens was most common in coastal areas of the

eastern bank at depths to 25 m, but extended to 40 m where cold

water flowed onto the edge of the shelf. In May through June (Fig.

6), C. proridens was restricted to shallow water, with commercially

exploitable aggregations present. By August (Fig. 7A), fish spread

out and moved to somewhat deeper (60-85 m) water; they were

most abundant east of long. 89°30'W, and below 30 m depth.

Bottom waters from May to August are the coldest of the year on

the bank, due to upwelling. In November. bottom temperatures are

at their maximum for the year, and fish were found in fairly shallow

water (Fig. 7B), with large aggregations only in the eastern and

western parts of the bank. In December (Fig. 7B), a similar pattern was seen, with fish well spread out across the bank. In general,

aggregations formed in periods of lowest bottom temperatures.

Seasonal changes in size composition of C. proridens also occur

on Campeche Bank (Dubovitsky 1977a). From December to March,

large individuals were always present on the eastern bank where

bottom waters were cold, though the numbers of large individuals

decreased from December to March. Large fish were present in

the West Florida Shelf, Darcy and Gutherz (unpubl.) reported C.

arctifrons most common in 9-18 m of water, with a mean depth

of 15.8 m, in the surveyed area. In addition to vegetated bottom,

C. arctifrons is common on hard bottom (Randall and Caldwell

1966) and rock bottom (Darcy and Gutherz unpubl.).

See 2.1, 2.21, 2.22, and 3.51.

6
22

20

N

f

N

I

88 86

YUCATAN

90

.--------......,;'..,-~-_...,-~----..,..24 N

B

/-{ I

'" 0 /' -" .,.~-9..... _._-.•-.---\, _ ......... ~.. O"'l::

- ...... ,_ ... I'" .........Cl -, 0' .--.. ••• -_~

• • •• • 00..\ \

,'..

... ~7

• 3

., .}

o 0 o 0

CO

W 92

,

,

,

22

20

N

I

N

r

88 86

o 0

• <100

• 100-300

• 300-500

YUCATAN

90

o •••

,

,

,

.'

,

\

0' 0 • 00

.--------/::"C'--a..';;'}~...----T24 N

B J(\0.1 /_1 0 d -/ . ~"- ... \

'" _.----....-------. -'.\ - -- .-::..~ ..--_. :."\~

~--/_.j •• • Q. \', ' .. \

" :1 I I~

.;-

.;

W 92

A

L..-,....---r----.,......-..,....-.I...I'--~-.....,...--l18

Figure 6.-Distribution of Calamus pToridens and catch rates on Campeche Bank;

A. May 1969; B. June 1969. (From Dubovitsky 1977a, fig. 2)

Figure 7.-Distribution of Calamus proridens and catch rates on Campeche Bank;

A. August 1969 (circles) and August 1971 (triangles); B. November 1970 (circles)

and December 1970 (triangles). (From Dubovitsky 1977a, fig. 2)

Calamus proridens is found primarily on sponge and coral bottoms

(live bottom) (Struhsaker 1969; Darcy and Gutherz unpub1.). Smith

et a1. (1979) reported C. proridens from natural and artificial reefs

off Clearwater, Florida, and Smith (1976) listed it as a secondary

reef fish of the northeastern Gulf of Mexico. The species also inhabits offshore platforms in the northern Gulf (Hastings et aI. 1976).

Hard substrates and sessile invertebrate growths probably provide

shelter and harbor food organisms of C. proridens.

See 2.1,2.21,2.22,3.51,3.52,4.13, and 4.2

2.4 Hybridization

No hybrids of either C. arctifrons or C. proridens are known.

3 BIONOMICS AND LIFE HISTORY

3.1 Reproduction

3.11 Sexuality

Nothing is known of sexuality of C. arctifrons, though it is probably a hermaphroditic species like many other sparids. Calamus

proridens is known to be a protogynous hermaphrodite, with

individuals changing from females to males al about 24 cm (measurement unspecified) (Dubovitsky 1977b). Because of this sex

reversal, females are consistently smaller than males (Fig. 8).

See 3.12 and 3.15.

7
32 0

0 0

E 28 0 0 0

U 0 0 0

r:P ...., 0 Do 0 0 .t= a 0 0 ODD

OJ 24 0 a Do C a a a a (J) a a -l a 00 00 a a 0 a 20 a 00 00 a a a a

a 16

N 0 J F M A M J J A

Month

Figure 8.-Mean lengths of males (squares) and females (circles) of Calamus proridens from Campeche Bank. Type of length measurement was unspecified. (From

Dubovitsky 1977a, fig. 5)

3.12 Maturity

Maturation size and age of C. arctifrons are unknown. Alth.;mgh

Dubovitsky (1977a, b) did not state a size of maturation for C. proridens, he did identify females as small as about 17 cm (measurement unspecified), and males as small as about 23 cm. The sreat

majority of adult females collected by Dubovitsky (1977a) were

ripe and in prespawning condition, but never more than half of the

ovaries had eggs in ripening condition. Male maturation stages were

difficult to assess.

See 3.11 and 3.16.

3.13 Mating

ably included C. arctifrons, were most abundant in winter and

spring. Reid (1954) hypothesized early spring spawning at Crystal

River, based on small individuals caught in June and July.

Calamus proridens-Spawning season of C. proridens along the

coast ofthe United States is not known, but larval surveys by Houde

et al. (1979) in the eastern Gulf of Mexico indicated highest abundance ofsparid larvae, probably including C. proridens, in winter

and spring.

On Campeche Bank, several studies of C. proridens have indicated

a long spawning period. Juarez (1975) stated that the spawning

season lasts from summer into winter, with maximum spawning

intensity in winter. Dubovitsky (1977a) reported spawning from

December to August. with a peak from January to March. Based

on visual inspection and histological examination of ovaries, individuals are apparently partial spawners (Dubovitsky 1977a), and

may spawn more than oncc a year. No dense spawning aggregations have been reported, but localized areas of prespawning

(characterized by numerous large, transparent oocytes occupying,

visually, about 20% of the total ovary volume) C. proridens have

been identified (Fig. 9) (Dubovitsky 1977a). Large, ripe individuals

occurred from December to June, with greatest abundance in

December and February on the shaJJow eastern bank, and in January

on the western bank at depths >35 m. Small prespawning fish were

observed from January through August, with peaks in January,

March, and July. Ripe, small fish were observed in offshore areas

at depths >40-50 m, and only on the central bank. Sequential spawning of size groups was also reported by Dubovitsky (1977a), with

different spawning locations for different size groups. Over the

spawning season, a gradual diminution ofsize ofspawners occurred.

From late May to the beginning of June, average size was 22 cm

(measurement unspecified); in early June it was 20 cm; by late July,

18 cm; and by early August, 17 cm.

YUCATAN

Mating in C. arctifrons and C. proridens has not been described.

Dubovitsky (1 977a) stated that C. proridens apparently does not

form dense spawning aggregations.

3.14 Fertilization

Fertilization is probably external.

3.15 Gonads

Gonads of C. arctifrons have not been described. Dubovitsky

0977a, b) histologically examined gonads of C. proridens, and

found successive sex inversion from female to male. The size at

which the ovaries regress and testes develop is very variable, but

there is a pronounced sex inversion at about 23-24 cm (measurement unspecified). By the end of the spawning season (middle to

late summer), asymmetry of the ovaries is often observed, the left

ovary being larger.

See 3.12 and 3.16.

3.16 Spawning

A

B

W92 90

r

88

23N

22

21

Calamus arctijrons-Spawning season of C. arctifrons is not known

with certainty. However, unidentified sparid larvae collected in the

northeastern Gulf of Mexico by Houde et al. (1979), which prob8

Figure 9.-Locations and months of collection of Calamus proridens in pre-spawning condition on Campeche Bank; A. December to March; B. May to August.

(From Dubovitsky 1977a, fig. 6)
-j 400

E 200

i

....: .

• • •

3.3 Adult phase

3.31 Longevity

Calamus arctifrons reaches a maximum size of about 22 cm

(measurement unspecified) and is common to 20 cm (Randall and

Vergara R. 1978). Calamus proridens reaches 44 cm and is common to 37 cm (Randall and Vergara R. 1978). Life spans of these

species are unknown.

3.32 Hardiness

60 C

40

20

D F M A M J

Month

A SON

Calamus arctifrons is apparently subject to toxic effects of red tides,

because Springer and Woodburn (1960) found several dead individuals on the beach in Pinellas County, Florida, following an

outbreak.

See 2.3 and 3.35.

Figure 10.-Seasonal changes in reproductive activity of Calamus proridens and

related factors on Campeche Bank. A. Bottom temperatures, 1969-1971. B. Total

plankton biomass between the 20-m and 200-m isobaths for lat. 91°W (dotted

line) and lat. 87°20'W (solid line). C. Percentage of pre-spawning females captured. Plankton productivity figures are from Jr6mov (1967). (From Dubovitsky

1977a, fig. 7)

Spawning of C. proridens on Campeche Bank is temperaturedependent, beginning with the appearance of cold-water inflows

onto the eastern bank in December (Dubovitsky 1977a). Plankton

productivity is also temperature-dependent, with maximum productivity in January through August (Jromov 1967). High plankton

productivity at the time of spawning assures that larval stages will

have adequate food supply. The relationships between temperature,

plankton productivity, and spawning of C. proridens on Campeche

Bank are shown in Fig. 10.

See 2.21,2.23,3.51, and 4.13.

3.17 Spawn

Eggs of C. arctifrons are probably pelagic, since most sparids produce pelagic eggs with little or no pigment (Breder 1962). Eggs

of C. proridens are known to be pelagic (Dubovitsky 1977a).

3.33 Competitors

Competitors of C. arctifrons and C. proridens are not known, but

probably include other sparids, haemulids, lutjanids, labrids, and

other fishes that feed on benthic invertebrates. Associates of C. arctifrons and C. proridens on the West Florida Shelf are discussed

in section 4.6. Whether these associates are actually competitors

is not known.

3.34 Predators

Predators of C. arctifrons and C. proridens are unknown, but probably include sharks, sphyraenids, and large serranids and lutjanids.

3.35 Parasites, diseases, injuries, and abnormalities

Calamus arctifrons has been reported parasitized by intestinal trematodes, including Proctoeces lintoni, Lepocreadium opsanusi,

Pseudocreadium anandrum, and Pachycreadium crassigulum

(Nahhas and Cable 1964). Red tides apparently prove fatal to some

individuals of C. arctifrons (Springer and Woodburn 1960).

3.4 Nutrition and growth

3.2 Preadult phase 3.41 Feeding

3.21 Embryonic phase

Embryonic development of C. arctifrons and C. proridens have not

been studied. Developing eggs are probably pelagic.

3.22 Larvae and adolescent phase

Larvae of C. arctifrons and C. proridens have not been described

in the literature. Young of Calamus spp. are difficult to distinguish.

Houde et al. (1979) were unable to separate species of Calamus

from plankton collections in the eastern Gulf of Mexico. Calamus

proridens is known to have pelagic larvae (Dubovitsky 1977a); C.

arctifrons larvae are probably also pelagic.

9

Feeding of C. arctifrons is unknown, but dentition suggests benthic feeding habits. Sessile and motile invertebrates are probably

bitten or picked off hard substrates and seagrass blades, and other

prey picked from the surface of softer substrates.

Calamus proridens probably feeds in a manner similar to that

described for C. arctifrons. On Campeche Bank, feeding intensity

varies over the year (Fig. II), with major feeding concentrated

around the spawning and postspawning periods (Dubovitsky 1977a).

Primary feeding grounds on Campeche Bank are the inshore parts

of the plateau (Dubovitsky 1977a). At the outer edge of the bank,

intense feeding, as measured by mean index of stomach fullness,

was noted only in the period just prior to spawning.

See 3.42.
1.6

• 0

0

X t:,.

(1) 1.2 0

"C 0 c: • • 0

• 0 (JJ 0.8 t:,. (JJ t:,.

(1) 0 .~

c: t:,.0

t:,.t:,. .0 u.. 0.4 • t:,.

t:,.; 0 •

0 /Y) .... arI\I". A

N D J F M A M J A

Month

Figure II.-Feeding intensity ofsmall mature (circles) and large mature (triangles)

specimens of Calamus proridens by month on Campeche Bank. Solid symbols refer

to specimens collected in >30-35 m depths, open symbols refer to specimens collected in depths <30-35 m. (From Dubovitsky 1977a, fig. 9)

3.42 Food

Both C. arctifrons and C. proridens probably feed primarily on invertebrates. Beebe and Tee-Van (1928) reported unidentifiable

animal and vegetable materials from stomachs of C. arctifrons, including 1halassia leaves. Whether the leaves were consumed

deliberately or incidentally is not known. Calamus aretifrons at

Crystal River conatined copepods, amphipods, mysids, shrimp,

bivalves, and gastropod mollusks (Mitrella sp. and Bittium sp.) and

polychaetes (Reid 1954). Calamus proridens examined by Beebe

and Tee-Van (1928) contained small crustaceans. Dentition of C.

proridens suggests feeding on hard-shelled benthic crabs and bivalved mollusks (Dubovitsky 1977a). On the western part of

Campeche Bank, south of lat. 20oN, the main food of young C.

proridens is pink shrimp, Penaeus duorarum (Dubovitsky 1977a).

3.43 Growth rate

Growth rate of C. arctifrons is unknown. Growth rate of C. proridens is reportedly rapid in the first year of life (Dubovitsky 1977a),

though Dubovitsky did not publish growth curves. There is apparently substantial variation in timing of growth and spawning within

the population, and scale increment formation is variable, possibly

due to differences in spawning and feeding among fish of different

size groups (Dubovitsky I977a). Growth rate differs from female

to male phase of the fish; egg development reportedly slows growth

of individuals functioning as females (Dubovitsky I977b), ?fter

which the growth rate increases again. Dubovitsky based these conclusions on histological examinations of gonads and analysis of

length frequencies, but there is little data presented in his publications on which to critically evaluate them.

See 3.44.

10

2.0

•

X 1.6

(1)

"C •

c: •

1.2 • -c:

(1) 0 • • 0

- t:,.•• t:,. c:

()

0 0.8 0 • • t:,. 0

- t:,. a3

u.. .~ t:,.

0.4- t:,. • •

• • •

0 • t:,.~ t:,.

0 n 0

N D J F M A M A

Month

Figure 12.-Seasonal changes in fat content of mature Calamus proridens from

Campeche Bank. Circles represent small specimens, triangles represent largt

specimens. Solid symbols refer to specimens caught in depths >30-35 m, open

symbols refer to specimens caught at depths <30-35 m. (From Dubovitsky 1977a,

fig. 8)

3.44 Metabolism

When C. proridens individuals are functioning as females, growth

rate, as determined from length frequency analysis, slows due to

metabolic changes (Dubovitsky 1977b). When fish change from

female to male, growth rate increases. Fat content of the body cavity

increases in mature fish to a maximum in November and December,

just prior to spawning (Fig. 12) (Dubovitsky 1977a). Fat content

decreases through the spawning season and is low by the end of

spawning. Fat build-up is rapid in July and August in large postspawning fish. During spawning, fat content is highest in fish living

in deep water.

See 3.43.

3.5 Behavior

3.51 Migrations and local movements

No migrations of C. arctifrons have been reported.

Migrations of C. proridens on Campeche Bank have been reported

by Dubovitsky (1977a). Eggs and larvae move primarily from east

to west across the bank, and move towards shallow water (Fig. 13).

Small fish move gradually from west to east in December through

March. The majority of mature fish migrate into coastal waters of

the central bank in January, dispersing eastward by March. Migration patterns in mature fish are strongly developed during the spawning period, with spawners generally moving offshore. After spawning, fish move into shallower water again. An extensive migration

of 19-21 cm (length unspecified) fish to inshore waters occurs in

January. Aggregations of C. proridens on Campeche Bank tend to

form when bottom temperatures are lowest.

See 2.3 and 3.52.
'0 100

/' 24 N "- (' '- J \ 80 r-) I N /

__ -r<----·'--:..);;----- --;~;~\" I -- - ,--'. /, , i "r"f" ,- -:7---" 60 r ' -,' ""l 5 - 22 de! (,.--' .","'4' " ,t~ '~ o-a "\"1\\ ". • ; f Ii .-1.,/ .- - 0-<> QQ .,: ~:.~/ //. l " \~',\ / /' 40

d ~I

\ \ {tt YUCATAN 20 20

dl t/

--~ 1 ..,. 2 0

=t~ 14 18 22 26 30 34

18

W92 90 88 86 LENGTH (em)

Figure 13.-Schematic representation of migrations and movements of Calamus

proridens on Carnpeche Bank. Arrows indicate movements of: I, eggs and larvae; 2, immature individuals; 3, spawning individuals; 4, post-spawning individuals. (From Dubovitsky 1977a, fig. 10)

Figure 14.-0verall sex ratio of Calamus proridens in the commercially exploited

population on Campeche Bank. (From Dubovitsky 1977b, fig. 2)

4.13 Size composition

Table 3.-Percentage of females in three size

ranges of Calamus proridens in the sexually

mature part of the commercial catch on

Carnpeche Bank, 1969-71. (From Dubovitsky

I97Th)

where S is the proportion of females, and L is the mean length of

the females examined. It is obvious from Fig. 14, however, that

the true relationship approaches linearity only over a restricted range

of length.

See 4.13.

Size composition of C. arctifrons has not been reported.

Size composition of C. proridens on Campeche Bank was examined by Dubovitsky (1973, 1977a, b) (Figs. 15-20; Table 4).

Size distributions presented by Dubovitsky (l977a) were often

bimodal, with the smaller size-class dominating. In stocks of mature

fish, females 18-22 cm (measurement unspecified) predominated

(Fig. 20). About 59% ofal! females in the commercially exploited

population were between 18 and 21 cm in length, and 65 % of the

males were between 25 and 28 cm. Size composition of females

and their proportions in the commercia! population are presented

in Fig. 21.

See 2.3.

37.9

39.5

22.6

% examined

specimens

99.3

78.5

19.7

%

females

14-20

21-25

26-34

Size

range' (em)

4.11 Sex ratio

Although schooling behavior is not well known in C. aretifrons and

C. proridens, most small porgies form aggregations (Randall and

Vergara R. 1978). Large individuals are more often solitary or found

in small groups. Dubovitsky (l977a) noted concentrations of C.

proridens on Campeche Bank, but stated that they did not form dense

spawning aggregations.

See 3.51.

4 POPULATION

Sex ratio of C. arctifrons is unknown.

On Campeche Bank, Dubovitsky (1977b) found that female C.

proridens always predominated, regardless of time or location. Sex

ratio observed was not constant from year to year, but averaged

1:2.7 males to females. About 73 % of the commercially exploited

population were females. Sex ratio changed drastically with size

of the fish examined (Fig. 14). Only about 1% of specimens <23

cm (measurement unspecified) were males (Table 3). Beyond 24

cm, the percentage of males increased sharply, to about 69% of

the total. Only females were found in the smallest specimens, and

only males in the largest. Dubovitsky (l977b) derived the following linear relationship between sex ratio and size of fish:

3.52 Schooling

4.1 Structure

S = 2.338 - 0.072 L

'Type of length measurement unspecified.

11
,. 15

10

5

O~""'~ - - __- ....._~p;iiiI...

14 18 22 26 30 34

LENGTH (em)

Figure 15.-Size composition of Calamus proridells on Campeche Bank (II = 665,

:r =22.73 em). Type of length measurement was unspecified. (From Dubovitsky

1973, fig. 3)

% 30

JAN

20

10

DEC

.. ....--- ....t~

<";.;-~..... , ,''-~

20

10

20

JAN

10

JUN

20

10

FEB-MAR

50

40

30

20

10

JUL

::~ 18 22 26 30

NOV

20 I'

I ' I \

10 I \

I

.f

18 22 26

NOV

30

LENGTH (em)

Figure 16.-Size composition of Calamus proridells on eastern Campeche Bank,

by month. Solid lines represent specimens collected at depths <30-35 m, dashed

lines represent specimens collected at >30-35 m. Cross-hatching represents

specimens with ovaries in pre-spawning condition. Type of length measurement

was unspecified. (From Dubovitsky 1977a, fig. 3)

12

LENGTH (em)

Figure 17.-Size composition of Calnmus proridells on central Campeche Bank,

by month. Solid lines represent specimens collected at depths <30-35 m,

dashed lines represent specimens collected at >30-35 m. Cross-hatching

represents specimens with ovaries in pre-spawning condition. Type of length

measurement was unspecified. (From Dubovitsky 1977a, fig. 3)
20

60

80

40

34

Females

Total

30

0-0 Males

18 22 26

LENGTH (em)

14

10

TOTAL

NUMBER

100

% 20

Figure 20.-Size distribution of male and female Calamus proridens in the

Campeche Bank commercial population with a histogram of lengths. (From

Dubovitsky 1977b, fig. 1)

DEC

JUL

NOV

10

10

10

20

20

% 20

18 22 26 30

LENGTH (em)

28 30

o DEC -MAR

OJUN-AUG

" NOV

o

o

26

o

22 24

LENGTH (em)

20

o

18

\

SO __.,.....__........__,..\..,....'1""'0..,.....,.....;__........"'I""""'l

70

80

60

". 90

DEC

20

1

10

I.---"""I"'=,....-,...-......,...-""T"""-~--r--\

Figure IS.-Size composition of Calamus proridens on northwestern Campeche

Bank, by month. Solid lines represent specimens collected at depths <30-35 m,

dashed lines represent specimens collected at >3()"35 m. Cross-hatching represents

specimens with ovaries in pre-spawning condition. Type of length measurement

was unspecified. (From Dubovitsky 1977a, fig. 4)

30

20

10

18 22 26

JUL

30

Figure 21.-Size distribution of female Calamus proridens and their frequency

in the sexually mature part of the Campeche Bank population, 1969-1971. Type

of length measurement was unspecified. (From Dubovitsky 1977b, fig. 3)

LENGTH (em)

Figure 19.-Size composition of Calamus proridens on western Campeche Bank,

by month. Solid lines represent specimens collected at depths <30-35 m, dashed

lines represent specimens collected at >30-35 m. Cross-hatching represents

specimens with ovaries in pre-spawning condition. Type of length measurement

was unspecified. (From Dubovitsky 1977a, fig. 3)

13
Table 4.-Size composition of Calamus proridens on Campeche Bank.

Numbers above horizontal lines refer to specimens caught at depths ,,35 m;

figures below lines refer to specimens caught at depths >35 m. (From Dubovitsky 1977a, Table 2)

ReI. freq. Percent length Size Modal of modal Mean

range' group group length distribution in sample

Months (em) (em) (%) (em) <20 em 20-25 em >25 em

Eastern Area

Dec. 18-34 21-24 30 25.9 "7 39 54

27-31 40

Jan. 18-32 22-23 24 25.0 "3 57 40

29-31 24

Feb.-Mar. 13-32 20-23 58 22.9 "8 70 22

Aug. 20-22 21

16-31 24-25 22 23.7 11 57 32

Nov. 18-30 19-22 55 220 25 54 21

Central Area

Jan. 15-29 19-22 51 21.2 28 58 14

16-26 18-19 51 18.6 67 30

Mar. 14-34 22.9 27 41 32

15-29 19-20 43 20.8 42 47 II

June 20-21

16-29 25 14 22.3 9 79 12

18-20 47

July 16-30 24-26 29 21.5 37 53 10

16-26 18 48 19.0 76 23 I

Nov. 15-33 23.6 18 47 :'5

16-30 18-20 58 20.6 47 44 9

Northwest Area

22-24 42

Dec. 19-29 26-27 26 24.6 .J. 59 40

17-27 19-21 48 21.6 26 64 10

24-26 27

Jan. 13-30 21-23 45 22.4 15 65 20

July 15-27 20-22 55 21.6 15 77 8

21-24 56

Nov. 19-30 26-28 32 24.3 .J. 63 36

15-30 18-21 56 20.8 40 51 9

Southwest Area

Dec. 17-29 19-21 56 20.7 34 63

Jan. 15-34 22-23 35 24.5 66 31

July 17-26 20-21 63 21.0 II 88 .1

14-26 16-19 72 18.2 78 21 1

'Type of length measurement unspecified.

; 1.5 ill z :I:: 1.0 C ;:) Cl Z

z 0.5

11.;:)

: '-r,..JD......J'-r,"'==...,.,--~-...,.---- II 0..rn8

, '~7, 28-SS,'6-"., m-~ •••" 1

m

)

4.14 Subpopulations

No subpopulations of C. arctifrons or C. proridens are known.

4.2 Abundance and density

Calamus arctifrons-Calamus arctifrons was reported very abundant around the Florida Keys by Jordan and Fesler (1893). Tabb

and Manning (1961) found C. arctifrons abundant on 7halassia flats

in Florida Bay; they were most common from October to April.

A trawling survey of the West Florida Shelf (Darcy and Gutherz

1984) collected C. arctifrons at 7.7 % of the stations sampled; this

species ranked tenth in biomass of all demersal fish sampled in 9-93

m in the survey area, and accounted for 3.1 % of the total demersal

fish biomass collected in that region. North of Tampa Bay, C. arctifrons ranked eighth in biomass in 9-35 m, and accounted for 5.6%

of the total biomass. Abundance was highest in shallow (9-18 m)

water (Fig. 22) and on rock bottom (Darcy and Gutherz unpubl.).

A plot of mean abundance by grid zone on the West Florida Shelf

(Fig. 23) shows maximum abundance in nearshore waters off the

northwest coast of Florida. Estimated standing stock of C. proridens

on the West Florida Shelf at depths >9 m is about 28,000 mt (Darcy

and Gutherz unpubl.). In shallower water in the eastern Gulf of

Mexico, C. aretifrons was reported common by Reid (1954), Grimes

(1971), Moe and Martin (1965), Tabb and Manning (1961), and

Grimes and Mountain (1971). The species is not common in the

West Indies (Beebe and Tee-Van 1928; Randall and Caldwell 1966).

See 2.1 and 2.3.

Calamus proridens-Calamus proridens abundance on the Atlantic coast ofthe United States is poorly known, but Struhsaker (1969)

reported it rather rare in the South Atlantic Bight.

In the Gulf of Mexico, C. proridens is not very common in the

extreme northern gulf (Walls 1975), but is quite common on the

northern central West Florida Shelf (Darcy and Gutherz unpubl.)

(Fig. 24). Smith (1976) reported that C. proridens is the most common sparid at depths >30 m in the eastern Gulf of Mexico, and

Smith et al. (1975) reported it as occasional on the Florida Middle

Ground. Calamus proridens occurred at 9.2 % of the stations sampled on the West Florida Shelf by Darcy and Gutherz (1984), and

ranked eighth in biomass (3.7 % of total demersal fish biomass) in

9-93 m over the entire survey area. It ranked seventh in biomass

(6.5 % of total demersal fish biomass) north of Tampa Bay in 9-35

m depths. Abundance was greatest in moderate (19-55 m) depths

(Fig. 25) (Darcy and Gutherz unpubl.); estimated standing stock

on the West Florida Shelf was about 34,000 mt.

On Campeche Bank, C. proridens is the most abundant sparid

(Dubovitsky 1973, 1977b); along with C. nodosus and C. bajonado,

C. proridens makes up the bulk of the commercial trawl fish catch

on Campeche Bank. It has been estimated to comprise 10% of the

benthic fish fauna of the bank (Sauskan and Olaechea 1974), and

is second only to the tomtate, Haemulon aurolineatum, in abundance (Olaechea and Sauskan 1974; Sauskan and Olaechea 1974).

According to Dubovitsky (1977a), C. proridens is the most important commercial fish on the bank. Catch rates of >500 kg/h trawling have been reported (Dubovitsky 1977a), but rates of<100 kg/h

are more usual (Sauskan and Olaechea 1974) (Table 5). Estimated

density on Campeche Bank by season, expressed in kilograms per

hectare, was:

Figure 22.-Index of relative abundance of Calamus arclifrons on the West Florida

Shelf, January 1978, expressed in terms of biomass and number of individuals.

(From Darcy and Gutherz, unpubl., fig. 12.)

14

Winter

1.79

Spring

5.50

Summer

5.65

Fall

1.74

X

3.74
r

26, +

ITIIJ 0.1-1.0 \ 1.1-10.0

10.1-50.0

50.1-100 "

• :> 100

24'

84' 82' 86'

r

26' ) +

[ll) •.1-1••

1.1-10.0

10.1-50.0

050.1- 100

24'''1----------+-:----------+-:-------'

860 84° 82°

Figure 23.-Mean catch rate of Calamus areti/rons by 20' x 20' grid zones on

the West Florida Shelf, January 1978. Catches are expressed in mean numbers

of individuals caught per to-min tow. (From Darcy and Gutherz, unpubl., fig. 13)

Figure 24.-Mean catch rate of Calamus proridens by 20' x 20' grid zones on

the West Florida Shelf, January 1978. Catches are expressed in mean numbers

of individuals caught per 10-min tow. (From Darcy and Gutherz, unpubl., fig. 15)

Table S.-Mean catch rates of Calamus proridens per hour

of trawling of SRT-M trawlers on Campeche Bank, and

percentage of the total fish catch in each part of the bank

accounted for by C. proridens. (From Sauskan and

Olaechea 1974.)

Mean % of Mean % of Mean % of

Depth catch total catch total catch total

(m) (kg/h) fish (kg/h) fish (kg/h) fish

15-30 45.5 12 95.0 14 69.9 6

30-50 53.5 15 54.0 8 17.1 6

50-120 0.0 0 13.9 5 60.9 14

w ll: 1.0 (,) w Z lD C J: 0.5 C ;:)

Z Z ... ;:)

OlDC 9-18 19-2728-5556-110111-193 Depth (m) ><ww III c> III z-

-~ c

J: ...

w 0

ll: iii 0.5

Figure 2S.-lndex of relative abundance of Calamus proridens on the West Florida

Shelf, January 1978, expressed in terms of biomass and numbers of individuals.

(From Darcy and Gutherz, unpubl., fig. 14)

Central East

Area of Campeche Bank

West

15
Table 6.-Estimated standing stock of porgies on Campeche

Bank, based on Cuban trawl surveys. Figures for 1974-75 represent two different estimates. (From Klima 1976)

Dates of surveys

1964-72 1974-75 1974-75 Mean

Standing stock (mt) 209,000 182,000 461,000 325,000

(Olaechea and Sauskan 1974). Sauskan and Olaechea (1974) estimated total mean abundance of C. proridens on Campeche Bank

at 88,000 mt. Estimates ofstanding stock of all porgies on Campeche

Bank are presented in Table 6.

Calamus proridens does not seem to be very common in the West

Indies (Randall and Caldwell 1966), though Beebe and Tee-Van

(1928) reported them quite common in Haitian fish markets.

See 2.1, 2.21, 2.22, 2.3, 3.16, 3.51, and 4.13.

thopristis chrysoptera; toadfish, Opsanus spp., seahorses and

pipefishes, Syngnathidae; mojarras, Gerreidae; seatrouts, Cynoscion spp.; gobies, Gobiidae; triggerfishes and fLiefishes, Balistidae;

puffers, Tetraodontidae; and porcupinefishes, Diodontidae. In

deeper water (>9 m), cluster analysis of demersal fish catches on

the West Florida Shelf (Darcy and Gutherz unpubl.) grouped conger eels, Hildebrandia sp.; spottail pinfish; crested cusk-eel, Ophidion welshi; sand perch, Diplectrumformosum; scrawled cowfish,

Lactophrys quadricomis; black sea bass, Centropristis striata; and

shortnose batfish, Ogcocephalus nasutus, with C. arctifrons.

Calamus proridens is a member of live-bottom communities in

shallow to moderate depths. On the West Florida Shelf, cluster

analysis grouped tomtate; gulf flounder, Paralichthys albigutta;

Atlantic croaker, Micropogonias undulatus; and the cusk-eel,

Lepophidium brevibarbe, with C. proridens in water >9 m deep

(Darcy and Gutherz unpubl.). On Campeche Bank, C. proridens

is often caught with C. penna, C. nodosus, and C. bajonado,

especially in summer (Olaechea and Sauskan 1974).

See 3.3.

4.3 Natality and recruitment

4.31 Reproduction rates

5

5.1

EXPWITATION

Fishing equipment

Reproduction rates of C. arctifrons and C. proridens are unknown,

but C. proridens is a sequential spawner based on ovarian examination and histology (Dubovitsky 1977a) and may spawn more than

once each spawning period.

See 3.15 and 3.16.

Calamus arctifrons is caught in beach seines (Evermann and Marsh

1902; Randall and Caldwell 1966) and bottom trawls, and on hookand-line and bottom longlines (Randall and Vergara R. 1978).

Calamus proridens is caught in fish traps, trawls, and on hookand-line (Randall and Vergara R. 1978).

4.32 Factors affecting reproduction 5.2 Fishing areas

Although there are no reports of factors affecting reproduction in

C. arctifrons, temperature probably determines spawning season.

Spawning in C. proridens is known to be temperature-dependent

and to coincide with maximum plankton productivity (Dubovitsky

1977a).

See 3.16.

4.33 Recruitment

Little is known of recruitment in C. arctifrons. Wang and Raney

(1971) collected a small specimen (26 mm SL) in March in Charlotte

Harbor estuary, Florida.

Calamus proridens as small as 18 mm SL were collected on

shallow grassflats off Lower Matecumbe Key, Florida, in April.

and specimens as small as 21 mm SL were collected in March

(Springer and McErlean 1962). On Campeche Bank, C. proridens

are recruited to the commercial stock between ages I and II, and

at lengths of about 14 cm (measurement unspecified) (Dubovitsky

1977a, 1977b).

See 2.21, 2.3, and 3.51

4.6 The population in the community and the ecosystem

Calamus aretifrons is a common member of grassbed communities

in shallow water, and live-bottom communities in somewhat deeper

water. Associates of C. arctifrons in shallow water probably include: pinfish, Lagodon rhomboides; spottail pinfish, Diplodus

holbrooki; sheepshead, Archosargus probatocephalus; pigfish, Or16

Calamus arctifrons is fished in coastal waters of the Gulf of Mexico from southern Florida to Louisiana (Randall and Vergara R.

1978), but there is no directed fishery.

Calamus proridens is fished in coastal waters of Florida,

Campeche Bay, and Cuba (Randall and Vergara R. 1978). Concentrations of C. proridens on Campeche Bank in summer were

mapped by Olaechea and Sauskan (1974) (Fig. 26).

See 2.1,2.22,2.3. and 3.51.

N 230 N

I

22

21

YUCATAN 20

W 92° 90 88

Figure 26.-Principal zones of concentration of Calamus proridens on Campeche

Bank in summer. Calamus bajonado, C. petlna and C. nodosus are also concentrated in the same areas. (From Olaechea and Sauskan 1974, fig. 3)
5.3 Fishing seasons

No specific infonnation regarding fishing seasons is available. Since

there is no directed fishery for e. arctifrons or e. proridens in the

United States, catches of these species are probably dependent upon

effort expended in pursuit of other species. On Campeche Bank,

e. proridens are present year-round, but are most concentrated,

and therefore most easily fished, when bottom temperatures are low

(Dubovitsky 1977a).

See 2.3 and 4.13.

5.4 Fishing operations and results

5.43 Catches

Calamus aretifrons and e. proridens are good food fish and are

marketed mostly fresh and frozen (Evennann and Marsh 1902; Randall and Vergara R. 1978). Moe (1963) stated that these species

are of minor commercial importance, but are sometimes caught by

commercial snapper fishermen off the west coast of Florida, and

are often used as fresh bait. Calamus proridens is one of the most

important commercial species on Campeche Bank (Sokolova 1965),

and is not uncommon in markets in Haiti (Beebe and Tee-Van 1928).

Catch statistics for e. arctifrons and e. proridens are not reported

separately (Randall and Vergara R. 1978). Total reported catch of

Calamus spp. from FAO fishing area 31 (western central Atlantic

Ocean) in 1975 was about 6,777 mt, but actual landings were undoubtedly much higher, since landings of porgies are usually combined with landings of other percomorph fishes (Randall and Vergara

R. 1978). Catches of all species of porgies combined in FAO fishing

area 31 were: 1970, 1,400 mt; 1971,3,600 mt; 1972,8,900 mt;

1973,3,600 mt (Klima 1976). Catches of porgies by individual nations fishing in the western central Atlantic area in 1970-73 are

shown in Table 7. Most of the reported catch was probably from

Campeche Bank. Porgies, of which e. proridens is the most common, comprised 30% of trawl survey catches by the Cubans on

Campeche Bank in the early 1970's (Klima 1976). Based on these

and other surveys, Campeche Bank could yield 33,000-65,000 mt

of porgies each year (Klima 1976). Sauskan and Olaechea (1974)

estimated the potential catch of e. proridens on the bank as 15,000-

20,000 mt. Porgies made up 56-58% of Cuban trawl catches on

Campeche Bank in 1973-74 (Klima 1976). Commercial catches on

the bank have been quite variable, possibly due to changes in environmental conditions (Klima 1976).

Recreational catch figures, like commercial catch figures, are

seldom reported for separate species of porgies. The total recreational catch of porgies in the Gulf of Mexico in 1960 was estimated

at 8,550,000 fish, of 5,792,000 kg (Clark 1962). Most (72 %) were

caught from boats, and the rest from shore. Recreational catches

of porgies reported in 1970 were 13,234,000 fish, of 9,671,000

kg, in the eastern Gulf of Mexico, and 16,230,000 fish, of

10,913,000 kg, along the Atlantic coast of the United States south

of Cape Hatteras (Deuel 1973). Landings are stratified by fishing

area and method in Table 8.

See 2.3 and 4.2

17

Table 7.-Reported catcbes of porgies from

the western central Atlantic fishing area,

1970-73, by fishing nation. (From Klima

1976)

Catch per year (mt)

Nation 1970 1971 1972 1973

Cuba 400 2,000 1,900 1,900

U.S.S.R. 0 400 6,000 300

Mexico 500 600 600 800

Table S.-Recreational catches of porgies in Ihe eastern Gulf of Mexico and

along the soulhern Atlantic coast of Ihe United States, 1970. Figures are in

thousands of individuals caught. (Data from Deuel 1973)

Fishing area Fishing method

Sounds, Private, Party, Bridge,

Geographical rivers, rented charter pier, Beach,

area Ocean bays boat boat jetty bank

South Atlami,;; 6,266 9,964 4,989 1,068 7,499 2,674

Eastern Gulf 905 12,329 6,579 128 4,587 1,940

of Mexico

6 PROTECTION AND MANAGEMENT

6.1 Regulatory measures

Calamus arctifrons is included in the Snapper/Grouper Complex

Fishery Management Plan of the South Atlantic Fishery Management Council. Both e. arctifrons and e. proridens are included

in the Reef Fish Fishery Management Plan of the Gulf of Mexico

Fishery Management Council.

ACKNOWLEDGMENTS _

I thank Donald M. Allen and Albert C. Jones of the National Marine

Fisheries Service, Southeast Fisheries Center, for critically reviewing this manuscript.
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18

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