NOAA Technical Report NMFS 26

Synopsis of Biological Data

on the Sand Perch,

Diplectrum formosum

(Pisces: Serranidae)

George H. Darcy

March 1985

FAO Fisheries Synopsis No. 143

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 propriety 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.
CONTENTS

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

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

1.1 Nomenclature.................................................................................... 1

1.11 Valid name.. .. .. .. . .. .. . .. .. . .. . 1

1.12 Objective synonymy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.2 Taxonomy....................................................................................... 1

1.21 Affinities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.22 Taxonomic status. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.23 Subspecies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.24 Standard common names, verncular names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

1.3 Morphology...................................................................................... 3

1.31 External morphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

*1.32 Cytomophology

*1.33 Protein specificity

2 Distribution........................................................................................... 4

2.1 Total area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2.2 Differential distribution 4

2.21 Spawn,larvae, and juveniles . .. .. .. . .. .. . 4

2.22 Adults........................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

2.3 Determinants of distribution changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

2.4 Hybridization..................................................................................... 6

3 Bionomics and life history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

3.1 Reproduction................................................................... . . . . . . . . . . . . . . . . . . 6

3.11 Sexuality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

3.12 Maturity............................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

3.13 Mating.................................................................................... 7

3.14 Fertilization.................. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

3.15 Gonads.................................................................................... 7

3.16 Spawning.................................................................................. 9

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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

3.31 Longevity.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

3.32 Hardiness..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

3.33 Competitors... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

3.34 Predators....................................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3.35 Parasites, diseases, injuries, and abnormalities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

*3.36 Physiology, biochemistry, etc.

3.4 Nutrition and growth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3.41 Feeding. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3.42 Food.......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3.43 Growth rate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

*3.44 Metabolism

3.5 Behavior......................................................................................... 13

3.51 Migrations and local movements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

3.52 Schooling....................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

3.53 Responses to stimuli 14

4 Population............................................................................................ 14

4.1 Structure................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

4.11 Sexratio................................................................................... 14

*4.12 Age composition

4.13 Size composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

4.14 Subpopulations....................................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

4.2 Abundance and density. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

4.3 Natality and recruitment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

4.31 Reproduction rates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

4.32 Factors affecting reproduction 17

4.33 Recruitment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

iii
4.4 Mortality and morbidity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

*4.5 Dynamics of population

4.6 The population in the community and the ecosystem 17

5 Exploitation........................................................................................... 17

5.1 Fishing equipment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

5.2 Fishing areas . . . . . . . . . . . . . . . 17

5.3 Fishing seasons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

5.4 Fishing operations and results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

5.41 Effort and intensity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

*5.42 Selectivity

5.43 Catches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

6 Protection and management. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

6.1 Regulatory measures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

*7 Pond fish culture

Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

*No information available.

iv
Synopsis of Biological Data on the Sand Perch,

Diplectrum formosum (Pisces: Serranidae)

GEORGE H. DARCyt

ABSTRACT

Information on the biology and fishery resources of a common western Atiantic serranid, Dip/ee/rum for.

mosum, is compiled, reviewed, and analyzed in the FAO species synopsis style.

INTRODUCTION

The sand perch, Diplee/rum formosum, is a small serranid occurring along the southeastern Atlantic coast of the United States,

throughout the Gulf of Mexico and much of the Caribbean Sea,

and along the coast of Central and South America south to about

Uruguay. Sand perch occur from shore to depths of over 130 m.

They are sometimes abundant and widely distributed over sand or

shell bottoms, particularly in the South Atlantic Bight, on the

West Florida Shelf, and on Campeche Bank. Because of their

abundance, they may be important as forage for larger predators,

such as snappers, groupers, porgies, and sharks. Sand perch enter

commercial landings primarily as by-catch of the shrimp and

bottomfish fisheries; many are discarded. Despite their small size,

sand perch are desirable recreational species due to their excellent

food quality. They are commonly caught by recreational and subsistence fishermen, especially on the west coast of Florida. This

paper summarizes information on the sand perch in the FAO

species synopsis format.

I IDENTITY

1.1 Nomenclature

1.11 Valid name

Diplectrum formosum (Linnaeus 1766) (Fig. I).

Sand perch, Dipleetrum formosum Linnaeus 1766:488-489,

type locality: Carolina. The name is derived from the Greek di-

(two) and plektron (cock's spur), referring to the preopercle, which

bears two clusters of spines, and the Greek formosus, meaning

beautiful.

1.12 Objective synonymy

The following synonymy is based on Bortone (1977):

Perea formosa Linnaeus 1766

Serranus radians Quoy and Gaimard 1824

Serranus faseicularis Valenciennes in Cuvier and Valenciennes

1828

'Southeast Fisheries Cenler, National Marine Fisheries Service, NOAA, 75

Virginia Beach Drive, Miami, FL 33149-1099.

Serranus irradians Valenciennes in Cuvier and Valenciennes

1828 (in part)

Diplectrum fasciculare Holbrook 1855

Centropristis fascicularis Gunther 1859

Centropristis radians Gunther 1859

Diplectrum radians Poey 1876

Serranus formosus Jordan 1885

Diplectrum formosum Goode and Bean 1886

Centropristis formosus Berg 1895

Haliperca formosa Miranda-Ribeiro 1904

Diplectrum formosus Jordan, Evermann, and Clark 1930

Diplectrum fascicularis Poey 1955

Diplectrum radiale Ringuelet and Aramburu 1960 (misidentification)

1.2 Taxonomy

1.21 Affinities

Suprageneric

Phylum: Chordata

Class: Osteichthyes

Superorder: Acanthopterygii

Order: Perciformes

Suborder: Percoidei

Family: Serranidae

Systematics of the family Serranidae have been discussea by

several authors. Jordan and Eigenmann (1890) reviewed the

American serranids and included six subfamilies. Three of these

(Grammistinae, Latinae, and Percichthyinae) are usually excluded

from Serranidae in recent literature. Kendall (1979) considered

the taxa Serraninae, Anthiinae, Epinephelinae, and

Liopropominae subfamilies, but Smith (1978) treated Anthiinae

as a separate family. Gosline (1966) considered only Serraninae,

Anthiinae, and Epinephelinae to be subfamilies of Serranidae.

Diplectrum belongs to the subfamily Serraninae.

Generic

The genus Diplectrum is composed of nine eastern Pacific and

three western Atlantic species (Rosenblatt and Johnson 1974;

Bortone 1977), and is endemic to tropical and subtropical shelf

waters of the Western Hemisphere. All species of Diplectrum are

hermaphroditic.
Figure I.-Adult Diplectrum !ormosum. (From Jordan and Eigenmann 1890.)

The genus was characterized as follows by Bortone (1977).

Small percoid fishes, moderately compressed; body depth

moderate to slender; jaws terminal; dorsal and ventral body profiles nearly equal. Caudal fin truncate or slightly forked; upper

caudal lobe usually somewhat longer than lower, sometimes extended as elongated filament. Dorsal fin continuous, only slightly

indented between spinous and soft dorsals. Villiform teeth present

in upper and lower jaws; both jaws with 5-10 small caniform

teeth; vomer with villiform teeth in wedge or V-shape; palatines

with thin, elongate band of villiform teeth. Preopercle forms

almost right angle between vertical and horizontal limbs, both

limbs serrated along their entire lengths; posterior preopercular

angle expanded as a spur of various shapes, composed of

numerous enlarged spines; opercle with 3 spines or projections

posteriorly, the middle spine the longest; posttemporal serrate; top

of cranium smooth. Pelvic fin origin under or in advance of pectoral fin. Branchiostegals 7; vertebrae 10+ 14. Predorsal fin supports

weakly developed. Dorsal fin X, 12; anal fin Ill, 7 or 8; pectoral

rays 14-19. Scales ctenoid; snout naked and top of head naked

posteriorly to vertical line with preopercular arm; dorsal fin

unscaled; pored lateral line scales with distal portion of pore projecting parallel to body midline.

Dipleetrum was erected by Holbrook (1855:32) with the type

species Serranus fasieularis Valenciennes = Perea formosa by

subsequent designation of Goode and Bean (1886:203). Synonyms

include Dipleetron Troschel 1858 (improper emendation) and

Haliperea Gill 1863 (type species Serranus bivittatus Valenciennes, by monotypy) (Bortone 1977). No subgenera were recognized by Bortone (1977).

Specific

The following species diagnosis is from Bortone (1977):

Preopercle with 2 clusters of radiating spines from upper and

lower posterior corners; scales small; cheek scale rows A (from the

posteroventral angle of the eye downward and posteriorly toward

the lower preopercular angle) 9-16 (mode 11); cheek scale rows B

(from the posterior serrated upper arm of the preoperclt: down2

ward and anteriorly to the distal corner of the maxillary) 12-18

(mode 16); 8-12 scales above lateral line; pectoral rays 16 modally; spinous-dorsal profile nearly parallel to dorsum.

Two other species of Dipleetrum, D. radiale and D. bivittatum,

occur in the western Atlantic. Dipleetrum formosum is distinguished from these species by having 2 spiny lobes on the preopercle,

its larger size as an adult, 16 (modally) pectoral rays (D. bivittatum

has 15, D. radiale 17), smallest scales and highest mean scale

counts, broadest lachrimals, and narrowest maxillaries (Bortone

1977; Smith 1978). As juveniles, D. formosum may be separated

from the other species by smaller eye diameter and by scale counts

(Bortone 1977).

Osteology of Do formosum was described in detail by Bortone

(1977); the bones of the skull are illustrated in Figure 2. According to Bortone, D. formosum is an apomorphic species with

respect to high scale counts, the enlarged preopercular spurs, and

other characters, and may represent a phylogenetic line long

separated from other members of the genus.

1.22 Taxonomic status

Dipleetrum formosum is generally considered a morphospecies

in recent literature.

1.23 Subspecies

Two morphological populations of D. formosum were identified

by Bortone (1977); these he recommended be recognized as subspecies. The northern subspecies, D. formosum formosum (Linnaeus 1766), occurs along the Atlantic and Gulf of Mexico coasts

of North America, and the southern subspecies, D. formosum radians (Quoy and Gaimard 1824), occurs in the Caribbean Sea and

along the coast of South America. The subspecies differ little in

meristic characters, except for gill raker counts, but differ in morphometry (Table 1). The number of gill rakers increases from

north to south, with a mean of 18.83 (range 17-22) in D. formosumformosum and 22.01 (19-24) in D. formosum radians.
Figure 2.-Right lateral view of skull of Diplec/rum formosum.

(From Bortone 1977.)

Table I.-Mean proportional data In thousandths of standard length (SL) for

two Diplectrum formosum subspecies. (From Bortone 1977, table 9.)

Subspecies

Character D. fonnosum fonnosum D. fonnosum radians

SL(mm) 140 119

TL 1.340 1,269

FL 1,191 1,183

Body depth 275 270

Head length 359 362

Postorbital length 191 192

Snout length 113 101

Lacrimal width 59 52

Maxillary width 26 27

Orbit length 74 83

Orbit width 59 63

Interorbital width 74 73

Upper jaw length 160 160

Lower jaw length 179 181

Cheek height 125 119

Cheek length 128 129

Caudal peduncle depth 134 125

Pectoral fin length 235 238

Pelvic fin length 216 210

Predorsal length 349 350

Pelvic origin to lower jaw 341 342

Pelvic origin to anus 306 314

Postanal length 402 401

Dorsal spine height

first 51 53

second 78 79

third 95 100

fourth 105 110

fifth 110 113

Anal spine height

first 25 29

second 47 53

third 70 74

1.24 Standard common names, vernacular names

The accepted common name of D. formosum in the United

States is sand perch (Robins et al. 1980), and standard FAO common names are: English, sand seabass; French, serran de sable;

3

Spanish, serrano arenero (Smith 1978). Other common names and

spelling variations that have appeared in the literature include:

squirrel-fish (Jordan and Eigenmann 1890), squirrel fish

(Baughman 1944), squirrelfish (Springer and Woodburn 1960),

sand fish (Breder 1948), sandfish (Hoese 1958), and sandperch

(Briggs 1958). In Brazil it is known as mixole or margarida

(Brandlio 1964).

1.3 Morphology

1.31 External morphology

The following morphological description is based on Miller and

Jorgensen (1973), Roux (1973), Bortone (1977), and Smith

(1978).

Body moderately small, slender, elongate. Body depth about 3.8

in SL; head about 2.5 in SL. Head and snout moderately long, the

snout flat and unsealed. Maxilla broad, completely exposed at

posterior end. Eye 5.3 times in head. Preopercle with 2 bony

lobes, one at the angle and one above it, each with radiating spines

connected together by a membrane for most of their length.

Number of spines in preopercular spurs variable, 6-15, but usually

9-12. Southern subspecies (D. f radians) has slightly longer

spines, tending to diverge earlier in ontogeny (75 mm SL) than

those of the northern subspecies (D. f formosum), which diverge

at about 90 mm SL. Horizontal and vertical preopercular arms

serrate. Opercle with 3 flat spines, the middle spine the longest.

Gill rakers of moderate length, 11-12, with 2-3 rudimentary

rakers at base of the first arch. Anterior nostril bears a short flap.

Dorsal and anal fins high. Dorsal fin single, unnotched, X, 12.

Anal fin III, 7, occasionally 6 or 8. Bases of dorsal and anal fins

with a few scales on the membranes, but not covered by thick,

scaly skin. Pectoral fins blunt, longer than pelvies, with 15-17

rays (mean 15.95). Caudal fin truncate in southern subspecies,

slightly forked in northern subspecies. Northern subspecies often

has elongated filament on upper lobe of caudal.

Scales ctenoid. About 80 rows of scales along length of body

above lateral line. Pored lateral line scales 49.76 mean (range

46-55). Scale rows in lateral line 76.78 mean (66-88); scale rows
00' 10' 70' 10' '0' '0' '0' '00'

,o·1'+----+-

Figure 3.-Distrlbutlon ofDiplectrumlormosum. (Based on Bortone 1977; Smith

1978.)

2.21 Spawn, larvae, and juveniles

'0·H------j---t--'i;:..;

lO'~--_+---!__-

2.2 Differential distribution

Eggs are presumably planktonic. Larvae are abundant in the

eastern Gulf of Mexico and are widely distributed in surface

waters overlying the West Florida Shelf (Houde 1982; Houde et

0·1H----."..~-__I

shallow embayments. Sand perch are considered characteristic of

the subtropical fish fauna in this region (Hoese 1973).

In the Gulf of Mexico, sand perch occur from the West Florida

Shelf to Campeche Bank (Sauskan and Olaechea 1974), but are

more common in the northeastern Gulf than in the northwestern

Gulf (Hoese and Moore 1977). In the northeastern Gulf, sand

perch occur from shore to reefs, ledges, man-made platforms

(Hastings et al. 1976), and live bottom offshore. They occur on the

Florida Middle Grounds, but are apparently not common there

(Smith et al. 1975). In the northwestern Gulf, they appear to inhabit deeper water farther offshore (Chittenden and Moore 1977).

Sand perch are found on offshore snapper and grouper banks off

both Florida (Bortone 1971a) and Texas (Baughman 1944).

Sand perch are not common in the Bahamas and West Indies.

Bohlke and Chaplin (1968) found them only near Eleuthera I. at

the northeastern end of the Great Bahama Bank. The species appears to prefer continental margins to insular environments.

See 2.21, 2.22, 2.23, and 4.2.

10'

...

.0·1'T----r---.,...---.,.....:r-__....,.........=--.,.---,r--~

...

Sand perch are distributed in waters of the continental shelf

from Virginia to southern Brazil or Uruguay (Fig. 3), including the

Gulf of Mexico, Bahamas, Cuba, Virgin Islands, and the southern

West Indies (Bohlke and Chaplin 1968; Randall 1968; Smith

1978). The most northerly record is from Chincoteague, VA,

though the species seldom occurs north of Cape Lookout, NC; the

Virginia record was taken during an unusually favorable period of

weather (Bortone 1977). Sand perch are not present in Bermuda

(Smith 1976). The southern end of the range is approximately Sao

Paulo, Brazil, with unconfirmed records as far south as Uruguay

(Bortone 1977).

Along the southern Atlantic coast of the United States, sand

perch are associated with open shelf habitat, particularly sand

bottom (Struhsaker 1969). They are widely distributed over such

bottom in 10-132 m of water from off Cape Fear, NC, to south of

Cape Canaveral, FL (Barans and Burrell 1976; Avent and Stanton

1979; Wenner et al. 1979a, c, d). They also occur inshore in

2.1 Total area

2 DISTRIBUTION

above lateral line 9.38 mean (8-12); scale rows below lateral line

21.36 mean (18-26); predorsal scale rows 17.20 mean (14-22).

Anterior chest scales embedded and slightly smaller than lateral

body scales. Scale development was examined by DeLamater and

Courtenay (1974).

Skeletal descriptions were provided by Bortone (1977). A

drawing of the skull appears in Figure 2.

Color pattern complex. Dorsum light brown, lateral surfaces

tan, ventral surface white. Lateral body surfaces with 2 dark

brown stripes in juveniles; in adults the stripes are interrupted to

form 5-7 vertical bars, sometimes more. Caudal spot distinct,

equal to eye diameter, present on upper half of caudal peduncle.

Anal fin with pale yellow spots, pectorals and pelvics pale bluish

white. Midlateral ventral area pink orange in adults. Upper

pharyngeal region yellow with blue flecks. Posterior branchial

cavity pale. Upper medial preopercular surface with small patch

of dark gray; lower medial surface pale white. Snout with 2 bright

blue lines from anterior edge of lachrimal to orbit, with bright

blue irregular lines below joining to form a single line running

irregularly across upper cheek. Additional blue lines on cheek,

opercle, and preopercle. Blue spots on maxillary. Lower jaw

bluish purple. Throat white, a bluish-purple blotch ventral and

posterior to lower jaw symphysis. Top of head brown, with 4-5

bluish-green lines. Seven to 9 longitudinal blue lines trimmed

with yellow along sides of body, bright blue above and paler

ventrally. Anterior dorsal fin with 2 longitudinal blue lines on an

orange field. Tips of dorsal spines bright red orange. Spinous dorsal more deeply forked than soft dorsal. Caudal fin with numerous

orange spots or rows of fused spots or vertical lines on a pale blue

field. Extended caudal filament orange. With preservation, colors

fade, orange and yellow becoming pale and blue becoming dusky.

Coloration may undergo ontogenetic and behavioral changes.

Small individuals have 2 longitudinal stripes as the dominant

lateral pattern, whereas in larger individuals these become broken

into what appears to be vertical bars (Bortone 1971a, b). Bohlke

and Chaplin (1968) reported that sand perch at rest show the

barred pattern and active individuals show the longitudinally

striped pattern.

See 1.21.

4
aU) (Fig. 4). In their survey of ichthyoplankton of the eastern

Gulf of Mexico, Houde et al. (footnote 2) found sand perch larvae

(1.1-16.0 mm SL) most common from spring to fall, though larvae

were present throughout the year; peak abundance was in spring

and summer, particularly in May. In winter, most larvae were

taken in the southern part of the survey area. Larvae were collected in water 9-155 m deep; 50% of the occurrences from stations < 30 m deep, and 90% from stations < 60 m deep.

Specimens were collected at surface water temperatures of 17.0°

to 32.0°C, but were most common at 23.00 ·29.0 0 C. Most were

collected in water of 35.00 / 00 salinity, though some were taken

at salinities as low as 31.0% 0 ,

Figure 4.-Stations at which Diplectrum formosum larvae were collected at least

once during 17 cruises to the eastern Gulf of Mexico, 1971-74. Small points Indicate stations at which no D. formosum larvae were collected. (From Houde et

aI. see text footnote 2, fig. 86.)

Juvenile sand perch are most common in shallow water. In the

South Atlantic Bight, Wenner et al. (1979a, b, c) found smallest

individuals « 10-15 cm FL) in shallow water, usually in 9-27 m.

In the northern Gulf of Mexico, sand perch smaller than about

15.0 cm SL are usually found in water < 15 m deep (Bortone

197Ia). Hastings (1972) observed juveniles near jetties in the

northern Gulf of Mexico, usually in summer and fall, though a few

individuals remained in shallow water year-round. Most juveniles

leave shallow water when the temperature drops below about

20°C (Hastings 1972).

See 2.3.

2.22 Adults

Adult sand perch occupy a variety of habitats from nearshore to

at least 130 m depth. Sand or sand-shell bottom is preferred,

especially near some form of cover, such as reefs, rocks, or manmade structures.

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

Ichthyoplankton abundance and di versity in the eastern Gulf of Mexico. Repon to

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

5

Nearshore, adults are found near jetties (Hastings 1972), reefs

(Starck 1968), mangrove lagoons (Cervigon 1966), and Thalassia

beds (La Monte 1952; Starck and Davis 1966; Bortone 1971 a).

Near rocks or jetties, sand perch usually lie on the bottom or hover

a few centimeters above it (Hastings 1972). Individuals often live

in holes in the bottom or under rocks.

Adults are somewhat more common offshore then in very

shallow water, with largest individuals inhabiting the deepest

water (Smith 1978; Wenner et al. 1979c). Hastings (1972) rarely

found large adults inshore in St. Andrew Bay, FL, especially when

water temperature dropped below 20°C. Sand perch> 15.0 cm

SL are usually found in water> 15 m deep (Bortone 1971 a).

Offshore, sand perch occur over sand, mud, or shell bottoms,

especially near the bases of reefs, rocks, or near upper edges of

depressions in the bottom (Moe and Martin 1965; Bortone 1971 a;

Hastings 1972; Smith 1976; Smith 1978). Baughman (1944)

reported sand perch from smooth sandy or muddy bottom off

Texas. Miller and Richards (1980) listed the sand perch as a reef

species of the South Atlantic Bight. Smith (1976) reported the

species from offshore reefs in 12-42 m in the eastern Gulf of

Mexico and termed it a "secondary reef" species. Sand perch are

found near artificial reefs and around offshore platforms, as well

as near natural limestone reefs (Bortone 1971 a; Hastings 1972;

Hastings et al. 1976; Smith et al. 1979). They have been reported

from water at least as deep as 132 m (Avent and Stanton 1979),

but are not common in depths> 50 m (Bortone 1977; Wenner et

al. 1979a, c, d; Darcy and Gutherz3).

See 2.3 and 4.2.

2.3 Determinants of distribution changes

Distribution of sand perch is influenced by water temperature,

season, salinity, depth, growth stage of the fish, and availability of

suitable habitat.

The sand perch is a subtropical species requiring warm water.

Northern and southern limits of the range are probably determined by low water temperature. Roessler (1970) reported the

temperature range of the sand perch as 18°·34°C, but individuals

have since been found at considerably lower temperatures.

MARMAP trawling in the South Atlantic Bight produced sand

perch from 15.7 °-20.1 °C water in winter-early spring 1975

(Wenner et al. 1979d) and 17.1°-22.0°C water in spring 1974

(Wenner et al. 1979b). Hastings (1972) found a few individuals in

shallow water as cold as 14°C in St. Andrew Bay; none were

found in water below 13°C. Few sand perch were found in

shallow water when the temperature fell below about 20°C

(Hastings 1972). Adults are particularly scarce at low temperatures.

Seasonal changes in water temperature affect local distribution.

In general, sand perch leave shallow water in winter to avoid low

temperatures. Although Wang and Raney (1971) observed no

seasonality in sand perch abundance in the Charlotte Harbor

estuary, FL, most other authors have indicated a decrease in abundance of sand perch inshore in the colder months. Near Cedar

Key, FL, Reid (1954) found sand perch from June to November,

but not during the cooler months of the year. Tabb et al. (1962)

reported that sand perch reach peak abundance in northern

'Darcy, G. H., and E. J. Gutherz. Abundance and distribution of commonly

trawled species of fish and invenebrates, West Florida Shelf, January 1978.

Manuscr. in prep. Southeast Fisheries Center, National Marine Fisheries Service,

NOAA, 75 Virginia Beach Drive, Miami, FL 33149-1099.
Florida Bay, FL, in fall and early winter. Roessler (1970) reported

that 100% of the sand perch caught in his study of Buttonwood

Canal, FL, were caught from September through November.

Hastings (1972) found juvenile and adult sand perch near St.

Andrew Bay jetties in summer and fall, but they were rare or absent in winter. Adults are not usually found inshore in water

cooler than 20°C; most juveniles and adults leave shallow water as

the temperature falls in late fall or early winter (November in St.

Andrew Bay) (Hastings 1972). Ogren and Brusher (1977) found

sand perch to be most abundant in St. Andrew Bay in fall, and

least abundant late winter through early summer (February-June).

Sand perch are able to tolerate temperatures well below 20°C and

are commonly found on offshore reefs in the northern Gulf of

Mexico at 15°C (Hastings 1972). Movement to deeper water may

protect the sand perch against rapid or extreme drops in

temperature.

Although seasonal changes in distribution of sand perch on offshore grounds are not dramatic, some seasonal shift of distribution

with depth may occur. In the South Atlantic Bight, sand perch are

apparently more common in summer than in winter (Hoese 1973;

Barans and Burrel1 1976). Although Barans and Burrel1 (1976)

stated that no seasonal change in depth distribution of sand perch

was discernible in their study, their data (Fig. 5) indicate a slight

shift in abundance towards shallower water (9-18 m) in summer,

and some preference for deeper water (27-55 m) in spring; peak

abundance was in 18-27 m throughout the year. Barans and Burrell speculated that territoriality may explain the relatively

uniform distribution of sand perch in shelf waters of the South

Atlantic Bight.

Sand perch are usually found in waters of relatively high salinity. Most records are from ful1 salinity seawater (35-36% 0) (Bortone 1971 a), though sand perch have been reported from salinities

at least as low as 18% 0 (Gunter and Hal1 1965, St. Lucie estuary,

FL). Tabb and Manning (1961) col1ected sand perch in 39% 0

water in Whitewater Bay, FL. In St. Andrew Bay, sand perch are

most common in high salinity areas (Ogren and Brusher 1977).

Growth stage of sand perch also affects their distribution. Larvae are present year-round in surface waters of the eastern Gulf of

Mexico, but are most abundant in spring and summer (Houde

1982; Houde et a1. footnote 2). In winter, larvae are much more

common over the southern West Florida Shelf than farther to the

north (Houde et a1. footnote 2), possibly indicating that spawning

is more continuous in the south, or that larval survival is reduced

in the north due to cold. Over the entire survey period, however,

there was no statistical difference in larval abundance between the

north and south (Houde 1982). Juveniles and adults occur both inshore and offshore, but adults occur over a wider depth range.

There is a tendency for size of fish to increase with depth (Moe

and Martin 1965; Bortone 1971a, 1977; Smith 1978; Wenner et

a1. 1979a, c). Wenner et aJ. (1979a) calculated mean sand perch

size for catches in three depth zones in the South Atlantic Bight:

9-18 m, 12±2 cm FL; 19-27 m, 15±1 cm FL; 28-55 m, 19±1 cm

FL. Results of other MARMAP surveys were similar (Fig. 6).

Sand perch are usually most abundant at moderate depths.

Barans and Burrel1 (1976) and Wenner et a1. (1979a, b, c, d) found

them to be most common at 19-27 m in the South Atlantic Bight.

Off Tampa Bay, Moe and Martin (1965) found sand perch most

common at 8-14 m. A trawling survey of the West Florida Shelf

(Darcy and Gutherz footnote 3) indicated that sand perch were

most abundant in 19-27 m in January 1978. Durand (1960)

reported sand perch from 20-70 m depth off the Guianas, with

peak abundance in 45-50 m.

When other physical parameters are suitable, presence or

absence of sand perch depends on availability of suitable habitat

(Bortone 1971 a). Sand or sand-shell substrate is preferred (Bortone 1971a; Barans and Burrell 1976). Although often seen over

open bottom, sand perch also associate with bottom irregularities

(Hastings 1972) and may be common near reefs, rock outcrops,

jetties, ledges, or over live bottom (Darcy and Gutherz footnote 3).

Sand perch are dominant members of the pink shrimp community

in the Gulf of Mexico, including Campeche Bank, but are rare or

absent on brown shrimp or white shrimp grounds (Hildebrand

1955; Chittenden and McEachran 1976). Pink shrimp grounds

typically have the sand and calcareous sediments that sand perch

prefer, whereas brown shrimp and white shrimp grounds usually

have soft, muddy sediments.

See 2.21, 2.22, and 4.2.

2.4 Hybridization

No hybrids of the sand perch are known.

3 BIONOMICS AND LIFE HISTORY

Figure 5.-Seasonal depth distribution of Dip/ectrum formosum in the South

Atlantic Bight. (From Barans and Burrell 1976, fig. 9.)

DEPTH STRATA

The testis begins to develop at approximately 90 mm SL (age 2)

and is normally fully mature in specimens of 125 mm SL (age 2 or

3) (Bortone 1971 a). Ovarian tissue is not distinguishable in

individuals smaller than about 90 mm SL, and is not usually

recognizable macroscopically in specimens < 125 mm SL (Bortone 1971 a). Mature eggs and sperm occur simultaneously in sand

perch> 150 mm SL, but whether an individual can function as

both male and female at the same time is unknown (Bortone

1971a).

3.11 Sexuality

The sand perch, like all members of the genus Diplectrum, is a

synchronous hermaphrodite. Separate sexes do not exist.

3.1 Reproduction

3.12 Maturity N: 502

SPRING

N "39B

FAll

IB 27 55 110 le3 366

N: 240

WINTER

N' 771

SUMMER

18 27 55 liD 163 366

100

80

60

:I: 40

0

5 20

...

0

.... ffi 100

0

a:: 80 '"

<l.

60

40

20

6
40

40 A 28-56m

11 30 Y=20Acm B

30 28-55 m n = 116

N = 162 20

20 10 (/)

...J 10 UJ 0 <t (J

::::> z 0 0 UJ 20 19-27m

50 IX: Y-14.7cm > IX:

0 19-27m :J

u..

0

II::

Z 40

30

N=211 (J

g

.....

z

UJ

10

"~ ILl 20 (J 0 IX: 1Il UJ

10 ll. 20 Dip/tic/rum formosum

::::>

Z ... c:::::::. 0

20 Oipl6Cfrum formosum 10

9-18m

10 N=83

5 10 15 20 25 30 5 10 15 20 25 30

FORK LENGTH (em) FORK LENGTH(cm)

Figure 6.-Length-frequency distribution of Diplectrum formosum by depth zone for A) spring 1974, and B) summer

1974, based on MARMAP groundflsh cruises in the South Atlantic Bight. (From Wenner et al. 1979b, fig. 10D; 1979c,

fig.7D.)

3.13 Mating possibly take place (Bortone 1971a). There is no evidence of

internal self-fertilization.

Mating patterns of sand perch are not well-known, but mating

pairs probably form and result in cross-fertilization (Bortone

1971a).

3.15 Gonads

Gonads of sand perch consist of an ovotestis. According to

Bortone (1977), the ovarian portion of the ovotestis appears

yellow and granular, and the testicular portion appears white and

fine-grained. The ovotestis is V-shaped with the ovarian portion

lying anterior to the testicular portion (Fig. 7). The lamellar

ovarian tissues surround the central lumen. Testicular tissue lies in

a wide diagonal band posterior to the ovarian region. The extreme

posteroventral portion forms an "ovarian sinus" (Bortone 1977).

3.14 Fertilization

Fertilization is external. Cross-fertilization probably occurs

between mated pairs, though external self-fertilization could

::

."

..

" 1

..."

c

Ul71 '972

FM.IIJJASON

1973

Figure 7.-Schematic gonad diagram of Diplectrum formosum. Cross-hatched

area, ovarian tissue; white area, testicular tissue; shaded area, posterior

ovarian-sinus region. Top figure is left lateral view, bottom figure is ventral

view, with gonopore represented as a circle. Posterior portion ofthe gonads is to

the right. (From Bortone 1977, fig. 8A.)

Cruis. Date

Figure 8.-Estimated mean abundance (number per 10 m' of sea surface) of

Diplectrum formosum larvae in the eastern Gulf of Mexico, 1971-73. (From

Houde et ai. see text footnote 2, fig. 85.)

7
24·l.':===~::;==-------;;~------;-!=-__--"J 86' 84' 82' 80'

·~Om"". .

"....,.

~\ ... .,

., +\ .. ., .. .,

.... ., '.\" ., . , ., ., .. ., .. .,

t ., .......

• • 0 •• - •

.. ., .. . .. \ .. ., '!' ••. .. .. +. +•• .. +. e. ".. +.

.!• ... :., .....

JIPlECTRU~1

FORMOSU,,",

AUGUST 971

• < I

° 1-10

• 11-100

• 101-1000 o >1000

24·l.'::=====~8~6==·=!..----8~47'-----8*:2""-----~80'

28'

26 0Number under 10m 2

+ 0

.-. - ~Om·. • •• .,

................. ......, . 't ., .

. \~ .. .. ;' . .... e'•••••,

., .... '.... ...........

••••••• ., .... . .. .. .\~ . .. .. .. ....\ . ... e: ••••• .... ... . .. -; .... "iat~/

JIPLECTRUM

FORMO$UM

MAY 1971

26' (.rN""U-m-b-'::""-u-nd-.-'-1-0-m-::2""

+ 0

<I

• 1-10

• 11-100

• 101-1000 o >1000

28'

30'

24·l.'::=====::;8~6;;·;==----·8"!:4;-;·'----·8-;;2;;·'----~80'

.-

,orn._..~~~ ..

'.

.......

. \\~..

.....\.

+ "\

+\

+\\

. l·

.,

< I

• 1-10

• 11-100

• 101 -1000 o >1000

~lPLECTRUM

FORMOSUM

24·l.':====~:=~----,;':-=------:!:c:-----."J 86' 84' 82' 80'

~OV'MB'R 1972 26' Lr-'N--'u'-m':"b'-.,-un-d-.-,-,0-m-"""'2

+ 0

28'

•

.om..... •••••

-..

.', ..

\

'''~\, .

.

DIPLECTRUM

FORMOSUH

< I

° 1-10

• 11-100

• 101-1000 o >1000

28'

FEBRUARY 1972 26·I.r-N

-

u

'-m'-b""o::',=un-d"".'-,-',-0-m--;:-'2

+ 0

30'

82' 80'

.0

84'

'0"'._.......... • ••• "

".

. "\...

.\,-

····.\••• 0\

.\ . .\ . -\ .

\ . !. •.

-; .. 10m2

86'

28'

JIPl£CTRUM

FORMOSUM

I:AV 1973

26' Number under

+ 0

· < I

° 1-10

.Ior"_..... • 11-100

• 101-1000

0 >1000

82' 80' 24'

•

JIPLECTRUM

FORMOSU'"

'om ..

-"'. +

.......

:.....,\-

.\

.. \" ° • JANUARY 1~73

26' LrN::"'u::"'m

- b

-.-,-un-d"-.-'-1:":0:"""m""2 0\..

+ 0 ,','. I

• 1-10 .:"i • 11-'00

• 101 -1000

o >1000

28'

24·l.':::====8~6:::·=----::8~4·;-------;;;'-;-----;}'

Figure 9.-Distribution and abundance of Dipl'ctrum formosum larvae in the eastern Gulf of Mexico, 1971-73, (From Houde et al. see text footnote 2,

fig. 87.)

8
24·t.=======;;s:::::!..--~t.:----=---.:J 8S' 84' S2' 80'

00""-"""""

.". ... .... . \-

~\ . .. .. \ . -\ . .\ •• +

\

.. 1+

+! .. •

i·'

DlPLECTRUM

FORMOSUM

30'

2S'

24·L.::====;JIS~.==:!---...,S~4"'.,......--- ...82:1=.-----1eo•

AGUST 1973 26·lJ':"N"'''''' ...b'-....r'''"U'''-ncler.,....--,.IO,....m-.;n

+ 0

• <I

• 1-10

• 11-100

• 101-1000 @ >1000

......----

~O"""~"""''''' " . + ..'" '\.

... .. \,4

",

" .. '\ ,

. . \ . .\. .-. +\ ....

\

.. 1-

+/ •

i·'

DIPlECTRUM

FORMOSUM

28'

26.j:P.l...... - ""ul...v-'1""97e;:3_---:"

N""'Hr_IOmZ

+ 0

• <I

• /-10

• 11-100

• 101-1000 @ >1000

•

50"'..........."'-

..'" "Ie

..

.. .. \ .

.......

• "\e++,

.\.. - .. \ .....

..

.'\, ... +

.. j.

-I

I .'

JI PLECTRUM.

FORMOSU'"

28'

~OVE"'ER B73

26·I.r.N-um""b:-.-r-U-nd:-..--:-lo""'m""'z;-'

+ 0

• <I

• 1-10

• 11-100

• 101-1000 @ >1000

24·1.!::::::===::;::;:::!----:=---~='=----:l 86' 84' 12' 10'

30'

Figure 9.-Continued.

3.16 Spawning 3.17 Spawn

Spawning of sand perch may occur at any time during the year,

but is at a peak in spring and early summer. Bortone (197Ia)

observed gonad maturation in sand perch from the northern Gulf

of Mexico beginning in March and April and reaching a maximum in May, though some ovotestes examined contained eggs as

late as September. Specimens examined by Bortone in November

and December had regressed ovotestes. The smallest sand perch

collected by Reid (1954) near Cedar Key (17-39 mm SL) were

from May and June. Smallest specimens collected by Bortone

(1971a) (11-22 mm SL) were caught in late May and June. Houde

et al. (footnote 2) found sand perch larvae year-round in surface

waters of the eastern Gulf of Mexico, but they were most abundant in spring and summer (Figs. 8, 9), particularly in April

through June (Houde 1982). According to Houde et al. (footnote

2), some spawning occurs in deep shelf waters (100-200 m),

though most larvae were colJected at stations in 50 m of water or

less.

Little is known concerning spawn of sand perch. Largest

ovarian eggs examined by Bortone (197la) were about 0.87 mm

in diameter. Eggs are presumably planktonic.

3.2 Preadult phase

3.21 Embryonic phase

Embryonic development of sand perch has not been described

in the literature.

3.22 Larvae and adolescent phase

Sand perch larvae are planktonic (Houde et al. footnote 2). Two

types of Diplectrum larvae were described by Kendall (1979),

Type 1 and Type 2. Neither type has been associated with a particular species, however, since Diplectrum species have overlapping

9
ranges and meristic features. Generalized DipLectrum larvae are

characterized by the following: Dorsal fin IX-X, 11-13; anal fin

III, 6-9; pectoral fin 15-18; vertebrae 10+14; predorsal pattern

0/0/0,2; body slender; dorsal and pelvic fin spine development

precocious; pigment located on jaw, c1eithrum, anus, anal fin base,

caudal peduncle, and on the dorsal and pelvic fin membranes.

According to Kendall (1979), Type 1 larvae are more common

and more characteristic of the genus than Type 2; D. formosum

probably has Type 1 larvae. Type 1 larvae exhibit most features of

other serranine larvae. Body shape of the larva approaches that of

the adult, and after fin ray formation they are among the most

slender serranine larvae. First dorsal fin spines and pelvic fin

elements form early; none of the early forming dorsal spines are

very elongate. Individual melanophores are more uniform in size

than those of Centropristis larvae. A series of about five spots occurs ventrally on the caudal peduncle, with at least one spot on the

base of the caudal fin. Type 1 larvae are illustrated in Figure 10,

and characteristic spine-bearing bones of the larvae are shown in

Figure II.

3.3 Adult phase

3.31 Longevity

Sand perch reach a total length of about 30 cm (Jordan and

Evermann 1896; Cervigon 1966; Randall 1968; Smith 1978).

though they are usually considerably smaller. The largest specimen examined by Bortone (1977) was 22.3 cm SL (28.5 cm TL)

and 6 yr old. Maximum age is probably 6 to 7 yr (Bortone

1977).

3.32 Hardiness

Sand perch can tolerate temperatures at least as low as 14°C

(Hastings 1972, St. Andrew Bay) and as high as 34°C (Tabb and

Manning 1961, Whitewater Bay). Gilmore et al. (1978) reported

that sand perch remained relatively active during cold waves in

the Tampa Bay, FL, area in 1976 and 1977 when temperatures as

low as 11.6°C were recorded in bottom waters on offshore reefs;

no sand perch were reported killed. Galloway (1941) found dead

sand perch near Key West, FL, following a cold wave in 1940;

water temperature was measured at 13.9°C.

Juveniles and adults usually leave inshore waters in winter and

return in spring (Bortone 1971 a). Larvae have been collected from

17.0° to 32.00 C (Houde et al. footnote 2).

Salinity tolerance of sand perch is at least 18% 0 (Gunter and

Hall 1965, Caloosahatchee estuary, FL) to 39% 0 (Tabb and Manning 1961, Whitewater Bay). Larvae are usually found at salinities

35% 0 (Houde et al. footnote 2). Reid (1954) reported mass

mortality of fish, including sand perch, following a hurricane near

Cedar Key in 1950 that lowered salinity from 23.5 to 9.7% 0 over

a 4-d period. Whether mortality was caused by the salinity drop,

increased turbidity, or both, is not known.

Sand perch were apparently killed or driven away from shallow

(12-18 m) reefs on the west coast of Florida by a red tide in 1971

(Smith 1976). Sightings of sand perch on the reefs dropped drasa

b

.. . .~. .. .•..

Figure 10.-Larvae of Diplee/rum sp., Type I, from the northwestern Atlantic Ocean: a) 5.8 mm notocord length; b) 10.0 mm SL. (From Kendall 1979, fig. 5.)

10
Figure H.-Spine-bearing bones of tbe opercular and posttemporal

regions of a 10.3 mm SL Dip/ectrum sp. Type 1 larva. (From Kendall

1979, fig. 7.)

opercular

posttemporal

subopercular

supracleithrum

tically following the red tide and none were sighted for almost a

year.

3.33 Competitors

Associates of sand perch in open-shelf habitat of the South

Atlantic Bight include longspine porgy, Stenotomus caprinus;

orange filefish, Aluterus schoepfi; searobins (Prionotus spp.); and

inshore lizardfish, Synodus foetens (Struhsaker 1969). Hoese

(1973) found sand perch closely associated with pearly razorfish,

Hemipteronotus novacula, off the Georgia coast. Extensive

MARMAP trawling in the South Atlantic Bight produced data

Table 2.-Closest associated species of Dip/ectrum formosum on sand bottom in

tbe Soutb Atlantic Blgbt based on MARMAP trawUng, 1973-75. Approximate

similarity indices are based on species cluster inverse analysis, Canberra-Metric

correlation on log-transformed data. (Data from Wenner et al. 1979a, b, c, d,

1980.)

Winterearly

Fall Spring Summer Spring Summer

Species 1973 1974 1974 1975 1975

Synodus foetens 0.52 0.52 0.45 0.25 0.53

Stephanolepis hispidus

(= Monacanthus hispidus) 0.41 0.55 0.42

Aluterus schoepfi 0.15 0.39 0.30 0.17

Batistes capriscus 0.15 0.00

P,ionotus carolinus 0.15 -0.05 0.00

Chaetodipterus faber 0.00

Lactophrys quadricornis 0.00 0.00

Raja eglanteria 0.20 0.17

StenolOmus aculeatus

(= S. caprin us) 0.15 -0.05 0.34 0.22

Ophidion grayi -0.10

Mustelus canis -0.10

Ancylopseua quadrocellata -0.10 0.00

Syacium papillosum -0.05 -0.05

Ccnt,···opristis ocyurus -005 -0.05

P"ano/us salmonicolor -0.05 0.00

Ophidion holbrooki -0.05

Trachinocephalus myops -0.05

Scorpaena calcarata -0.05

HemipteronOlus novacula 0.00

'No data.

11

appropriate for species cluster analysis (Table 2) (Wenner et al.

1979a, b, c, d, 1980). Species most closely associated with sand

perch in the South Atlantic Bight, based on MARMAP surveys,

include inshore lizardfish; planehead filefish, Monacanthus hispidus; and orange filefish. Most of these species are widely distributed over sand bottom in depths of < 55 m (Wenner et al.

1979a).

On the West Florida Shelf, a trawling survey in 9-193 m from

off Cape San Bias, FL, to the Dry Tortugas, FL, produced numerous specimens of sand perch (Darcy and Gutherz footnote 3).

Cluster analysis indicated that closest associates of sand perch on

the West Florida Shelf are scrawled cowfish, Lactophrys quadricornis; black sea bass, Centropristis striata; grass porgy, Calamus

arctifrons; shortnose batfish, Ogcocephalus nasutus; conger (Hildebrandia sp.); spottail pinfish, Diplodus holbrooki; and crested

cusk-eel, Ophidion welshi.

In the northern Gulf of Mexico, in 1-7 m of water, Bortone

(1971 a) observed sand perch associated with belted sandfish, Serranus subligarius; pygmy sea bass, Serraniculus pumilio; pigfish,

Orthopristis chrysoptera; spot, Leiostomus xanthurus; pinfish,

Lagodon rhomboides; cocoa damselfish, Pomacentrus variabilis;

slippery dick, Halichoeres bivittatus; crested blenny, Hypleurocheilus geminatus; and doctorfish, Acanthurus chirurgus. In deeper

water (7-36 m), associates of sand perch reported by Bortone included bank sea bass, Centropristis ocyurus; southern sea bass,

Centropristis striata melana; gag, Mycteroperca microlepis; red

snapper, Lutjanus campechanus; vermilion snapper, Rhomboplites

aurorubens; greater amberjack, Seriola dumerili; white grunt,

Haemulon plumieri; red porgy, Pagrus pagrus; and gulf flounder,

Paralichthys albigutta.

To what extent these associated species actually compete with

sand perch is not known. Most of the species listed above are

carnivores, as are sand perch, and some probably compete with

sand perch for food. Since sand perch are smaller than many of

the associated species, juveniles of some of the larger species of

serranids, lutjanids, and carangids may compete with sand perch

more than the adults of those species do.

A study by Bortone et al. (1981) comparing morphology, abundance, food, and feeding habits of D. formosum and D. bivittatum,

in the vicinity of the Dry Tortugas, did not reveal a mechanism for

reducing or eliminating competition between these often sym-
3.34 Predators

3.41 Feeding

3.4 Nutrition and growth

Frequency Average

Food item (%) number

Annelid worms

Polychaetes 4.5 < 0.1

Mollusks

Bivalves 2.6 < 0.1

Crustaceans

Isopods 1.3 < 0.1

Amphipods 11.1 0.4

Stomatopods 0.7 < 0.1

Decapods 67.0 7.1

Shrimp 56.5 6.0

Crabs 37.7 1.1

Echinoderms

Ophiuroids 1.9 < 0 1

Fishes

Perciforms 3.9 < 0.1

Scorpaeniforms 3.9 < 01

Pleuronectiforms 2.6 < 0.1

Tetraodontiforms 0.7 < 0.1

Unidentified 15.6 0.2

Plants 3.9 0.0

Unidentified material 12.3 0.0

Empty 17.5 0.0

3.42 Food

Table 3.-Stomach analysis of 154 specimens of Dip/ecIrum !ormosum from the northern Gulf of Mexico in

percentage frequency of occurrence and average

number of organisms per stomach. (From Bortone

1971a, table 2.)

Sand perch feed primarily on benthic crustaceans and fishes.

Longley and Hildebrand (1941) reported that sand perch fed

mainly on crabs and shrimp at Dry Tortugas. Lowe-McConnell

(1962) and Bohlke and Chaplin (1968) came to the same conclusion based on specimens from the coast of Guyana and the

Bahamas, respectively. At Cedar Key, Reid (1954) found that

70-110 mm SL specimens taken in fall contained caridean and

palaemonid shrimps, and mysids; samples from October contained

primarily penaeid shrimps. Copepods and fish also occurred in

specimens examined by Reid (1954). In Venezuela, Cervigon

(1966) reported shrimp and other decapod crustaceans, engraulid

fishes, amphipods, larval shrimp, and planktonic decapod crustacean larvae among foods eaten by sand perch.

A more thorough study of food habits of sand perch was conducted by Bortone (1971a) in the northern Gulf of Mexico (Table

3). Crabs and shrimp made up the major part of the diet. Amphipods were particularly important in the diets of young fish. Fishes

made up a moderate portion of the diet and included small sea

bass (Centropristis sp.), sea robins (Prionotus sp.), blennies, gobies,

flatfishes (Symphurus sp.), and filefish (Monacan thus sp.). Other

food items included hermit crabs, isopods, stomatopods, crab

zoeae, polychaetes, bivalve mollusks, and ophiuroids.

Another study of food habits of sand perch was conducted by

Bortone et al. (1981) near the Dry Tortugas; food habits of the

sand perch were compared with a sympatric congener, the dwarf

sand perch, D. bivittatum. Sand perch examined contained

primarily amphipods, natantian shrimp, and brachyuran crabs, by

number; and fish, shrimp, and crabs, by volume (Table 4). Among

crustaceans identified from the guts were Alpheusfloridanus, Thor

sp., Sicyonia brevirostris, Periclimenes sp., Solenocera sp., Portunus

sp., Speocarcinus lobatus, Caprella sp., Eurysquilla plumata,

Squilla chydaea, and Squilla heptacantha. Sand perch consumed

2000 2400

SUNSEl

1200 1600

TIME

SUNRISE

60

>- 50

Q.

40

'"' 30

Z

'"' 20 V

a=

'"' 10

Q.

0

2400 0400 0800

Parasites of sand perch include nematodes and digenetic trematodes. Nahhas and Powell (1971) found the trematode Lecithochirium parvum in the stomach of a sand perch from Pensacola Bay,

FL. The nematode Hysterothylacium reliquens has also been

reported from sand perch (Deardorff and Overstreet 1981).

Sand perch are apparently killed or driven off by red tides.

Smith (1976) noted a dramatic reduction in sand perch sightings

on shallow reefs off the west coast of Florida following a red tide

outbreak.

See 3.32.

3.35 Parasites, diseases, injuries, and abnormalities

Sand perch feed primarily on or near the bottom near low reefs

or rocks (Bortone 1971 a). These feeding habits are reflected in the

foods reported from sand perch stomachs. Bortone (1971 a)

described sand perch as aggressive and voracious feeders. No

seasonality of feeding was detected in Bortone's (1971a) study in

the northern Gulf of Mexico. Feeding activity appears to be greatest in mid-morning and mid-afternoon (Bortone et al. 1981) (Fig.

12).

See 3.42.

Figure 12.-Feeding ac1ivlty ofDip/eclrum!ormosum from orr the Dry Tortugas,

FL. (From Bortone et al. 1981, fig. 2.)

Predators of sand perch include snappers and groupers (Jordan

and Gilbert 1883; Jordan 1885); red porgy, Pagrus pagrus

(Manooch 1977); red drum, Sciaenops ocellatus (Overstreet and

Heard 1978); and bull shark (Branstetter 1981). Other large

piscivores, such as jacks, barracudas, and flatfishes, probably also

prey on sand perch. According to Bohlke and Chaplin (1968),

sand perch dart into holes to escape predators.

patric species. Jaw structures, gill raker structures, body size,

mouth size, and time of feeding were remarkably similar, and

there were few differences in the food habits observed. Some

competitive exclusion may occur due to competition that results

in segregation based on substrate preference (Bortone et al. 1981).

12
Table 4.-8tomach content analysis of 326 Diplec/rumformosum (100 empty). (From Bortone et 01. 1981,

table 1.)

Vol./fish Vol./item Vol. {JAIl/item

Food item % No. No./fish % Vol. (JAl/g) (JAl/g) per g fish

Nematoda 1.3 0.05 -'

Polychaeta 2.2 0.08 4.8 20 232 5.80

Unident. worms 0.7 0.03

Natantia 21.5 0.83 20.7 84 102 2.54

Brachyura 8.5 0.33 13.0 53 161 4.03

Stomatopoda 0.9 0.04 4.9 20 563 14.09

Mysidacea 0.7 0.Q3 0.1 1 17 0.42

Cumacea 0.3 0.01

Amphipoda 29.5 1.14 0.3 1 1 0.Q3

Unident. crustaceans 16.1 0.62 12.5 51 82 2.06

Mollusca 3.5 0.13 1.2 5 37 0.92

Ophiuroidea 6.4 0.23 5.8 24 95 2.37

Pisces 6.1 0.23 31.2 127 542 13.56

Unidentifiable 2.3 0.09 5.6 23 255 6.39

'Measured volume displaced < 0.1 ml.

5,.....---------------,

STANDARD LENGTH IN MM

nearly the same frequency of food items as the dwarf sand perch,

though the sand perch contained a more even distribution of the

items.

3.43 Growth rate

Otoliths can be used to age sand perch, annuli being formed in

March-April in the northern Gulf of Mexico (Bortone 1971 a).

Fish length and otolith radius are related as follows:

Y = 0.12002 + 0.02753X (r2 = 0.988)

where Y is otolith radius in millimeters and X is standard length in

millimeters (Bortone 1971 a). Growth rate is fastest in the first 3

yr of life, slowing after annulus 3 is laid down (Table 5, Fig. 13).

Length and weight of sand perch are related as follows:

log W = -4.67194 + 3.04051 log L

where W is body weight in grams and L is standard length in millimeters (n = 194; range = 21-223 mm SL, 0.3-331.0 g).

Table 5.-Mean annual increase in standard length (81.) of Diplec/rum

formosum in the northern Gulf of Mexico based on otolith readings.

(Data from Bortone 1971a, table 1.)

Year

5

15

10

I 5

lL.

a 5 0<

UJ

d)

::>

Z

5

10

5

5

50 100 150 200

6

5

3

2

o

SL increase (mm)

SL(mm)

2 4 5 6

34.7 43.2 59.5 37.8 21.0 11.2

34.7 77.9 137.4 175.2 196.2 207.4

Figure 13.-Lengtb-frequency histograms of Dipleclrum formosum aged by

otoliths for age-groups 0 through 6. (From Bortone 1971a, Og. 5.)

3.5 Behavior

3.51 Migrations and local movements

Sand perch are not known to undergo large-scale migrations,

but evidence indicates that there is seasonal inshore-offshore

movement. Both Bortone (/971 a) and Hastings (1972) observed a

decrease in sand perch abundance in shallow water as the temperature dropped in late autumn. A few juveniles remain in shallow

water all winter, but most juveniles and virtually all adults are ab13

sent from shallow water in winter (Hastings 1972). Juveniles and

adults return to shallow water in spring when waters warm. Since

sand perch are common on offshore reefs in winter, Hastings

(1972) theorized that the fish leaving shallow water winter offshore.

Local movements are probably slight. Sand perch are territorial,

living in holes in the bottom or under rocks; some holes they dig

themselves by vibrating their bodies rapidly to throw up loose

sand, others they find ready-made (Longley and Hildebrand 1941;

Bohlke and Chaplin 1968). When approached by divers or large

fish, sand perch dive into cover (Longley and Hildebrand 1941).

Young have been seen darting into depressions occupied by stone
crabs (Bortone 1971a). Bortone (197la) observed a sand perch in

an aquarium bury itself in the sand with only its eyes protruding,

indicating a possible escape reaction. One fish may occupy the

same hole for a long period of time (Bohlke and Chaplin 1968).

Each sand perch maintains a home territory which it defends

against intruders. Barans and Burrell (1976) theorized that this

territoriality could explain the fairly uniform distribution of sand

perch on the continental shelf of the South Atlantic Bight. Individuals have been shown to return to their home territories if

displaced (Bortone 1971a).

See 3.52.

3.52 Schooling

Sand perch do not appear to school; territoriality tends to space

the individuals apart. Size of home territories increases with increasing size of the fish (Bortone 1971 a). Individuals <40 mm TL

occur as close together as 1 m apart, whereas 100 mm individuals

are found about 3 m apart (Bortone 1971 a).

3.53 Responses to stimuli

Sand perch respond to falling water temperatures by moving to

deeper water. Strong currents may force them to leave their home

territories temporarily (Bortone 1971a). Light also affects activity

patterns, with greatest activity occurring during daylight and

general inactivity at night (Starck and Davis 1966; Bortone

1971a; Hastings et al. 1976) (Fig. 14). Roessler (1965) caught

more sand perch at night than during the day in Biscayne Bay, FL,

possibly because their reduced activity at night slowed their

escape response to the trawl.

See 2.3.

4 POPULAnON

4.1 Structure

4.11 Sex ratio

Sand perch are synchronous hermaphrodites. There are no

separate sexes.

4.13 Size composition

Larval sand perch collected by Houde (1982) had a standard

length mode of 2.1-3.0 mm during seasons when larvae were

abundant and spawning was at a peak (primarily spring and early

summer). During the rest of the year, the modal length increased

to 3.1-4.0 mm.

Length-frequency histograms based on MARMAP surveys of

the South Atlantic Bight over a 2-yr period indicate that the

modal size of sand perch in that area is about 20-22 cm FL and is

quite constant over time (Wenner et a1. 1979a, b, c, d, 1980) (Fig.

15). Summer and fall MARMAP surveys showed a small size class

of sand perch (mode about 5 cm FL) entering the population at

that time. Few individuals> 25 cm FL were collected.

See 2.3, 4.33, Figure 6.

4.14 Subpopulations

There is no direct evidence that subpopulations of sand perch

exist, though the two subspecies named by Bortone (1977) based

on morphometric differences may indicate subpopulations. To

what extent the subspecies are reproductively isolated is not

known.

4.2 Abundance and density

Figure 14.-Aetlvlty pattern ofDiplectrumformosum based on three trials on one

fish In an aquarium. Activity is the number of 10 em squares the fish traversed

per minute. (From Bortone 1971a, fig. 7.j

LIGHT DARK LIGHT

•

•

• . / •

V"\ r / J • • •

\ Ii •

\ . .

•'" '..... .I\J .-.,.-...... .

25

20

15

>-

I-

>

IU 10 <{

5

o

1200 1800 o

TIME OF DAY

0600 1200

14

Sand perch are often quite abundant and may occur over large

areas of bottom. Because they are not schooling fish, extremely

large catches of sand perch are seldom reported. Abundance of

sand perch may not be accurately reflected by trawl catches due to

their ability to dart into holes or under other cover.

In the South Atlantic Bight, sand perch are very common and

are among the most ubiquitous fishes of the open shelf. Struhsaker

(1969) collected sand perch at more than 50% of the stations he

sampled. Barans and Burrell (1976) reported sand perch catch

rates of 0.29 kg/30-min tow in winter to 0.69 kg/30-min tow in

summer, with greatest catch rates in 19-27 m. Sand perch composed about 2% of the total fish catch weight in Barans and Burrell's survey. Observations from research submersibles off the

central east coast of Florida (Avent and Stanton 1979) indicate

that sand perch are dominant members of the sand-bottom communities of the inner and outer continental shelf between lat.

27°30' and 28°30'N.

MARMAP groundfish surveys in the South Atlantic Bight have

provided much information on distribution and abundance ofsand

perch in that area. Sand perch were most abundant in 19-27 m in

all seasons surveyed (Fig. 16), but occurred in a high percentage of

samples in 9-55 m. They contributed the highest proportion of the

total fish catch by number and by weight in 28-55 m (Table 6).

The sand perch was the most abundant serranid in the MARMAP

survey area (Wenner et a1. 1979b). Minimum standing stock estimates of sand perch in the South Atlantic Bight for fall 1973 and

spring 1974 appear in Table 7. Density estimates for spring 1974

were 3.1 individuals/ha and 0.361 kg/ha for log-transformed data
60 2.0 1.0

A

1.5 r- A

40 ~. 1.0 .... 0.5 . ..

20 0.5 ..

h .. .. -r'1

II)

...

c(

Q

-

>

Q

Z

...

o

o

z

100

80

60

40

20

60

40

20

60

40

20

60

40

20

B

c

o

E

•III

•

I

:::)

z

III

U

Z

C

CI

Z

:::)

al

C

III

>

C

...

III

•

...

o

)(

III

CI

Z

2.5

2.0-

1.5·

1.0

0.5

3_0

2.5

2.0

1.5

1.0

0.5-

1.5

1.0

0.5

3.0-

2.5

2.0-

1.5-

1.0-

0.5-

....

....

....

-

....

....

rB

c

D

E

1.0

~0.5

1.0

0.5

1.0

0.5

"1.0

0.5

4ft

4ft

C

I

o

al

III

U

Z

C

CI

Z

:::)

•

C

III

>

C

...

III

•

...

o

)(

III

CI

Z

5 10 15 20 25 30 9-18 19-27 28-55 56-110

FORK LENGTH (eM)

Figure 15.-Length-frequencies of Dipleetrum !ormosllm in the South Atlantic

Bight: A. FaU 1973; B. spring 1974; C. summer 1974; O. winter-early spring

1975; E. summer 1975, based on MARMAP groundllsh cruises. (Based on Wenner et al. 1979a, fig. 14B; 1979b, fig. 100; 1979c, fig. 70; 1979d, fig. 80; 1980,

fig.4F.)

15

DEPTH (M)

Figure 16.-lndlces of relative abundance of Dipleetrum !ormosllm from the

South Atlantic Bight, 1973-75. A. FaU 1973; B. spring 1974; C. summer 1974; O.

winter-early spring 1975; E. summer 1975, based on MARMAP groundllsh

cruises. Shaded columns represent indices based on biomass, open columns

represent indices based on numbers. (From Wenner et at. 1979a, fig. 14A;

1979b, fig. IOC; 1979c, fig. 7C; 1979d, fig. 8C; 1980, fig. 4E.)
Table 6.-Percentage of total catch by number and weight of Dipleetrum formosum in the South Atlantic Bight based on MARMAP groundftsh surveys. (Data

from Wenner et al. 1979a, b, c, d, 1980.)

Depth range (m)

Date of 9-18 19-27 28-55 56-110

survey %No. %Wt. %No. %Wt. %No. %Wt. %No. %Wt.

Fall 1973 1.1 2.8 2.7 10.5 11.1

Spring 1974 0.5 2.5 4.2 6.8 4.6 6.7 0 0

Summer 1974 7.9 6.3 10.8 11.4 23.4 21.6 0 0

Winter-early

spring 1975 1.7 3.5 0.7 7.8 2.0 0 0

Summer 1975 2.8 7.6 6.3 10.1 22.8 ) 1.6 0 0

'Not given.

(Wenner et a1. 1979b). Estimated density ofsand perch in summer

1975 was 6.6 individuals/ha and 0.38 kg/ha, for an estimated

standing stock of 3.71 x 107 individuals or 2.14 x 103 t (metric

tons) in the South Atlantic Bight at 9-55 m (Wenner et al. 1980).

In the eastern Gulf of Mexico, sand perch have been reported

locally common in shallow water, as well as on offshore reefs and

on live bottom on the West Florida Shelf. Large individuals are

frequently caught from bridges and bulkheads in Tampa Bay and

on reefs offshore (Springer and Woodburn 1960). Hastings (1972)

reported sand perch common near jetties and on offshore reefs in

the northeastern Gulf. In St. Andrew Bay, sand perch comprised

2.6% of trawl catches in a survey by Pristas and Trent (1978).

Sand perch are among the dominant fish on natural and artificial

reefs off Clearwater, FL (Smith et al. 1979). A groundfish survey

of the West Florida Shelf (Darcy and Gutherz 1984) indicated that

sand perch are most common in 9-35 m of water in the area between Tampa Bay and Dry Tortugas, (Fig. 17) where they made

up 9.9% of total fish biomass in the survey tows. Sand perch were

present at 35.5% of the stations sampled between Cape San Bias,

FL, and the Dry Tortugas. Over the entire survey area, sand perch

ranked fifth in biomass of all fish species; 5.1% of total fish

biomass was contributed by sand perch. Abundance of sand perch

was greatest in 19-27 m, both in number of individuals caught and

biomass (Darcy and Gutherz footnote 3) (Fig. 18). Hourglass

investigations of the West Florida Shelf indicated that sand perch

may be abundant enough to be of some commercial importance

(Joyce 1968).

Sand perch larvae are among the most abundant and most frequently caught fish larvae in the eastern Gulf of Mexico (Houde et

al. footnote 2); 55.5% of all serranid larvae collected were sand

perch (Houde 1982). Houde et a1. (footnote 2) collected sand

perch larvae on all 17 cruises to the eastern Gulf, though they

t 26°

MEAN CATCH

PlR TOW

0 1-'

10-49

fI 50-9'

• 100+

24° 84° 82° 86°

Figure 17.-Mean catches ofDipleetrumformosum by 20' latitude)( 20'Iongltude

grid zones on the West Florida Shelf based on bottom trawling by the Oregon II,

Cruise 85, January 1978. Catches are per 10-mln tow of a 12.2 m semi-balloon

shrimp trawl. Isobaths are 18.5, 37, and 185 m.

were most abundant from spring to fall. Mean larval abundance

ranged from 0.3 to 29.8 larvae under each 10 m2 of surface area,

with maximum abundance in May. Although larval abundance indicates large sand perch biomass in the eastern Gulf, relatively

high fecundity may mean that the actual biomass is not as large as

the larval abundance indicates (Houde et a1. footnote 2). Because

of their abundance, sand perch larvae may be useful indicators to

monitor effects of environmental changes in the Gulf of Mexico

ecosystem (Houde 1982).

Table 7.-Mlnlmum standing stock estimates of Dipleetrum formosum in the South Atlantic

Bight, 9-55 m In fall 1973 and spring 1974. The Bliss approximation has been used for the

estimate on naturallng transformed data. Data have not been adjusted for vulnerability of this

species to the 3/4 Yankee trawl and should be regarded as minimum estimates. LCL and UCL

are lower and upper 9O'Jl, conftdence limits, respectively. Biomass Is expressed in metric tons.

(Data from Wenner et al. 1979a, table 17; 1979b, table 14.)

Fall 1973 Spring 1974

Standing Standing

stock LCL UCL stock LCL UCL

Number

untransformed 1.47 x 10' 0.91 X 10' 2.03x1O' 1.56x 10' 1.I9xlO' 1.94 x 10'

transformed 1.51 x 10' 1.09x \0' 2.05xl0' 1.75xlO' 1.33x\O' 2.28x 10'

Biomass

untransformed 1.17 x 10' O.72x 10' 1.63 x 10' 2.11 x 10' 1.50x 10' 2.71 x 10'

transformed 1.12 x 10' 0.83x10' 1.45 x 10' 2.03x 10' 1.59x 10' 2.54x 10'

16
2.5 ,---

IX

... r--- CD 1.5 - :::)

Z

0.5 n 9-18 19-27 28-55 56-110 111-193 DEPTH 1M)

CIl

CIl

< 0.5

o

iii

1.0

Figure 18.-lndex of relative abundance of Diplec/rum formosum on the West

Florida Shelf, January 1978. (Darcy and Gutherz see text footnote 3.)

Sand perch are not very common in the northwestern Gulf of

Mexico (Hoese and Moore 1977). They rarely occur on brown

shrimp grounds in the western Gulf (Hildebrand 1954).

On Campeche Bank, sand perch are very common in 11-19 m,

and are among the most abundant fishes of the pink shrimp

grounds (Hildebrand 1954, 1955). During MEXUS-GOLF()4

trawling investigations of Campeche Bank, sand perch were the

third largest component (4.9%) of the fish catch during the day,

and the eighth largest component (1.9%) at night. The average

total biomass of sand perch on Campeche Bank was estimated at

16,000 t by Sauskan and Olaechea (1974).

Off northern South America, sand perch are common in 45-50

m of water on shell and sand bottoms (Durand 1960).

See 2.1, 2.2, and 2.3.

4.3 Natality and recruitment

4.31 Reproduction rates

The reproduction rate of sand perch is not known. Houde et al.

(footnote 2) stated that sand perch may have high relative fecundity (ova produced per gram of somatic tissue).

4.32 Factors affecting reproduction

Reproduction is apparently influenced by season (water temperature). Highest larval densities in the eastern Gulf of Mexico

were reported in spring through fall, particularly in May (Houde et

al. footnote 2), indicating a spawning peak as waters warm (Figs.

8, 9). Some spawning occurs year-round, particularly in the

southeastern Gulf (Houde 1982; Houde et al. footnote 2).

See 2.3 and 3.16.

4.33 Recruitment

Small sand perch « 10 ::m FL) appear in summer and fall trawl

catches in the South Atlantic Bight (Wenner et a1. 1979a, c,

4MEXUS-GOLFO. 1979. Report on MFXUS-GOLFO research activities.

Mimeogr.. 11 p. Southeast Fisheries Center. National Marine Fisheries Service,

NOAA, 75 Virginia Beach Drive, Miami, FL 33149-1099.

17

1980), indicating that small individuals are recruited to the

demersal fish community during that period (see Figs. 6, 15).

See 2.1, 3.16, and 4.13.

4.4 Mortality and morbidity

Sand perch are known to be killed by low temperatures (Galloway 1941), and are suspected of being killed by red tides (Smith

1976). Mortality data for larval sand perch from the eastern Gulf

of Mexico were presented by Houde et al. (footnote 2) (Fig. 19).

Approximately 98% mortality occurs between 2.8 and 7.8 mm

SL.

See 3.32.

4.6 The population in the community and the ecosystem

Sand perch are important members of open-shelf and livebottom communities of the southeastern United States because of

their abundance and their widespread distribution on the shelf.

Associated fishes include sharks, rays, lizardfishes, cusk-eels, sea

basses, porgies, spadefishes, razorfish, scorpionfishes, searobins,

flatfishes, cowfish, filefishes, and trunkfishes (Table 2). A

generalized diagram of the trophic position of sand perch appears

in Figure 20.

Seasonal inshore-offshore migrations of some sand perch may

effect transfer of energy to inshore fish communities in spring and

to offshore communities in fall.

See 3.33, 3.34, and 3.51.

5 EXPLOITAnON

5.1 Fishing equipment

Sand perch are caught on handlines, in fish traps, and in bottom

trawls (Smith 1978). They are a common catch of recreational

fishermen using hook and line from shore, bridges, small private

boats, and party boats. Hastings (1972) reported that they are

fished by charter boats off the Florida Panhandle. Shrimp trawlers

sometimes catch sand perch (Siebenaler 1952; Anderson and

Gehringer 1965; Anderson 1968; Keiser 1977).

5.2 Fishing areas

Sand perch are caught throughout their range (Smith 1978),

though there appears to be no directed fishery. Shrimp trawlers

fishing off North Carolina sometimes land small quantities of sand

perch (Keiser 1977). Sand perch also enter shrimp catches off

northeastern Florida and off Key West (Siebenaler 1952) and industrial bottomfish catches in the northern Gulf of Mexico (Roithmayr 1965).

Recreational catches of sand perch are greatest in Florida,

though they are occasionally caught by headboat fishermen off the

Carolinas in 28-93 m (Huntsman 1976). They are commonly

taken on offshore reefs and on snapper-grouper banks off the west

coast of Florida (Moe and Martin 1965; Bortone 1971 a), as well

as inshore in bays, tidal creeks, and from bridges (Sutherland

1977; Smith 1978). All reported recreational landings of sand

perch in 1979 were from Florida (U.S. Department of Commerce

1980); 10.4% of the catch was from the Atlantic coast and 89.6%

from the Gulf coast. Throughout Florida, 66.2% of reported sand

perch were caught> 4.8 km (3 mi) from shore, 5.5% were caught

in open water < 4.8 km from shore, 2.9% were caught in enclosed
01 PLECTRIJIl FORPlOSUI1

1971

DIPLECTRUM FORMOSUM

1972

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4000

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DIPLECTRUM FORPlOSUM

1973

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Figure 19.-Apparent mortaDty rates ofDip/ectf1lmlormosum larvae

determined by lchtbyoplankton survey cruises to the eastern Gulf of

Mexico, 1971-73. (From Houde et al. see text footnote 2, tlg. 162.)

2.0 4.0 &.0 &.0 '0.0 12.0 '4.0

18
Figure ZO.-Schematic diagram of the major trophic relationships ofDip/ectrum

formosum in the southeastern United States. Direction of arrows is from prey or

food item to consumer.

coastal waters, and 25.2% were from unknown areas. On the

Atlantic coast of Florida most were caught < 4.8 km from shore,

whereas on the Gulf coast most were caught beyond 4.8 km.

See 2.1 and 5.43.

5.3 Fishing seasons

No well-defined fishing seasons for sand perch exist. Hastings

(1972) reported that charter boats off Destin and Panama City.

FL, sometimes target sand perch in winter when larger species are

not available. Sand perch are available offshore year-round, but

are not common inshore in winter; inshore catches are probably

low in winter. In St. Andrew Bay, most reported sand perch

catches from bridges are in October and November (Sutherland

1977).

5.4 Fishing operations and results

5.41 Effort and intensity

There is little effort information available for sand perch.

MARMAP groundfish cruises have provided some catch/effort

data from the South Atlantic Bight. In fall 1973, mean catch/30-

min tow was 7.56 individuals, with a mean weight of 0.563

kg/tow (based on log-transformed catch data) (Wenner et al.

1979a). Spring 1974 catch rates were 8.8 individuals/30-min tow,

and 1.022 kgltow based on transformed catch data (Wenner et al.

1979b).

See 5.43.

19

5.43 Catches

Separate catch statistics are not reported for sand perch (Smith

1978), though recreational fishing surveys have collected some

data on noncommercial catches. Sand perch enter commercial

catches primarily as by-catch of fisheries directed at sciaenid

bottomfishes (Roithmayr 1965) and shrimp (Siebenaler 1952;

Anderson and Gehringer 1965; Anderson 1968; Keiser 1977).

Only small numbers are actually landed. Some are retained for use

as bait for larger fish species, such as groupers (Bortone 1971 a).

Sand perch are of considerable importance as recreational fish,

though they are seldom targeted. They are caught from shore,

bridges, jetties, and small boats inshore, and from larger private,

charter, and party boats offshore (Moe and Martin 1965; Bortone

1971a; Hastings 1972; Huntsman 1976; Sutherland 1977; Smith

1978). Because of their small size, sand perch are not often the

preferred catch, but they are excellent food fish and fight gamely

on light tackle (Jordan and Evermann 1904; La Monte 1952;

Hastings 1972). They are sometimes sought when other larger

species are not available (Hastings 1972).

Recreational catches of sand perch reported in 1979 were entirely from Florida waters (U.S. Department of Commerce 1980).

On the east coast of Florida, 190,000 sand perch were reported

caught by recreational fishermen in 1979; 1,643,000 were

reported from the west coast of Florida. Most were caught from

private or rental boats, some from man-made structures, and a few

from party boats. Most sand perch caught on the east coast of

Florida were caught nearshore, whereas on the west coast most

were farther offshore. The average sand perch kept by recreational

fishermen and recorded in the survey was about 0.22 kg.

See 5.2.

6 PROTECTION AND MANAGEMENT

6.1 Regulatory measures

Sand perch are not regulated under the Fishery Management

Plans of the South Atlantic, Gulf of Mexico, or Caribbean Fishery

Management Councils at this time.

ACKNOWLEDGMENTS

I thank D. M. Allen and A. C. Jones of the National Marine

Fisheries Service, Southeast Fisheries Center, for critically

reviewing the manuscript.

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"GPO 593 - 081 (1985)