C-55./3 :A//V*S C/^C </0*/

404

/

£

C

%

J& 0"co

o

A*

^rss o*

NOAA Technical Report NMFS Circular 404

Revision of the Sea Basses
of the Genus Diplectrum
(Pisces: Serranidae)

Stephen A. Bortone

September 1977

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

NOAA TECHNICAL REPORTS
National Marine Fisheries Service, Circulars

The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic
distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels
for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing
grounds, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the
development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service
and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyzes, and publishes statistics on
various phases of the industry.

The NOAA Technical Report NMFS Circular series continues a series that has been in existence since 1941. The Circulars are technical
publications of general interest intended to aid conservation and management. Publications that review in considerable detail and at a high
technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management in-
vestigations appear in the Circular series.

NOAA Technical Report NMFS Circulars are available free in limited numbers to governmental agencies, both Federal and State. They are
also available in exchange for other scientific and technical publications in the marine sciences. Individual copies may be obtained (unless
otherwise noted) from D825, Technical Information Division. Environmental Science Information Center, NOAA, Washington. D.C. 20235. Re-
cent Circulars are:

365. Processing EASTROPAC STD data and the construction of ver-
tical temperature and salinity sections by computer. By Forrest R. Miller
and Kenneth A. Bliss. February 1972, iv + 17 p., 8 figs., 3 app. figs. For
sale bv the Superintendent of Documents, U.S. Government Printing Of-
fice. Washington. D.C. 20402.

366. Key to field identification of anadromous juvenile salmonids in 'he
Pacific Northwest. By Robert J. MacConnell and George R. Snyder.
January 1972, iv + 6 p., 4 figs. For sale by the Superintendent of
Documents, U.S. Government Printing Office, Washington, D.C. 20402.

377. Fishery publications, calendar year 1970: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. December 1972, iv + 34 p., 1 fig.
For sale by the Superintendent of Documents. U.S. Government Printing
Office. Washington, D.C. 20402.

378. Marine flora and fauna of the northeastern United States.
Protozoa: Ciliophora. By Arthur C. Borror. September 1973, iii + 62 p., 5
figs. For sale by the Superintendent of Documents, U.S. Government
Printing Office, Washington, D.C. 20402.

367. Engineering economic model for fish protein concentration
processes. By K. K. Almenas. L. C. Durilla, R. C. Ernst, J. W. Gentry, M.
B. Hale, and J. M. Marchello. October 1972, iii + 175 p.. 6 figs., 6 tables.
For sale bv the Superintendent of Documents, U.S. Government Printing
Office. Washington, D.C. 20402.

368. Cooperative Gulf of Mexico estuarine inventory and study,
Florida: Phase I, area description. By J. Kneeland McNulty, William N.
Lindall, Jr.. and James E. Sykes. November 1972, vii + 126 p., 46 figs., 62
tables. For sale bv the Superintendent of Documents, U.S. Government
Printing Office. Washington, D.C. 20402.

369. Field guide to the anglefishes (Pomacanthidae) in the western
Atlantic. By Henry A. Feddern. November 1972, iii + 10 p., 17 figs. For
sale bv the Superintendent of Documents, U.S. Government Printing Of-
fice, Washington, D.C. 20402.

370. Collecting and processing data on fish eggs and larvae in the
California Current region. By David Kramer, Mary J. Kalin, Elizabeth
G. Stevens, James R. Thrailkill, and James R. Zweifel. November 1972.
iv + 38 p., 38 figs., 2 tables. For sale by the Superintendent of
Documents, U.S. Government Printing Office, Washington, D.C. 20402.

371. Ocean fishery management: Discussion and research. By Adam A.
Sokoloski (editor!. (17 papers, 24 authors.) April 1973, vi + 17.3 p., 38
figs., 32 tables, 7 app. tables.

379. Fishery publications, calendar year 1969: Lists and indexes. By Lee
C. Thorson and Mary Ellen Engett. April 1973, iv + 31 p.. 1 fig. For sale
bv the Superintendent of Documents, U.S. Government Printing Office.
Washington, D.C. 20402.

380. Fishery publications, calendar year 1968: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. May 1973, iv + 24 p., 1 fig. For
sale by the Superintendent of Documents, U.S. Government Printing Of-
fice. Washington, D.C. 20402

381. Fishery publications, calendar year 1967: Lists and indexes. By Lee
C. Thorson and Mary Ellen Engett. July 1973, iv + 22 p., 1 fig. For sale
by the Superintendent of Documents, U.S. Government Printing Office,
Washington. D.C. 20402.

382. Fishery publications, calendar year 1966: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. July 1973, iv + 19 p., 1 fig. For
sale by the Superintendent of Documents, U.S. Government Printing Of-
fice, Washington, D.C. 20402.

383. Fishery publications, calendar year 1965: Lists and indexes. By Lee
C. Thorson and Mary Ellen Engett. July 1973, iv + 12 p.. 1 fig. For sale
bv the Superintendent of Documents, U.S. Government Printing Office,
Washington, D.C. 20402.

372. Fishery publications, calendar year 1971: Lists and indexes. By
Thomas A. Manar. October 1972. iv + 24 p.. 1 fcg. For sale by the
Superintendent of Documents, U.F. Government Printing Office,
Washington. D.C. 20402.

384. Marine flora and fauna of the northeastern United States. Higher
plants of the marine fringe. By Edwin T. Moul. September 1973, iii + 60
p.. 109 figs. For sale by the Superintendent of Documents, U.S. Govern-
ment Printing Office. Washington, D.C. 20402.

374. Marine flora and fauna of the northeastern United States.
Annelida: Oligochaeta. By David G. Cook and Ralph O. Brinkhurst. May
1973. iii + 23 p., 82 figs. For sale by the Superintendent of Documents.
U.S. Government Printing Office, Washington, D.C. 20402.

385. Fishery publications, calendar year 1972: Lists and indexes. By Lee
C. Thorson and Mary Ellen Engett. November 1973. iv + 23 p.. 1 fig. For
sale by the Superintendent of Documents, U.S. Government Printing Of-
fice. Washington, D.C. 20402.

375. New Polychaeta from Beaufort, with a key to all species recorded
from North Carolina. By John H. Day. July 1973, xiii + 140 p., 18 figs.. 1
table. For sale bv the Superintendent of Documents, U.S. Government
Printing Office. Washington, D.C. 20402.

386. Marine flora and fauna of the northeastern United States. Pyc-
nogonida. By Lawrence R. McCloskey. September 1973, iii + 12 p., 1 fig.
For sale bv the Superintendent of Documents, U.S. Government Printing
Office, Washington, D.C. 20402.

376. Bottom-water temperatures on the continental shelf. Nova Scotia
to New Jersey. By John B. Colton. Jr. and Ruth R. Stoddard. June 1973.
iii + 55 p.. 15 figs.. 12 app. tables. For sale by the Superintendent of
Documents. U.S. Government Printing Office. Washington, D.C. 20402.

387. Marine flora and fauna of the northeastern United States.
Crustacea: Stomatopoda. By Raymond B. Manning. February 1974, iii +
6 p., 10 figs. For sale by the Superintendent of Documents, U.S. Govern-
ment Printing Office. Washington, D.C. 20402.

Continued on inside back cover

NOAA Technical Report NMFS Circular-404

^qJ3 spa

rMENl Of

Revision of the Sea Basses
of the Genus Di plectrum
(Pisces: Serranidae)

Stephen A. Bortone

September 1977

c

c

Q

U.S. DEPARTMENT OF COMMERCE

Juanita M. Kreps, Secretary

National Oceanic and Atmospheric Administration

Robert M. White, Administrator

National Marine Fisheries Service

Robert W. Schoning, Director

The National Marine Fisheries Service (NMFS) does not approve, rec-
ommend 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 pro-
motion 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

Materials and methods 1

Genus Diplectrum Holbrook 1855 2

Osteology 3

General osteology of Diplectrum 3

Comparative osteology of Diplectrum spp 4

Notes on Serranus osteology 8

Comments on the systematic status of Diplectrum and Serranus 8

Status of the subgenus Haliperca Gill 9

Comments on the definition of Serraninae 9

Comments on the definition of Serranidae 9

Gonad morphology 9

Key to species of Diplectrum 10

Diplectrum formosum (Linnaeus 1766) 11

Diagnosis 15

Description 16

Northern form 16

Southern form 16

Color 16

Size 19

Habits 19

Distribution 19

Variation 20

Systematic comments 20

Specimens examined 20

Diplectrum bivittatum (Valenciennes 1828) 22

Diagnosis 22

Description 22

Gulf of Mexico 23

Caribbean and South America 23

Intermediate population from South Florida, Florida Keys, Tortugas, and Cuba 23

Color 23

Size 24

Habits 24

Distribution 24

Variation 24

Systematic comments 26

Specimens examined 26

Diplectrum radiale (Quoy and Gaimard 1824) 26

Diagnosis 27

Description 27

Color 27

Size 27

Habits 27

Distribution 27

Variation 27

Specimens examined 27

Diplectrum pacificum (Meek and Hildebrand 1925) 28

Diagnosis 29

Description 29

Color 29

Size 30

Habits 30

Distribution 30

Variation 30

Specimens examined 30

iii

Diplectrum rostrum (Bortone 1974) 30

Diagnosis 30

Description 30

Color 31

Size 31

Habits 31

Distribution 31

Variation 31

Specimens examined 32

Diplectrum conceptione (Valenciennes 1828) 32

Diagnosis 32

Description 32

Color 33

Size 33

Habits 33

Distribution 33

Variation 33

Specimens examined 33

Diplectrum maximum (Hildebrand 1946) 33

Diagnosis 33

Description 34

Color 34

Habits 34

Distribution 34

Variation 34

Specimens examined 35

Diplectrum sciurus (Gilbert 1892) 35

Diagnosis 35

Description 35

Color 35

Size 36

Habits 36

Distribution 36

Variation 36

Specimens examined 36

Diplectrum euryplectrum (Jordan and Bollman 1890) 36

Diagnosis 36

Description 36

Color 36

Size 37

Habits 37

Distribution 37

Variation 37

Specimens examined 37

Diplectrum labarum (Rosenblatt and Johnson 1974) 37

Diagnosis 37

Description 37

Color 38

Size 38

Habits 38

Distribution 38

Variation 38

Specimens examined 38

Diplectrum eumelum (Rosenblatt and Johnson 1974) 39

Diagnosis 39

Description 39

Color 40

Size 41

Habits 41

Distribution 41

iv

Variation 41

Specimens examined 41

Diplectrum macropoma (Giinther 1864) 41

Diagnosis 41

Description 42

Color 42

Size 42

Habits 42

Distribution 42

Variation 42

Specimens examined 42

Zoogeography 43

Acknowledgments 44

Literature Cited 45

Figure

1. Diplectrum formosum, Florida 3

2. Diplectrum formosum, 165 mm SL, North Carolina 4

3. Diplectrum formosum, jaw suspension and suborbital bones 4

4. Diplectrum formosum, caudal complex and 10th, 11th, and 12th vertebrae 5

5. Diplectrum formosum, hypobranchial apparatus and pectoral girdle 5

6. Squamation of cheek and serration of preopercle in Diplectrum species 6-7

7. Ontogenetic development of serrae on left preopercles of Diplectrum formosum 8

8. Schematic gonad diagram of Diplectrum species 10

9. Standard length and location found of Diplectrum species 12-14

10. Distribution map for Diplectrum formosum, D. sciurus, and D. conceptione 20

11. Distribution map for Diplectrum bivittatum, D. maximum, and D. euryplectrum 24

12. Distribution map for Diplectrum radiale and D. pacificum 28

13. Distribution map for Diplectrum rostrum and D. macropoma 31

14. Distribution map for Diplectrum eumelum and D. labarum 39

15. Scatter diagram of number of pseudobranch filaments versus standard length for Diplectrum
macropoma and D. eumelum 40

Table

1. Number of pectoral rays in Diplectrum species 16

2. Number of scale rows in lateral line of Diplectrum species and populations 17

3. Scales above lateral line of Diplectrum species 17

4. Scales below lateral line of Diplectrum species 18

5. Number of predorsal scale rows of Diplectrum species and populations 18

6. Number of cheek scale rows A of Diplectrum species 18

7. Number of cheek scale rows B of Diplectrum species 19

8. Total number of gill rakers of Diplectrum species 21

9. Mean proportional data in thousandths of standard length (SL) for Diplectrum species and
populations 25

Revision of the Sea Basses of the Genus Diplectrum

(Pisces: Serranidae)

STEPHEN A. BORTONE1

ABSTRACT

Twelve species of Diplectrum are recognized: three western Atlantic (D. formosum, D.
bivittatum, and D. radiate) and nine eastern Pacific species (D. paeificum, D. conceptione, D. max-
imum, D. rostrum, D. labarum, D. eumelum, D. sciurus, D. macropoma, and D. eury plectrum). All
species are hermaphroditic and possess an ovarian sinus in the posteriormost gonadal area. Diplec-
trum osteological characters aid in defining the genus and in comparing its taxonomic status with
Serranus. Definitions of the Serraninae and Serranidae are modified as a result of Diplectrum os-
teological characters. The genus is basically restricted to the warm tropical-subtropical coastal areas
of the new world. Diplectrum bivittatum, however, often occurs at insular localities, and D. con-
ceptione occurs in colder, temperate Peruvian waters. Diplectrum formosum is considered to have a
northern subspecies, D. f. formosum, and a southern form, D. f. radians; D. bivittatum is also com-
posed of two subspecies: D. b. bivittatum from South American and Caribbean areas and D. b. ar-
cuarium from the Gulf of Mexico. An intermediate population occurs in the vicinity of southern
Florida. Diplectrum radiate and D. paeificum are "geminate species" and similar morphologically.
Diplectrum mexicanum is considered a junior synonym of D. macropoma.

INTRODUCTION

MATERIALS AND METHODS

The genus Diplectrum is composed of nine Pacific
(Rosenblatt and Johnson 1974) and three Atlantic
species distributed in the tropical-subtropical areas of
the Western Hemisphere. The species are colorful sea
basses which bear an enlarged preopercle. All species
thus far examined are considered to be synchronous her-
maphrodites. Because of their small size (usually less
than 300 mm) they are presently of little commercial im-
portance, yet as they are abundant at some localities
they may have potential as a food or industrial protein
resource. Considerable confusion occurs in the literature
because the genus has never been adequately defined.
Species recognition has been difficult owing to inac-
curate literature accounts and mislabeled museum
material. Robins and Starck (1961) performed a
preliminary investigation on the genus Serranus. This
was the first major step in clarifying the subfamily Ser-
raninae since the revision of the Serranidae by Jordan
and Eigenmann (1890). Revisions of such genera will
allow ichthyologists to begin assembling our knowledge
and permit a system of order in the Serranidae (Smith
1971).

Herein is presented a systematic revision of the genus
Diplectrum. The purposes of this paper are to define the
genus, to make a preliminary evaluation of the status of
the genus among the Serraninae, to describe the species
recognized as associated with the genus, and to sum-
marize our current knowledge of the life history and
zoogeography of the species.

'Faculty of Biology, University of West Florida, Pensacola, FL 32504.

Under the subsections entitled "Specimens examined"
the following institutional abbreviations are used:
BMNH, British Museum (Natural History); CAS,
California Academy of Sciences; FMNH, Field Museum
of Natural History; FSBC, Florida Department of
Natural Resources; FSM, Florida State Museum; FSU,
Florida State University; GCRL, Gulf Coast Research
Laboratory; GMBL, Grice Marine Biological
Laboratory; INP, Instituto Nacional de Pesca, Mexico;
LACM, Los Angeles County Museum; MCZ, Musem of
Comparative Zoology; MNHN, Museum National d'His-
tiore Naturelle; MZUSP, Museu de Zoologia da Univer-
sidad de Sao Paulo; RMNH, Rijksmuseum van Natuur-
lijke Historie; SIO, Scripps Institution of Oceanography;
SOSC, Smithsonian Oceanographic Sorting Center;
TABL, Tropical Atlantic Biological Laboratory; UA,
University of Arizona; UCLA, University of California at
Los Angeles; UCR, Universidad de Costa Rica; UMML,
Rosenstiel School of Marine and Atmospheric Sciences,
University of Miami; UNC, University of North
Carolina, Institute of Marine Sciences; UPR, University
of Puerto Rico; USNM, National Museum of Natural
History, Washington, D.C.; UTMSI, University of Tex-
as, Marine Sciences Institute; UW, University of
Washington.

When complete information is available for specimens
examined, the abbreviation for the museum is followed
by its museum catalog number, followed by a comma,
the number of specimens in the lot, and the range of
standard length in millimeters in parentheses. Type
material is labeled holotype, paratype, lectotype, or
paralectotype immediately after the catalog number.

Uncataloged museum specimens are referred to by field
numbers, cruise numbers, or other field data in place of
catalog numbers.

All measurements were made with dial calipers and
recorded to 0.1 mm. except standard, total, and fork
lengths greater than 100 mm which were recorded to the
nearest millimeter. Counts and measurements were
made and recorded according to the methods of Hubbs
and Lagler (1964) with the following additions and
modifications: counts for cheek scale rows A, from the
posteroventral angle of the eye downward and pos-
teriorly toward the lower preopercular angle; counts for
cheek scale rows B, from the posterior serrated upper
arm of the preopercle downward and anteriorly to the
distal corner of the maxillary (Ginsburg 1948); predorsal
scale rows, immediately below the predorsal midline
from the base of the first dorsal spine forward; gill rakers,
counted on the left anteriormost arch and include all
rudiments of gill rakers and tubercles; preopercular spur
spines, counted between the upper and lower outside
angles of the preopercular spur; only rays of the left pec-
toral fin were counted; all pseudobranch filaments were
counted on the left side; interorbital width, as the least
fleshy width of the interorbital area; cheek length, from
the distal maxillary edge to the nearest border of the ver-
tical propercular arm; maxillary width, at the broadest
part of the distal maxillary area; lacrimal width, from
the anteroventral corner of the eye to the nearest
anteroventral lacrimal edge; orbit width, the least dis-
tance between fleshy edges; lower jaw length, the length
of mandible; pelvic origin to anus, distance from an-
terior pelvic fin base to base of anal spine; postanal
length, from the anterior anal spine base posteriorly to
the center of the hypural base.

Standard length (SL) is given in millimeters but all
other measurements are presented in thousandths of SL.
Data in the species accounts are presented in the fol-
lowing manner: character and mean value, followed in
parentheses by standard deviation, range, and number of
specimens. For meristic characters, the standard devia-
tion and number of specimens are not included in the
description but are found in the appropriate tables.
Proportional measurements are presented only for
specimens larger than 75 mm SL. Counts for meristic
characters are presented for all fishes regardless of stan-
dard length.

Color descriptions were taken from live and/or preserv-
ed specimens and are indicated as such.

Osteological terminology generally follows that of
Smith (1971), Monod (1968), and McAllister (1968), and
the abbreviations used are as follows: AR, articular; B,
branchiostegals; BB, 1-3, basibranchials; BO, basioc-
cipital; BS, basisphenoid; CB, 1-5, ceratobranchials;
CH, ceratohyal; CL, cleithrum; COR, coracoid; CP, 1-4,
pre-ural centrum; E, 1-3, epurals; ECT, ectopterygoid;
EH, epihyal; END, endopterygoid; EPO, epiotic; EXO,
exoccipital; FR, frontal; H, hyoid; W, IV, hypurals; HA,
haemal arch; HB, 1-3, hypobranchials; HS, haemal
spine, GH, glossohyal; IC, intercalar; LAC, lacrimal; LE,
lateral ethmoid; LH, lower hypohyal; MET, metaptery-

goid; PA, parietal; PAL, palatine; PAS, parasphenoid;
PH, parhypurals; PRO, prootic; PT, posttemporal; PTO,
pterotic; PTS, pterosphenoid; QU, quadrate; R, radial;
RA, retroarticular; SC, scapula; SCL, supracleithrum;
SE, supraethmoid; SO, supraoccipital; SS, subocular
shelf; ST, stegural; SYM, symplectic; UH, upper
hypohyal; V, vomer.

Osteological examination was conducted from radio-
graphs on skeletons prepared by maceration, cleaning by
dermestid beetles, soaking in ammonium hydroxide
(Konnerth 1965), or staining with alizarin and clearing
with trypsin. The method employed for clearing and
staining was the procedure of Taylor (1967) but modified
in that the staining was performed prior to clearing. Ver-
tebral counts were separated into precaudal and caudal
by a plus sign. Method for counting dorsal fin supports
followed that of Smith and Bailey (1961).

Specimens examined osteologically

Diplectrum formoswn. North Carolina: Uncat., 17 (42-
210). Florida Gulf: LACM 33307-1, 1 (205); LACM
33306-1, 1 (228); LACM 33306-3, 1 (210); LACM
33306-2, 1 (210); LACM 33308-1, 1 (164); Uncat., 3
(ad.).

D. bivittatum. Cuba: USNM 110282, 1 (ad). Florida:
Uncat., 1 (103). Texas: Uncat., 1 (ad). Honduras:
TABL 105156, 1 (91); TABL 105160, 1 (85). Panama:
Uncat., 1 (juv). Colombia: UMML 15397, 1 (75).

D. pacificum. Mexico: SIO 65-164, 1 (69). Panama:

Uncat., 4 (117-207); Uncat., 1 (juv).
D. macropoma. Panama: Uncat., 2 (115-117); UMML

26246, 2 (74-84); UMML 25075, 1 (62); UMML 25094,

2 (50-61).
D. rostrum. Bahia de Banderas, Mexico: SIO 65-160, 6

(61-105).
D. conceptione. Peru: USNM 213578 and 211385, 3 (89-

100).
D. maximum. Baja California: SIO 62-106, 1 (juv.).
D. sciurus. Gulf of California: SIO 70-70, 1 (109);

USNM 46571, 1 (paralectotype).
D. euryplectrum. Costa Rica: Uncat., 5 (122-

139). Panama: UMML 26244, 2 (60-73).
D. labarum. Golfo de Tehauntepec, Mexico: SIO 63-

524, 4 (juv.).
D. eumelum. Costa Rica: Uncat., 2 (209-228). Colom-
bia: USNM 211393, 2 (74-88).
Serranus phoebe. North Carolina: Uncat., 1

(53). Cuba: USNM 110279, 1. No locality: UMML

638, 2 (ad.).
S. tabacarius. Cuba: USNM 26662, 1 (ad.).
S. tigrinus. Cuba: USNM 110283, 1 (ad.).
S. notospilus. Carolina: Uncat., 3 (juv.).

GENUS DIPLECTRUM HOLBROOK 1855

Diplectrum Holbrook 1855:32 (type-species: Ser-
ranus fasicularis Valenciennes = Perca formosa
Linnaeus as subsequently designated by Goode
and Bean 1886:203).

Diplectron Troschel 1858:24 (improper emenda-
tion).

Haliperca Gill 1863:236 (type-species: Serranus bi-
vittatus Valenciennes, by monotypy).

Troschel (1858:80) invalidly emended Diplectrum as
Diplectron. This error was later copied by Canestrini
(1860:303, 306). Vieillot (1816:50, as cited by Neave
1939:106) had earlier applied Diplectron as a genus to the
class Aves. Berg (1895:45) wrongly considered the generic
names Diplectrum and Diplectron as homonyms and
therefore thought Diplectrum (Pisces) to be preoccupied
by Diplectron (Aves). Berg then applied the generic
name Centropristis to formosus (= formosum) because
of priority. The generic name Diplectrum is to be retain-
ed for Pisces under Article 56a of the International Code
of Zoological Nomenclature (International Commission
of Zoological Nomenclature 1964).

Description. — Small percoid fishes, moderately com-
pressed, moderate to slender depth with terminal jaws.
Dorsal and ventral body profiles nearly equal. Caudal fin
truncate or slightly forked. Upper caudal lobe usually
somewhat longer than lower. Upper lobe sometimes ex-
tended as elongate filament. Dorsal fin continuous, only
slightly indented between spinous and soft dorsal fins.
Villiform teeth present in upper and lower jaws. Both
jaws also with an outer row of 5-10 small caniniform
teeth. Vomer with villiform teeth in wedge- or V-shaped
pattern. Palatines with thin elongate band of villiform
teeth. Preopercle forms almost right angle between
horizontal and vertical limbs. Both limbs serrated entire
length (except vertical limb in D. euryplectrum).
Posterior preopercular angle expanded as spur of varied
shape, composed of numerous enlarged spines. Oper-
culum with three spines or projections posteriorly, the
middle spine the longest. Posttemporal serrate. Top of
cranium smooth. Midline suture of frontal bones flat.
Epiotic-parietal-supraoccipital area next to supraoc-
cipital crest nearly flat. Supraoccipital crest short and
low, usually with supportive stay close to crest edge
paralleling body line. Epiotic posttemporal facet elon-
gate and narrow with a single posterior projection.
Parietal "wings" elongate and broad. Epiotic posttem-
poral facet not delineated medially by a ridge. Parietal
ridge present but low, not extending forward upon fron-
tals. Lacrimal broader than distal maxillary tip except
in D. sciurus. Supramaxillary absent. Pelvic fin origin
under, or in advance of, pectoral fin. Branchiostegals 7,
vertebrae 10 + 14. Predorsal fin supports weakly de-
veloped, with a formula of 0-0-0-2- or 0-0-2- . Dorsal fin
formula X, 12. Anal fin formula III, 7 or 8. Pectoral rays
14-19. Snout naked and top of head naked posteriorly to
vertical line with upper preopercular arm. Inter- and
subopercles naked. Scales ctenoid. Small scales present
in rows on some species along rays and spines at anal fin
base. Scales absent from dorsal fin. Pored lateral-line
scales with distal portion of pore projecting parallel to
body midline.

All species are synchronous hermaphrodites. Ovarian

tissue comprises the anterior portion of the gonadal
lobes; testicular tissue present posteriorly in a broad
band in varied patterns. Posteriorly and ventrally an
"ovarian sinus" of possible accessory-gland function oc-
curs.

Color pattern in the genus is varied but basically the
snout and cheeks bear spots or lines of yellow, orange, or
blue. Dorsal and caudal fins have a combination of
orange and blue patterns of spots and stripes; either
spots or stripes may be absent. Body is brown dorsally,
pale ventrally, lateral body surface is traversed ver-
tically with dark bars and horizontally, with two dark
stripes. Portions of this pattern are of varied intensity
and distinctness.

The genus is endemic to the western hemisphere and is
generally found in continental coastal areas in tropical-
subtropical life zones.

OSTEOLOGY

General osteology of Diplectrum
(Figs. 1-5)

Vomer with a narrow chevron-shaped patch of
villiform teeth. Villiform medially and short caniniform
teeth laterally present in narrow bands on dentary and
premaxillary bones. Glossohyal teeth absent. Villiform
teeth present on palatines in a thin elongate patch.
Lateral ethmoid with three facets for articulation (see
Gosline 1966). Interorbital area narrow. Frontals with
deep grooves dorsally leading from anterior edge and ter-
minating in a large elongate midfrontal foramen. Dorsal
junction of frontals a broadened surface. Top of cranium
smooth, without ridges, area rounded or somewhat flat-
tened. Epiotics with a single elongate narrow posttem-
poral facet. The facet not sharply delineated from
remaining portion of epiotic by a ridge. Epiotic-parietal-
supraoccipital surfaces lateral to the supraoccipital crest
nearly flat. Parietal ridge low and short, terminating at

Figure 1 . — Diplectrum formosum, Florida; right lateral view of skull.

Figure 2.—Diplectrum formosum, 165 mm SL, North Carolina: 1) left
lateral; B) ventral; and C) dorsal views of neurocranium. See text
page 2 for abbreviations.

frontal border. Posterior frontal area with evidence of
parietal-ridge extension forward and curving laterally,
not a true ridge but as a corner on skull. Posttemporal
fossae large. Pterotic "wings" elongate and broad.
Supraoccipital crest short and low, extending posteriorly
and ventrally to exoccipitals. Anteriorly, supraoccipital
crest base only slightly expanded, terminating at
posterior frontal edge. Intercalar with prominent
posterior projection for articulation with posttemporal.
Parasphenoid straight or nearly so. Prootic-basioc-
cipital-parasphenoid area expanded as prominent otic
capsule. Six subocular bones (Fig. 3). Subocular shelf
composed of third and occasionally a portion of the
fourth subocular bones. Predorsal fin supports thin and
weakly developed, the formula (Smith and Bailey 1961)
0-0-0-2- or 0-0-2-. Vertebral column with 10 + 14 ver-
tebrae. The eleventh vertebra bears the first haemal
spine generally curving posteriorly, slightly flattened dis-
tally, but concave on proximal anteroventral surface
(Fig. 4). Caudal fin type-Vb of Monod (1968). Ceratohyal
with shallow, gently curved dorsal surface.
Branchiostegals seven; 5'/2 on the ceratohyal, IVs on the
epihyal (Fig. 5). The first three attach ventrally to

ceratohyal, remaining branchiostegals having lateral at-
tachment. Operculum with three prominent points or
projections posteriorly. Posttemporal serrate.

Preopercle forming nearly right angle between horizon-
tal and vertical arms. Horizontal arm always serrate.
Preopercular spur with many elongate spines and ex-
panded in various shapes.

Comparative osteology of Diplectrum spp.

Skeletons of various preparations for all species of
Diplectrum except D. radiale were examined. Figures of
D. formosum were prepared and are presented for com-
parative purposes. Although D. formosum may be one of
the more specialized species, comparison will be made on
its osteological morphology and differences with other
species will be noted. The basic osteology for the genus
has been presented above and the only special os-
teological aspects are considered here.

Vomerine-teeth patch forms a broad wedge-shaped
pattern in D. formosum. Diplectrum sciurus has a nar-
rower triangular vomerine-tooth patch. The anterior
parasphenoid bends slightly downward in D. formosum
but is nearly straight for the other species. The lateral

Figure 3.— Diplectrum formosum: A) jaw suspension, left lateral; B)
suborbital bones above, right lateral; below, dorsal. See text page 2
for abbreviations.

B

HA

Figure 4. — Diplectrum formosum: A) caudal complex, left laterial; B) 10 th, 11th, and 12th vertebrae, left lateral. See text pages 5-6 for abbrevia-
tions.

Figure 5. — Diplectrum formosum: A) hypobranchial apparatus, left lateral; B) hypobranchial apparatus, dorsal; C) pectoral girdle, left lateral.

See text pages 5-6 for abbreviations.

parietal ridges are low in all species but slightly more
developed in D. maximum, D. eumelum, and D.
pacificum. Diplectrum pacificum and D. maximum
show some evidence of a medial curl of the ridge near the
frontal parietal borders. In D. formosum the supraoc-
cipital crest originates at an abrupt angle, is higher, and
its posterior edge approaches the exoccipitals at a low
angle. In all other species of Diplectrum the crest
originates at a low angle, its dorsal profile is nearly paral-
lel to the central axis, and its posterior edge is at an
abrupt angle. Also the supraoccipital stay (see Gosline
1966) is absent in D. formosum; its presence in other

Diplectrum is variable between species. The stay is
greatly reduced in D. biuittatum. In D. labarum the stay
runs closely along the dorsal crest edge. In other Diplec-
trum the ridge is present, strong, and parallels the cen-
tral axis. Anterior supraoccipital-crest base is not
noticeably thicker in D. formosum but appears slightly
thickened for all other Diplectrum, Predorsal formula is
0-0-0-2- for all Diplectrum (the third predorsal support
may be missing in a few individuals) except in D. bivit-
tatum, D. conceptione, and D. euryplectrum which have
a formula of 0-0-2- . First haemal spine with a low
protuberance on the posterior surface for D. formosum,

Figure 6.— Squamation of cheek and serration of preopercle in species of Diplectrum: A) D. formosum, 143 mm SL,
UMML 3314, Jacksonville, Fla.; B) D. bivittatum, 104 mm SL, UMML 12324, St. Augustine, Fla.; C) D. pacificum,
133 mm SL, USNM 183980, Gulf of California; D) D. radiale, 197 mm SL, UMML 28749, Trinidad; E) D. rostrum, 117 mm
SL, CAS 25295, Colombia; F) D. conceptione, 145 mm SL, USNM 213478, Ecuador; G) D. euryplectrum, 122 mm SL,

D. bivittatum, and D. sciurus. Posterior haemal spine
smooth in Diplectrum except in D. pacificum where the
posterior haemal spine curve is somewhat exaggerated.
Lacrimal very broad, broadest in D. formosum, except
for D. sciurus which may be considered to have a narrow
lacrimal width. The maxillary is thin, broadest toward
middle in D. formosum. In other species it is broadest
distally but the shaft is thick and somewhat spatulate.
D. sciurus has a broad maxillary tip and the shaft is thin.
The subocular shelf is rectangular, somewhat more
square in D. maximum, D. conceptions, and D. bivit-
tatum. The shelf is generally restricted to the third sub-

ocular bone, but in some individuals the fourth flattens
out medially to form the posterior shelf edge. The post-
temporal is variously serrate in all species. Diplectrum
bivittatum and D. maximum have the least developed
serrae; D. conceptione has enlarged serrae; and fewer ser-
rae are found in D. eumelum, D. euryplectrum, and D.
labarum.

Preopercles vary (Fig. 6), however, the species can be
placed in several morphological groups with regard to
this character. With its broad spur and double cluster of
spines, D. formosum forms a category by itself. A second
group with broad preopercular spurs is composed of D.

UMML 26251, Panama; H) D. eumelum, 189 mm SL, USNM 213813, Colombia; I) D. macropoma, 127 mm SL, USNM
213591, Colombia; J) D. labarum, 134 mm SL, SIO 65-257, outer Baja California; K) D. maximum, 302 mm SL, USNM
200373, Peru; L) D. sciurus, 126 mm SL, SIO 70-70, Gulf of California.

euryplectrum, D. eumelum, and D. macropoma.
Decidedly truncate preopercles are seen in D. labarum
and D. sciurus. Diplectrum pacificum, D. radiale, andD.
rostrum may be grouped together with narrow, truncate
spurs. Diplectrum conceptione and D. maximum form a
final group having a narrow spur with fewer spines.
Diplectrum bivittatum could justifiably be placed in
either the D. pacificum or D. conceptione group.

In juveniles less than 75 mm SL the preopercular spur
is not yet differentiated into the adult form. Figure 7 il-
lustrates developmental changes in the preopercle for D.
formosum. Preopercular spurs of smaller Diplectrum
basically resemble the smallest D. formosum preoper-
cular spur depicted.

Notes on Serranus Osteology

an irregular ridge. Epiotic-parietal-supraoccipital area
not flat, but irregular and uneven. A groove or
longitudinal depression along the junction of the fron-
tals. Pterotic "wings" short and narrow. Horizontal and
vertical preopercular arms join at an obtuse angle and
the preopercular spur is never developed. Lacrimal nar-
row, equal to or narrower than maxillary width. Predor-
sal fin support formula 0-0-0-2-, the interneurals strong.
Lateral supraoccipital crest stay not close to distal crest
edge.

Differences have been emphasized in this partial os-
teological summary of Serranus. However, the notable
character differences between the genera should serve as
a means to define the genus Diplectrum more suc-
cinctly. I wish to stress the need for more complete os-
teological studies of Serranus.

Owing to the obvious affinities of Serranus and Diplec-
trum some characters for the former genus are presented
to clarify the latter. Inferences on relationships are made
below. Only a few species and specimens of Serranus
were examined, but a few obvious features are noted.

Vomer broad and U-shaped. Midlateral parietal ridges
are prominent, extending upon the frontal area anterior-
ly and laterally. Epiotic bears two posterior projections
and the posttemporal facet is broad. This facet is also
sharply delineated from the remainder of the epiotic by

Comments on the Systematic Status of
Diplectrum and Serranus

The genera Diplectrum and Serranus are obviously
similar in many morphological aspects. In the list of 16
characters used by Robins and Starck (1961:261) to
define the genus Serranus only four do not apply to
Diplectrum; those four characters are: "12) Lateral-line
pores opening through tubes directed dorsally . . . ."
[directed parallel to body lines in Diplectrum]; "13) Dor-

Figure 7. — Ontogenetic development of serrae on left
preopercles of Diplectrum formosum: A) 41.7 mm SL,
North Carolina; B) 59.5 mm SL, North Carolina; C)
70.7 mm SL, North Carolina; D) 79.8 mm SL, Florida
Keys.

sal and anal fins with small wedges of scales . . . ." [pre-
sent only on anal fins of Diplectrum]; "16) Size moderate
. . . probably never more than 200 millimeters in stan-
dard length." [commonly reaching 200 mm SL or more in
Diplectrum]; "17) No filamentous spines or rays . . . ."
[D. labarum has filamentous spines]. Admittedly the
above differences are minor, however, if osteological
characters are considered, greater differences between
these genera are apparent. In particular, these are the
shape of the posttemporal facets of the epiotics, parietal
ridge system, fusion line of frontals, preopercular spur
shape and arm angle and width of lacrimal. The gonads
are not similar in patterns of "territorial her-
maphroditism." Smith (1965) has depicted S. tigrinus
(and I have observed this condition in S. fasciatus, S.
phoebe, and S. notospilus) with a thin band of testicular
tissue. All Diplectrum have a wide band of testicular
tissue. Also, but tentatively, the presence of an "ovarian
sinus" with possible accessory-gland function is ap-
parently restricted in the Serraninae to Diplectrum (Bor-
tone in press a).

Serraninae is composed of a closely aligned mor-
phological group of genera. Other workers attempted to
separate these in keys with the character of pelvic fin
origin in relation to pectoral fin (e.g., Norman 1966:227-
228; Po below or behind P] for Serranus; P2 in advance of
Pj for Diplectrum). Actually this character is variable
and contributes little to the separation of these genera.
Overall, the principal characters in distinguishing Ser-
ranus from Diplectrum are the obvious preopercular spur
and the wide lacrimal bone of Diplectrum. Other
previously mentioned characters may also be useful in
separation.

On the basis of scale morphology, McCully (1961) sug-
gested that Diplectrum spp. be placed in the genus Dules
(= [in part] Serranus of Robbins and Starck 1961). The
problem of the systematic position of Diplectrum and
Serranus has not been resolved here. Further studies
such as comparative osteology are needed on Serranus
and other Serraninae to resolve this question. However,
on the basis of present evidence, it is desirable to retain
Diplectrum and Serranus as distinct genera.

Status of the Subgenus Haliperca Gill

Gill (1863:263) designated S. biuittatus (= D. bivit-
tatum) as the type-species of his genus Haliperca. Later
Jordan et al. (1930:318-319) considered Haliperca as a
subgenus of Diplectrum to include radiale (= [in part]
D. bivittatum), D. pacificum, D. macropoma ( = [in
part] D. eumelum), D. sciurus, and D. euryplectrum.
Jordan et al. (1930:318-319) then limited the subgenus
Diplectrum to D. formosum.

Diplectrum formosum is an apomorphic species with
regard to its high scale meristics, the enlarged double
preopercular spur, and other characters and may repre-
sent a long-separated phylogenetic line (Bortone 1973);
relationships of the remaining Diplectrum are not easily
interpreted. Rather than retain Haliperca as a subgenus

for a group of uncertain lineage and relationships, or es-
tablish more subgeneric names of equally tenuous status,
the genus Haliperca is considered as a junior synonym of
Diplectrum. Subgenera are not utilized for any species or
species group at the present time. Further consideration
of subgeneric status for major lineages may be necessary
in the future when histological data on the gonads be-
come available.

Comments on the Definition of Serraninae

Katayama (1960:119-120) summarized and defined the
subfamily Serraninae based on the genus Chelidoperca
Boulenger 1895. The osteological features of Diplectrum
required modifications of Katayama's definition. Thus,
gill rakers for Serraninae must be considered short or
long, and in some species (e.g., D. sciurus) close together.
Anal rays for Serraninae are III, 7 or 8 and not simply III,
7. The suborbital shelf for Serraninae is developed on the
second, third, or fourth subocular bones and not always
on all three of these as Katayama (1960) stated.

Comments on the Definition of Serranidae

In his admirable 1966 work Gosline considered and
redefined the Serranidae, and although his definition is
reasonably complete and in general agreement with
specimens examined here, a comment is in order. Gos-
line (1966:120, table 1) listed characters which he believ-
ed would separate the well-defined Percichthyidae from
the Serranidae. He thought the Serranidae could be
defined by the presence of the supportive stay
(longitudinal ridge of Gosline) of the supraoccipital crest,
but in the present study D. formosum, D. bivittatum,
and Serranus phoebe lack the supportive supraoccipital
crest stay. This character, therefore, may not always lead
to the distinction of the Serranidae from the Per-
cicthyidae.

GONAD MORPHOLOGY

All Diplectrum examined in this study were "syn-
chronous hermaphrodites" as defined by Smith (1959,
1965) in a condition referred to as "territorial her-
maphroditism" by D'Ancona (1952). The gonads of D.
rostrum (Bortone 1974), D. formosum (Bortone 1971),
and D. pacificum (Bortone in press a) have been sec-
tioned and determined to be ovotestes. In these three
species, ovarian tissue appears yellow and granular, and
testicular tissue appears as fine white tissue. The
posteriormost region will be considered histologically in
other works (Bortone in press a) and the area is con-
sidered to be a region of possible accessory gland func-
tion. As a general structure it is termed an "ovarian
sinus." The structure is similar to the "oviducal pouch"
described by Smith and Atz (1969) for Pseudogramma
bermudensis but differs in many gross aspects. For in-
stance in Diplectrum the pouch does not open into the
body cavity. Smith (1965) used gross inspection to ex-

amine the pattern of hermaphroditism in serranid fishes.
This method is considered reliable for general mor-
phological purposes.

Schematic gonad diagrams (Fig. 8) suggest overall
similarity for the genus but also suggest certain mor-
phological affinities among species. In general the
anterior lobes of the U-shaped ovotestis are ovarian tis-
sues consisting of a central lumen which is surrounded by
lamelli. The testicular tissue is generally a wide, diagonal
band more posterior in orientation dorsally. The postero-
ventral portion of the ovotestis is composed of the "ovar-
ian sinus."

Morphologically there are several groups. One group
consisting of D. formosum, D. pacificum, D. radiale, D.
rostrum, D. euryplectrum, and D. eumelum (Fig. 8A, C,
D. E. G. H) shows a schematic pattern with regard to
shape and position of testicular and "ovarian sinus"
areas. Another group of species which are similar to
ovotestis morphology is D. sciurus and D. labarum (Fig.
8J, L). In these species the "ovarian sinus" portion is evi-
dent as in the first group but the testicular tissue is
somewhat extended into rounded anterior lobes which
begin to invade the ovarian tissue. Diplectrum macro-
poma (Fig. 81) is in a group by itself in that no "ovarian
sinus" region is visible grossly, yet the anterior testicular
lobes are present as in the second group. A fourth group
is composed of the remaining species: D. bivittatum, D.

conceptione, and D. maximum (Fig. 8B, F, K).
Testicular tissue in this group is present in very evident
anterior projections. Also the "ovarian sinus" region is
more ventral and slightly more anterior in position.

No apparent gross morphological differences were
found at the subspecific level between D. b. arcuarium
andD. b. bivittatum orD. f. formosum andD. f. radians.

KEY TO THE SPECIES OF DIPLECTRUM

(for specimens larger than 75 mm SL)

la Atlantic Ocean specimens 2

lb Pacific Ocean specimens 4

2a Preopercle with two clusters of spines radiat-
ing from upper and lower posterior corners
(Fig. 6A). Scales small; cheek scale rows A,
9-16 (modally 11); cheek scale rows B, 12-18
(modally 16); 8-12 scales above lateral line.
Pectoral rays modally 16. Spinous-dorsal-fin
profile nearly parallel to dorsum . . . D. formosum

2b Preopercle with a single cluster of spines
(Fig. 6B, D). Scales larger; cheek scale rows
A, 12 or less; cheek scale rows B, 14 or less.
Scales above lateral line, 9 or less. Pectoral

Figure 8. — Schematic gonad diagram of species of Diplectrum. Cross-hatched area, ovarian tissue; white area, testicular tissue; stippled area,
posterior ovarian-sinus region. Top of each paried figure is a left lateral view of the gonad. The bottom figure is a ventral view (the gonopore is
represented by a circle); the posterior portion of the gonad is to the right. A) D. formosum; B) D. bivittatum; C) D. pacificum; D) D. radiale; E) D.
rostrum; F) D. conceptione; G) D. euryplectrum; H) D. eumelum; I) D. macropoma; J) D. labarum; K) D. maximum; L) D. sciurus.

10

rays modally 15 or 17. Anterior dorsal fin
elevated not parallel to dorsal profile 3

3a Second dorsal spine short, 1.7-2.1 into third.
Preopercular spur square or rounded, its up-
per corner never pointed (Fig. 6D). Orbit
length small, 71-95%o SL. Pectoral rays
modally 17. Cheek scale rows A, 9-12 (modally
10). Upper caudal lobe without an extended
filament D. radiale

3b Second dorsal spine long, 1.2-1.6 into third.
Preopercular spur with a gentle curve to the
lower angle, its upper angle most often point-
ed (Fig. 6B). Orbit length large, 83-103%o
SL. Pectoral rays modally 15. Cheek scale
rows A, 7-9 (modally 8). Upper caudal lobe
often with an elongate filament . . . D. bivittatum

4a Total gill rakers 30 or more on first arch.
Maxillary width equal to, or greater than,
lacrimal width D. sciurus

4b Total gill rakers fewer than 30, maxillary

width less than lacrimal width 5

5a Anal rays 8. A dark lateral band of pigment
along soft-dorsal-fin base. Vertical preoper-
cular arm above spur not serrate . . D. euryplectrum

5b Anal rays 7 (except for D. labarum from out-
er Baja California in which 18^ have 8 anal
rays). No dark pigment present on soft-
dorsal-fin base. Vertical preopercular arm
serrate 6

6a Second, third, fourth, and occasionally fifth
dorsal spines with tips extended as long
black filaments (only on specimens larger
than 75 mm SL) D. labarum

6b Second, third, fourth, and fifth dorsal spines

with tips not extended 7

7a Second anal spine stronger than third and
long, 0.9-1.2 into third. Preopercle as an
elongate bony flap composed of thin spines
(or consisting of 2-4 strong spines), not as a
squared spur (Fig. 6F, K). Pelvic fins dusky ... .8

7b Second anal spine much shorter than third
(1.2 or more). Second anal spine not appreci-
ably stronger than third. Preopercle with a
definite squared spur and not as a thin spiny
flap nor as a cluster of 2-4 strong spines. Pel-
vic fins pale 9

9b Cheek scale rows A, 5-8. Cheek scale rows B,
7-10

11

8a Gill rakers 23-28. Preopercle with 2-4 very

strong spines at its angle (Fig. 6F) . . D. conceptione

8b Gill rakers 17-21. Preopercle spur as a bony
flap with 5-8 long thin spines at its angle
(Fig. 6K) D. maximum

9a Cheek scale rows A, 8-12. Cheek scale rows

B, 10-14 10

10a Snout with a distinct pale bar from the an-
teroventral corner of the eye, forward and
downward to the anteroventral edge of the
lacrimal. Soft dorsal fin with a distinct row
of pale spots on upper half of membrane on
a dark field. Gill rakers 22-25. Preopercular
spur spines 7-11 (Fig. 6E) D. rostrum

10b Snout with a vague spot, not as a well-de-
fined bar. Soft dorsal fin with 2-3 irregular
rows of pale spots on a dark field. Gill rakers
normally 15-22, occasionally 23 or 24. Pre-
opercular spur spines 5-8 (Fig. 6C) . . D. pacificum

11a Caudal peduncle depth 2.9-3.5 in head
length. Snout with 3-5 distinct pale spots.
Snout blunt. Length of posterior dorsal and
anal rays nearly equal to length of anterior
rays D. eumelum

lib Caudal peduncle depth 2.5-2.8 in head
length. Snout without distinct pale spots or
wavy cheek bar. Snout pointed. Length of
posterior dorsal and anal rays much shorter
than anterior rays D. macropoma

Diplectrum formosum (Linnaeus 1766)

Figures 1-5, 6, 7-8, 9, 10

Perca formosa Linnaeus 1766:488-489 (original descrip-
tion, Carolina); Gmelin 1788:1322 (copy of original
description); Linnaeus 1789:1322 (copy of original de-
scription); Cuvier and Valenciennes 1828:230 [in syn-
onomy with Haemulon formosum = Haemulon
plumieri (Lacepede)]; Goode and Bean 1886:203 (type
examined).

Serranus radians Quoy and Gaimard 1824:313-314, pi.
58, fig. 2 (original description, Brazil); Boulenger
1895:295-296 (catalog); Devincenzi 1924:223 (descrip-
tion).

Serranus fascicularis Valenciennes in Cuvier and Valen-
ciennes 1828:245-246 (original description, Brazil);
Cuvier 1829:139 (generic description); DeKay 1842:23
(description); Storer 1846:28-29 (general account);
Canestrini 1860:306 (systematics); Poey 1866:195
(Cuba); Jordan and Gilbert 1882:534 (not seen); Jor-
dan and Gilbert 1883:273 (Pensacola, Fla.); Goode
1884:410, pi. 163 (South Carolina); Goode 1887:46
(South Carolina); Diaz 1893:52 (synonomy).

Serranus irradians Valenciennes in Cuvier and Valen-
ciennes 1828:244-245, pi. 30 (in part, original descrip-
tion, Brazil); Castelnau 1855:3 (color notes, Rio de
Janeiro).

Diplectrum fasciculare Holbrook 1855:32-35, pi. 5, fig. 2
(generic description, South Carolina); Holbrook
1860:35-38, pi. 5, fig. 1 (South Carolina); Gill 1862:30
(southeast United States); Gill 1864:93 (name only);
Poey 1868:282 (generic designation); Poey 1871:54

11

A

'j?

***p«.

— ■■"

\.rk£

/*■

Figure 9.— A) Diplectrum formosum, 139 mm SL, (uncat.) northern Gulf of Mexico;

B) D. bivittatum, 117 mm SL, UMML 16302, Venezuela;

C) D. pacificum, 140 mm SL, USNM 183980, Gulf of California;

D) D. radiale, 107 mm SL, UMML 16302, Venezuela;

12

y

~**&.

Figure 9.— Continued.

E) D. rostrum, 131 mm SL, USNM 208197, Colombia;

F) D. conception^ 145 mm SL, USNM 213578, Peru;

G) D. euryplectrum, 63 mm SL, UMML 26251, Panama;
H) D. eumelum, 180 mm SL, USNM 213813, Colombia;

13

—

■

Figure 9. — Continued.

I) D. macropoma, 127 mm SL, USNM 213591, Colombia;
J) D. labarum, 135 mm SL, LACM 32094-1, outer Baja California;
K) D. maximum, 93 mm SL, SIO 64-63, outer Baja California;
L) D. sciurus, 109 mm SL, SIO 70-70, Gulf of California.

14

(generic designation); Gill 1873:28 (distribution);

Goode 1880:115 (Florida).
Centropristis fascicularis Giinther 1859:83 (catalog).
Centropristis radians Giinther 1859:83 (catalog).
Diplectrum radians Poey 1876:23 (synonomy); Diaz

1893:52 (name only).
Serranus formosus Jordan 1885:125 (color description,

Key West); Jordan 1887:82 (distribution, catalog);

Adams and Kendall 1891:292, 309 (Florida); Henshall

1891:387 '(Key West).

Diplectrum formosum Goode and Bean 1886:203 (ex-
amined type of Perca formosa); Jordan and Eigen-
mann 1890:397-398, pi. 65 (description and account);
Henshall 1895:215 (Key West); Jordan and Evermann
1896:1204, 1207-1208 (systematics, synonomy); Smith
1896:175 (Florida); Evermann and Kendall 1900:75
(Florida); Fowler 1903:329 (color notes, Florida); Jor-
dan and Thompson 1905:240-241 (color notes,
Florida); Fowler 1907:263 (Florida); Smith 1907:282
(North Carolina); Jordan and Evermann 1923:398-399
(general account); Meek and Hildebrand 1925:472-474
(key, description, and synonomy); Fowler 1927:252
(Florida); Fowler 1929:453 (Florida); Borodin 1934:113
(Florida Keys); Gunter 1935:39, 40 (Louisiana); Bere
1936:608 (parasites); Fowler 1940a:14, fig. 20 (Florida)
Fowler 1941a:770 (Rio de Janeiro); Fowler 1941b:86
(Florida); Galloway 1941:119 (lethal temperature)
Longley and Hildebrand 1941:103-104 (life history
Florida); Fowler 1942:158 (Brazil); Baughman 1944:89
90 (Texas); Fowler 1945a:200, 310 (synonomy
southeast United States); Breder 1948:166 (general ac-
count); La Monte 1952:104 (popular account); Out-
door Writers Association of America 1952 (check list);
Pearse 1952a (parasites); Hildebrand 1954:304
(Campeche); Hildebrand 1955:208, 221-223, tables 2-4
(Campeche); Joseph and Yerger 1956 (Florida); Briggs
1958:272 (distribution); Hoese 1958:331 (Texas); Mig-
dalski 1958:355 (popular account); Robins 1958 (check
list, Florida); Bailey et al. 1960:25 (check list);
Springer and Woodburn 1960:34 (ecological data,
Florida); Clem et al. 1961:726-766 (tissue culture);
Durand 1961:46, 50 (ecological data, Guyana); Smith
and Bailey 1961:361 (osteology); Tabb and Manning
1961:618 (Florida); Lowe 1962:695 (Guyana); Freihofer
1963:131, table 2 (neuroanatomy); Gabrielson and La
Monte 1963:51 (popular account); Brandao 1964:19
(Brazil); Tyler 1964:9 (Florida); Anderson and Geh-
ringer 1965:table 12 (collection data, Florida); Bullis
and Thompson 1965:44 (collection and distribution
data); Christensen 1965 (Florida); Gunter and Hall
1965:31 (collection data, Florida); Moe and Martin
1965:138 (biological data, Florida); Roessler 1965:314,
table 2 (ecological data, Florida); Roithmayr 1965:20,
table A-l (northern Gulf of Mexico); Cervigon
1966:321-323 (description, biology, Venezuela); Moe
1966:11, table 2 (Florida); Moe et al. 1966:32-33
(Florida); Starck and Davis 1966:320 (Florida); Ander-
son 1968:1-60 (trawl data from S. Atlantic coast of
United States); Beaumariage 1968:8 (utilized for fish

meal, Florida); Bohlke and Chaplin 1968:267
(Bahamas); Randall 1968:76, fig. 87 (general account,
Caribbean); Starck 1968:19 (Florida); Wolf and Quim-
bey 1969:298, table 8 (tissue culture); Bailey et al.
1970:34 (check list); Bortone 1970:1-55 (life history,
Florida); Godcharles 1970:11 (Florida); Mago Leccia
1970:94 (Venezuela); Anonymous 1971:61-62 (popular
account); Bearden and McKenzie 1971:13 (collection
data, South Carolina); Bortone 1971a:l-27 (life his-
tory, Florida); Bortone 1971b:28 (behavior and color
notes, Florida); Dahl 1971:224, fig. 295 (Colombia);
Ewald et al. 1971:54 (Venezuela); Godcharles 1971:34,
table 8 (collection data, Florida); Grimes and Moun-
tain 1971:40-41 (Florida); Nahhas and Powell 1971:4
(parasites); Smith 1971:108 (synonomy); Swingle
1971:33 (Alabama); Bortone 1972:55 (life history,
North Carolina); Mountain 1972:60 (collection data,
Florida); Gallaway et al. 1972:89, 91 (key); Miller and
Jorgenson 1973:310 (meristics); Cliburn 1973:18 (key);
Dahlberg 1975:58, fig. 117 (Georgia); Walls 1975:169
(northern Gulf of Mexico).

Centropristis formosus Berg 1895:45-46 (synonomy, Uru-
guay).

Haliperca formosa Miranda-Ribeiro 1904:25 (Brazil);
Miranda-Ribeiro 1915:29 (Brazil); Miranda-Ribeiro
1918:92-93 (Brazil).

Diplectrum formosus Jordan et al. 1930:318-319 (check
list); Gunter 1941:195 (Louisiana); Baughman
1950:249 (Texas); Buller 1951:15 (southeast United
States); Reid 1954:37-38 (life history, Florida).

Diplectrum fascicularis Poey 1955:186-187, pi. 46 (de-
scription, Cuba).

Diplectrum radiate Ringuelet and Aramburu 1960:36, 62
(misidentification).

I have not seen the type-specimen of D. formosum, but
I accept the authority of Goode and Bean (1886:203) who
examined the type at the Linnaean Society of London in
Burlington House. They confirmed that the Linnaean
type was what had been referred to on the eastern coast
of the United States as D. fasciculare (= S. fascicularis
Valenciennes in Cuvier and Valenciennes 1828:245-246).

Valenciennes (in Cuvier and Valenciennes 1828) also
described Serranus irradians and although the descrip-
tion is of D. formosum the syntypes are of two species
(i.e., MNHN A464, A461, 7018 are D. formosum; the
other syntype MNHN 462 is D. radiale). MNHN A464 is
designated the lectotype for Serranus irradians.

Diplectrum radians (Quoy and Gaimard 1824:313-314,
pi. 58, fig. 2) represents the first description of D. for-
mosum from Brazil.

Diagnosis. — As an adult, D. formosum is easily recog-
nized because it is the only Diplectrum whose preopercle
has two centers of diverging spines. These form a double
pointed spur consisting of upper and lower projections.
Diplectrum formosum modally has 16 pectoral rays while
D. bivittatum and D. radiale have 15 to 17 each respec-
tively (Table 1). Also D. formosum possesses small
scales. This is reflected in the species having the highest

15

Table 1. — Number of pectoral rays in Diplectrum species.

Species

14

15

16

17

18

19

X

s

n

D. form os um

22

236

8

15.95

0.32

266

D. biuittatum

Gulf of Mexico

3

131

28

15.16

0.41

162

S. Florida

1

28

20

15.39

0.53

49

Caribbean and

South America

3

145

129

15.45

0.52

277

Total

7

304

177

15.35

0.51

488

D. radiate

25

73

4

16.79

0.49

102

D. pacificism

11

101

14

17.02

0.44

129

D. rostrum

3

29

1

16.94

0.35

33

D. conceptione

4

38

13

18.16

0.54

55

D. maximum

1

14

16.93

0.26

15

D. sciurus

4

67

1

16.96

0.26

72

D. eunplectrum

2

47

85

17.62

0.52

134

D. labarum

Outer Baja California

6

17.00

6

Gulf of California

28

9

16.24

0.43

37

Panama and Costa R

ca

2

28

1

16.97

0.31

31

Total

30

43

1

16.61

0.52

74

D. eumelum

1

95

3

17.02

0.20

99

D. macropoma

3

130

21

17.12

0.38

154

mean scale numbers (Tables 2-7) of any Diplectrum. Ju-
venile D. formosum may be confused in general appear-
ance with other Atlantic Diplectrum but eye size (small
in D. formosum) and scale counts will separate the spe-
cies. As an adult, D. formosum has the broadest lacrimal
and narrowest maxillary bones.

Description. — Within D. formosum there are two
morphological populations, a northern subspecies from
coastal Gulf of Mexico and Atlantic North America and a
southern subspecies from coastal Caribbean and Atlan-
tic South America. The two forms are described
separately for morphometric characters. Meristic charac-
ter differences for both forms except gill raker counts are
slight; therefore, these statistics are considered the same
for both.

D, X. 12 (13); A, III, 7(6-8); P, , 15.95 (15-17); pored
lateral-line scales, 49.76 (46-55); scale rows in lateral
line, 76.78 (66-88); scales above lateral line, 9.38 (8-12);
scale rows below lateral line, 21.36 (18-26); predorsal
scale rows, 17.20 (14-22); cheek scale rows A, 11.77 (9-
16); cheek scale rows B, 15.60 (12-18).

Northern form.— Gill rakers, 18.83 (17-22). Standard
length, 140.4 (36.7, 77.6-220, 49). Total length, 1,340
(50.12, 1241-1451, 44); fork length, 1,191 (22.79, 1,136-
1,308, 49); body depth, 275 (15.09, 250-318, 45); head
length, 359 (12.58, 330-390, 49); postorbital length, 191
(8.34, 176-210, 49); snout length, 113 (8.77, 93-130, 48);
lacrimal width, 59 (6.76, 45-70, 39); maxillary width, 26
(2.05, 22-29, 39); orbit length, 74 (6.42, 62-89, 49); orbit
width, 59 (5.76, 48-72, 49); interorbital width, 74 (4.42,
64-82, 38); upper jaw length, 160 (7.10, 145-175, 49);
lower jaw length, 179 (8.12, 154-198, 49); cheek height,
125 (9.85, 104-143, 49); cheek length, 128 (7.39, 115-145,
49); caudal peduncle depth, 134 (4.86, 122-146, 49); pec-
toral fin length, 235 (10.82, 207-257, 49); pelvic fin

length, 216 (10.53, 185-238, 39); predorsal length, 349
(10.62, 321-368, 39); pelvic origin to lower jaw, 341 (13.29,
313-379, 32); pelvic origin to anus, 306 (24.09, 254-357,
49); postanal length, 402 (13.90, 374-428, 48). Dorsal
spine height: first, 51 (6.40, 35-68, 46); second, 78 (7.19,
58-90, 48); third, 95 (7.36, 79-108, 45); fourth, 105 (8.89,
84-125, 46); fifth, 110 (8.58, 91-131, 43). Anal spine
height: first, 25 (4.41, 16-37, 46); second, 47 (6.80, 35-65,
48); third, 70 (8.37, 50-92, 46).

Southern form.— Mean gill rakers, 22.01 (19-24). Stan-
dard length, 119.1 (28.1, 76-167, 50). Total length, 1,269
(17.77, 1,232-1,312, 46); fork length, 1,183 (12.80, 1,149-
1,207, 50); body depth, 270 (16.06, 238-299, 50); head
length, 362 (15.31, 329-394, 48); postorbital length, 192
(9.66, 173-214, 50); snout length, 101 (7.89, 84-121, 50);
lacrimal width, 52 (5.59, 41-62, 36); maxillary width, 27
(2.09, 21-31, 37); orbit length, 83 (6.14, 71-94, 50); orbit
width, 63 (5.71, 54-78, 50); interorbital width, 73 (5.18,
63-83, 47); upper jaw length, 160 (7.22, 141-174, 50);
lower jaw length, 181 (7.00, 165-195, 50); cheek height,
119 (8.97, 98-135, 50); cheek length, 129 (6.50, 116-144,
50); caudal peduncle depth, 125 (5.28, 111-137, 50); pec-
toral fin length, 238 (7.81, 220-263, 50); pelvic fin length,
210 (9.57, 182-232, 49); predorsal length, 350 (10.20, 335-
370, 37); pelvic origin to lower jaw, 342 (16.66, 318-382,
50); pelvic origin to anus, 314 (20.07, 276-353, 50);
postanal length, 401 (312, 371-422, 50). Dorsal spine
height: first, 53 (5.08, 37-66, 50); second, 79 (6.34, 61-90,
49); third, 100 (7.93, 82-114, 49); fourth, 110 (8.09, 91-
125, 46); fifth, 113 (8.29, 94-133, 49). Anal spine height:
first, 29 (3.55, 20-35, 49); second, 53 (7.84, 40-78, 49);
third, 74 (7.67, 60-93, 50).

Preopercular spur with two clusters of diverging
spines. The number of spur spines variable, from 6 to 15,
but more often from 9 to 12 between upper and lower
clustered tips. The southern form with slightly more

16

Table 2. — Number of scale rows in lateral line of Diplectrum species and populations.

Species

48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70

D. formosum
D. bivittatum
D. radiate
D. pacificum

Gulf of California

Mexican coast and
Costa Rica

Panama and Colombia
Total
D. rostrum

Northern

Southern
Total
D. conceptione
D. maximum
D. sciurus
D. euryplectrum
D. labarum

Gulf of California

Panama and Costa Rica
Total
D. eumelum
D. macropoma

1

1

2

3

1

2

1

2

9

16

30

:■'

53

51

49

51

46

37

26

9

1

1

1

2

7

7

5

10
1

16
1

1 1
2

10

1

12
6

6

5

6
8

1

10

3

1

1

1

4

2

2

6

2

2

1

2

6

3

5

8

9

4

6

3

5

2

1

7
2

3
3

6

3
1

13
4

15

5
2

10

3
1

14

1

1 1

15

3

12

1
3

4
2

2

3
2

1

1
1

4

2

1

7
3
3
4

4
2
1
2

1
1

2

7
1
4

3
5
3

4

4

8

■j

7

2
6

1
8

2

1

4

3

6

14

15

Id

19

12

9

13

9

6

4

1

1

4

3

2

5

1

6

4

3

7

1

1 2

6

2

5

3

4

2

2

2

2

1 2

6

2

6

4

8

5

4

7

t

6

6

3

7

1

1 3

4

5

15

9

16

12

U

7

9

2

4

2

12 22 24 21 19 15

Species

71

74 75 76 77 78 79 80 81 82 83 84 85 86 87

D. formosum
D. bivittatum
D. radiate
D. pacificum

Gulf of California

Mexican coast and
Costa Rica

Panama and Colombia
Total
D. rostrum

Northern

Southern
Total
D. conceptione
D. maximum
D. sciurus
D. euryplectrum
D. labarum

Gulf of California

Panama and Costa Rica
Total
D. eumelum
D. macropoma

9 20 19 29 20 20 23 22 29 13 19 14

5 1

1

76.78

3.82

259

64.69

2.95

126

63.36

3.07

100

1

1 1

1 1

67.58

5 4
1

5 6

1 1

2.40

43

65.40

2.72

25

63.79

2.82

52

65.48

3.13

120

63.10

2.07

21

67.08

2.60

13

64.62

2.98

34

68.82

3.34

55

66.33

3.13

15

70.76

3.60

68

64.12

3.39

130

61.27

2.95

41

55.87

3.07

31

58.94

4.02

72

57.42

3.01

104

53.86

2.70

147

Table 3. — Scales above lateral line of Diplectrum species.

Species

10 11

12

D. formosum
D. bivittatum
D. radiate
D. pacificum
D. rostrum
D. conceptione
D. maximum
D. sciurus
D. euryplectrum
D. labarum
D. eumelum
D. macropoma

17 81 49 11

18

150

76

6

9

55

33

2

1

52

51

12

16

16

1

10

32

6

1

8

4

2

14

32

24

1

50

61

12

1

11

30

8

1

3

(in

36

2

1

9

73

51

2 9.38

0.81

160

7.28

0.63

250

7.28

0.66

99

7.64

0.68

116

7.54

0.56

33

7.92

0.58

48

8.47

0.83

15

8.17

0.76

71

7.71

0.67

124

6.98

0.68

50

6.39

0.63

102

6.32

0.60

133

17

Table 4.— Scales below lateral line of Diplectrum species.

Species

11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

D. formosum
D. bicittatum
D. radiate
D. pacificum
D. rostrum
D. conceptione
D. maximum
D. sciurus
D. euryplectrum
D. labarum
D. eumelum
D. macropoma

2 18 31 32 32 33

6

34

74

66

46

15

3

1

3

13

29

35

13

6

2

6

28

45

25

4

5

8

10

Id

1

2

17

19

13

2

1

1

1

4

2

5

1

1

IS

25

21

6

4

10

34

34

28

8

2

4

10

12

13

4

1

9

38

42

12

9

38

55

17

4

1

2 21.36 1.60 157

18.69

1.25

244

18.57

1.20

100

18.88

1.00

110

16.76

1.06

33

19.93

1.09

55

21.13

1.73

15

19.18

0.96

71

17.81

1.19

118

16.93

1.29

45

15.54

0.85

102

14.60

1.15

132

Table 5. — Number of predorsal scale rows of Diplectrum species and populations.

Species

10 11 12 13 14 15 16 17 18 19 20 21 22

D. formosum

4

IS

34

27

34

is

D. bivittatum

Gulf of Mexico

1

13

31

48

14

9

S. Florida

2

4

5

3

4

3

Carribbean and

South America

10

28

42

41

13

2

Total

3

17

46

79

60

53

13

2

D. radiate

2

6

19

32

27

8

5

1

D. pacificum

1

4

17

28

22

24

6

6

2

D. rostrum

2

9

10

4

7

D. conceptione

11

9

16

11

1

3

2

D. maximum

1

2

6

3

1

D. sciurus

2

2

12

11

11

2

9

1

D. euryplectrum

Mexico

1

4

10

11

6

1

1

Panama

1

2

10

8

10

11

3

4

Colombia

2

8

11

11

Galapagos

s.

1

1

Total

2

10

17

31

19

17

13

4

4

D. labarum

1

5

19

6

7

3

1

D. eumelum

18

:«;

30

8

3

2

D. macropoma

Gulf of California

1

2

9

13

4

1

1

Panama

1

3

7

9

13

12

8

2

( 'oltimhia

1

5

11

14

14

3

Total

1

7

16

30

36

20

13

8

3

1 17.20

1.64

147

12.76

1.08

116

12.57

1.60

21

14.18

1.13

136

13.45

1.36

273

15.25

1.33

100

14.85

1.60

110

14.16

1.25

32

13.98

1.56

53

16.43

1.55

14

15.48

1.70

50

13.70

1.24

34

14.82

1.70

4d

11.97

0.93

32

15.50

2

13.73

1.81

117

13.62

1.30

42

11.46

1.11

97

13.81

1.28

31

14.96

1.62

55

12.92

1.18

48

13.96

1.66

134

Table 6. — Number of cheek scale rows A of Diplectrum species.

Species

10 11 12 13 14 15 16

D. formosum
D. bicittatum
D- radiate
D. pacificum
D. rostrum
D conceptione
D. maximum
D. sciurus
D. euryplectrum
D- labarum
D. eumelum
D. macropoma

10

1 1

1 19 86 40 40 10

68

305

97

24

62

13

3

2

42

70

7

1

17

15

1

6

24

17

8

2

10

3

24

41

4

23

87

16

3

42

27

1

11.77 1.24 202

93 52 1

8.06

0.59

470

9.95

0.69

102

9.70

0.64

122

8.52

0.56

33

8.49

0.88

55

0.07

0.59

15

7.66

0.63

71

7.97

0.68

130

7.34

0.60

74

5.98

0.42

102

6.27

0.59

157

L8

Table 7.— Number of cheek scale rows B of Diplectrum species.

Species

10 11 IS

13 14 15 16 17 18

D. formosum
D. bivittatum
D. radiale
D. pacificum
D. rostrum
D. conceptione
D. maximum
D. sciurus
D. euryplectrum
D. labarum
D. eumelum
D. macropoma

6 36 67 104 43

1

4

140

225

81

4

5

28

54

9

1

5

10

68

39

4

1

25

7

1

6

is

22

5

3

1

7

6

12

1
29

20

3

in

51

55

11

1

1

L9

20

4

7

51

39

5

3

62

65

5

in

15.60

1.08

263

11.86

0.75

455

11.73

0.78

97

12.21

0.80

126

11.18

0.46

33

11.60

1.05

55

11.47

0.74

15

13.44

1.02

72

10.49

0.87

131

10.61

0.69

44

8.41

0.69

102

8.53

0.61

135

elongate spines tending to diverge earlier during on-
togeny at about 75 mm SL as opposed to 90 mm SL for
the northern form. Horizontal and vertical preopercular
arms serrate. Gill rakers of moderate length, the first at
the angle reaching to the third below in the northern
form and to the fourth below in the southern form. An-
terior nostril with short flap. Anterior chest scales em-
bedded and slightly smaller than lateral body scales.
Caudal fin slightly forked in the northern form, the upper
lobe nearly always extended as an elongate filament.
Southern form with caudal fin nearly truncate, upper
lobe never extended as an elongate filament.

Color. — Color notes are here presented for live specimens
of the northern form of D. formosum from North
Carolina and Florida. Dorsum light brown, lateral sur-
face tan, ventral surface white. Lateral body surface in
juveniles with two dark brown stripes. In adults these
stripes interrupted and appear as 5-7 vertical bars.
Caudal spot distinct, equal to eye size, and present on
upper half of caudal peduncle. Anal fin with pale yellow
spots, pectoral and pelvic fins pale. Midlateral ventral
area pink-orange in adults. Upper pharyngeal region yel-
low with blue flecks of pigment in adults. Posterior
branchial cavity pale. Upper medial opercular surface
with small patch of dark gray pigment, lower medial
opercular surface pale white.

Snout with two bright blue lines from anterior lac-
rimal edge to orbit with bright blue irregular lines below
joining to form a single line running irregularly across up-
per cheek. Three or four more blue lines on cheek,
preopercle, and opercle. Maxillary with blue spots.
Lower jaw bluish purple. Throat white, a bluish-purple
blotch ventral and posterior to lower jaw symphysis. Top
of head brown with four to five bluish-green lines. Seven
to nine longitudinal blue lines trimmed with yellow along
sides of body. These bright blue above and paler ven-
trally. Anterior dorsal fin with two longitudinal blue lines
on an orange field. These blue lines irregular anteriorly,
developing into three posteriorly, and into four lines on
the soft dorsal. Tips of spines bright red-orange. Caudal
fin with numerous orange spots or rows of fused spots or
vertical lines on a pale blue field. Extended caudal fila-
ment orange. All colors fading with preservation. Orange
and yellow becoming pale and blue becoming dusky.

In preservative the following differences between the
northern and southern forms are noted: in the southern
form lines on cheek and snout more wavy; dark brown
vertical body bars broader, more intense, extending fur-
ther ventrally; caudal spot larger than eye, extending
below midcaudal peduncle line and vertical in orien-
tation; blue lines on top of head and seven to nine body
stripes not discernible; base of pectoral fin brown, not
pale as in the northern form; blue dorsal fin lines tending
to be more irregular, terminating in three, rather than
four, stripes; caudal fin spots more irregular; definite
dark brown spot dorsally on caudal peduncle.

Size. — The species attains a moderate size and 224 mm
SL is the maximum size known.

Habits. — Diplectrum formosum is found in coastal areas
over sand bottom at the base of reefs and upper edges of
depressions in the substrate (Bortone 1971a). The species
is recorded from depths of 1-80 m but is most often found
at depths of 7-50 m. Larger adults tend to be found in
deeper water while juveniles are more commonly taken
from shallow water (Bortone 1971a).

Distribution. — Diplectrum formosum is a continental
species found throughout the Gulf of Mexico, around
peninsular Florida, and northward along the east coast of
the United States to Chincoteague, Va. A record from a
New York market (USNM 15844) is not confirmed to be
from New York and most probably came from the North
Carolina area as part of an exported fish catch. The
species has been recorded from the Bahamas (Bohlke
and Chaplin 1968:267). This is the only plotted literature
record (Fig. 10) not verified in the present study. One
specimen is from the Virgin Islands (FSBC 5393). The
species is also found along the northern coast of South
America, and southward to about Sao Paulo, Brazil.
Literature accounts (e.g., Berg 1895:45-46; Cuvier and
Valenciennes 1828:244-245) of the species occurring
south to Uruguay cannot be confirmed at this time.

The northern form would include North American
specimens examined. The southern form includes
specimens from South America and the Virgin Islands.
The type-locality is the Carolinas.

19

Figure 10.— Distribution map for Diplectrum for-
mosum (circles); D. aciurus (triangles); and D. con-
ceptione (squares). [Localities are based on the
materials examined in this study. 1

Variation. — Among meristic characters the number of
gill rakers shows the most noticeable variation (Table 8).
The northern form shows a mean of 18.83. There is slight
variation at different localities with North Carolina
specimens having a mean of 18.18, those in the Florida
Keys with 19.31, and those off Yucatan with 19.80. This
slight variation may be temperature induced. In the dis-
tribution of the species a sharp separation between
southern and northern forms is also shown in mean gill
raker count; from Venezuela southward higher mean
counts increase from 21.67 to 22.48 in Brazil.

The northern subspecies has a considerably greater
total length than the southern form, because of the
elongate filamentous upper caudal lobe of the former.
Snout length tends to be greater for the northern form,
and eye size is generally larger in the southern form. All
dorsal and anal spines measured tend to be slightly
longer for the southern form. Color descriptions
presented herein reflect discernible differences between
the two forms. It cannot be established what sig-
nificance the differences may have, however, until live
southern form D. formosum are examined. The more
exaggerated and earlier formed double-spur preopercle of
the southern form D. formosum is also an important
character for separating the subspecies.

Systematic comments. — The differences as well as
similarities of the two forms are readily apparent. The
question of separation of the two forms into different
species seems at this point somewhat arbitrary in view of
the obvious overlap in characters and lack of live
specimens for color comparisons. It does seem, however,

that the significance of these differences should be em-
phasized and recognized. Therefore it is recommended
that the two forms be considered as subspecies: D. for-
mosum formosum (Linnaeus 1766), the northern form,
and D. formosum radians (Quoy and Gaimard 1824), the
southern form.

Specimens examined. — A total of 352 specimens from 29

to 224 mm SL. NEWYORK: USNM 15844, 1 (145). VIRGINIA:
USNM 14.3388, 1 (120). CAROLINAS: UNC 2687, 2 (192-200); UNC 194,
1 (82); UNC 6058, 27 (150-210); BMNH 207, 2 (167-224); GMBL 70-113. 2
(187-109); GMBL 67-1, 1 (125); MCZ 10255. 1 (196); MCZ 2860, 1 (178);
MCZ 10254, 1 (202). JACKSONVILLE, FLA.: UNC 5225, 27 (68-175);
SIO 65-522, 1 (167). S. FLORIDA, KEYS AND CUBA: USNM 4676, 2
(100-151); USNM 13057, 1 (153); UMML 2081, 4 (55-104); UMML7778, 1
(83); UMML 17409, 1 (136); UMML 5195, 1 (57); FMNH 46165, 2 (165-
168); UMML 3223, 15 (49-147); MCZ 45033, 1 (184); TABL (Key Bis-
cayne, 13 Dec. 1967), 4 (55-72); CAS SU1422, 6 (65-169); FSBC 336, 5 (57-
115); MCZ 2932. 1 (113); MCZ 2929, 1 (105); MCZ 21768, 1 (164); MCZ
21766, 1 (155); MCZ 21767, 1 (145); MCZ 21765, 1 (155); MCZ 38654, 1
(183); MCZ 2863, 2 (143-178); MCZ 46663, 1 (155); UMML 16240, 1
(80). EASTERN GULF OF MEXICO: UMML 1814, 1 (92); FMNH
33952, 1 (94); FMNH 33953, 1 (88); FSBC 2955, 7 (75-155); FSBC 4667, 7
(94-150); FSBC 6376, 17 (42-88); MCZ 45116, 1 (147); TABL (Sanibel Is.,
14 Aug. 1959), 1 (68); MCZ 38656, 1 (197). NORTHERN GULF OF
MEXICO: USNM 142852, 8 (29-45); FSU 17893, 4 (41-65); FSU 17890, 3
(81-95); FSU 14724, 1 (100); FSU 269, 2 (71-78); FSU 18508, 35 (33-93)
MCZ 35111. 3 (184-198); UW 16762, 1 (124); UW 16648, 4 (95-116)
USNM 155208, 2 (135-138); GCRL 812, 3 (94-107); GCRL 1213, 1 (137)
GCRL1444, 1 (72). TEXAS: USNM 143389, 1 (124). MEXICO: USNM
134192, 1 (167); FMNH 45476, 2 (155-162); MCZ 38, 1 (177); INPLB776,
1 (140); GCRL 2981, 2 (102-107); UTIMS 923, 1 (155); UTIMS 924, 4 (66-
130); UTIMS 919, 1 (124); UTIMS 916, 2 (51-84). VIRGIN ISLANDS:
FSBC 5393, 1 (62). COLOMBIA: USNM 211406, 3 (150-157); USNM
211403, 3 (145-148). VENEZUELA: UNC (Cubagua, 11 Oct. 1971), 9
(109-167); MCZ 40942, 7 (52-92); MCZ 41088, 1 (160); UPR 2665. 3 (108-
144); USNM (lat. 10°30'N, long. 70°30'W, 15 Nov. 1958), 5 (72-104);

20

Table 8. — Total number of gill rakers of Diplectrum species and populations.

2

29
19

5
9

2
2

1

10

11
21

2

1

22
18
25

5
5

12
6

7

7

5

1

2
2

3

93

89

36

6

4

1

6
5

32
11

31
11

28
9

10

1

1

11
10

9

11

7
12

1
9

3

93

90

42

38

35

28

10

7

54

70

40

15

1

6

19

24

12

1

1

2

28

102

118

38

8

5

1

2

29

108

144

L16

90

46

15

2

23

46

18

12

5

1

5

28

75

56

8
1

2

21

5

2
5

H

1

3

2

7

1
3

20

58

70

6

17

42

58

20

2

6

47

38

11

1

1

13

53

71

30

11

3

2

Species 15 16 17 18 19 20 21 22 23 24 25 26 27 28

D. formosum

North Carolina

Jacksonville

S. Florida

W. Florida

N. Florida

Mississippi, Loui-
siana, Texas

Yucatan

Northern total

Southern total

Venezuela

Guyana

Brazil
Total
D. bivittatum

Gulf of Mexico

S. Florida

Caribbean and
South America
Total
D. radiale 1

D. pacificum
D. rostrum

D. conceptione 5 6 23 13

D. maximum
D. sciurus

D. euryplectrum 20

D labarum
D. eumelum
D macropoma

Species 32 33 34 35 36 37 38 39 40 41

D. formosum

North Carolina

Jacksonville

S. Florida

W. Florida

N. Florida

Mississippi. Loui-
siana. Texas

Yucatan

Northern total

Southern total

Venezuela

Guyana

Brazil
Total
D. bivittatum

Gulf of Mexico

S. Florida

Caribbean and
South America
Total
D. radiale
D. pacificum
D. rostrum
D. conceptione
D maximum

D. sciurus 1 3 3 28 33 42 26 12 1 2

D. euryplectrum
D. labarum
D. eumelum
D macropoma

18.18

0.61

38

18.43

0.63

30

19.31

1.08

52

18.86

0.69

35

18.74

0.68

53

19.00

0.92

8

19.80

1.08

15

18.83

0.92

231

22.01

0.74

108

21.67

1.01

36

21.77

1.07

30

22.48

1.09

42

19.74

1.68

339

22.03

1.01

187

20.68

1.00

63

19.67

1.05

302

20.58

1.49

552

18.25

1.17

107

20.25

1.02

177

23.44

0.80

32

25.28

1.17

56

19.28

1.14

1 I

36.64

1.51

151

23.68

1.06

190

21.52

1.05

145

19.55

0.80

103

19.93

1.16

184

21

BMNH 1904-3.15:26-7. 1 (163); RMNH 23349, 2 (162-170). CURACAO:
RMNH 22193. 1 (188); RMNH 22194. 1 (177). GUYANA: BMNH 1961-
8.31:122-127. 3 (129-150): FMNH 64975. 5 (80-143); USNM 15681, 3 (128-
1541. SURINAM: USNM 1S5497. 2 (142-151); USNM 185306, 7 (115-
135). FRKNCH GUIANA. FMNH 64966. 2 (130-146); UMML 11593, 1
il3D. BRAZIL: MCZ 2864. 5 (142-153); USNM 185028, 2 (44-66);
MZUSP 9059. 1 (117); MZUSP 9055. 1 (125); MZUSP 8989, 1 (114);
MZUSP 9060. 1 (144); MZUSP 9058, 1 (127); MZUSP 9057, 1 (99);
MZUSP 9056. 1 (119): MNHN 7022. 1 (133); MNHN A464. lectotype, 1
IS MNHN A461. 1 (123): MNHN 7018, 1 (104); MCZ 44166, 2 (105-
16); USNM 1567S0, 5 (101-140); USNM (Oregon 4241), 2 (121-141);
I SNM 100853. 1 (123); MZUSP 9066, 1 (80); MZUSP 9067, 1 (80);
MZUSP 906S. 1 (69); MZUSP 9069, 1 (67); MZUSP 9070, 1 (76); MCZ
10245. 9 (83-131); MZUSP 9061, 1 (97); MZUSP 9062, 1 (94); MZUSP
906:;. 1 (92); MZUSP 9064. 1 (77); MZUSP 9065, 1 (87); UMML 115411, 2
1120-123): BMNH 1923-7.30:80-82. 1 (129).

Diplectrum bivittatum (Valenciennes 1828)

Figures 6, 8, 9, 11

Serranus bivittatus Valenciennes in Cuvier and Valen-
ciennes 1828:241-242 (original description, Mar-
tinique and Havana); Storer 1846:27 (general ac-
count); Gill 1863:236 (generic affinity with Haliperca);
Poey 1866:185 (Cuba); Poey 1868:282 (description,
Cuba); Boulenger 1895:297 (catalog).

Centropristis bivittatus Gtinther 1859:82 (catalog).

Haliperca bivittatus Poey 1871:52-54 (systematics).

Serranus radialis Jordan 1886:376 (in part, dis-
tribution); Jordan 1887:82 (catalog, West Indies);
Metzelaar 1919:54-55 (synonomy, West Indies).

Diplectrum radiale Jordan and Eigenmann 1890:398-399
(in part, systematics); Jordan and Evermann
1896:1203-1205 (in part, systematics); Evermann and
Marsh 1902:159 (Puerto Rico); Fowler 1907:263 (in
part, distribution); Meek and Hildebrand 1925:472,
475-477, pi. 46, fig. 1 (systematics, Panama); Nichols
1929:255 (Puerto Rico and Virgin Islands); Jordan et
al. 1930:318 (in part, check list); Longley 1932:286
(biology, Florida); Beebe and Tee-Van 1933:135-136
(description, Bermuda); Baughman 1950:249 (Texas);
Briggs 1958:272 (in part, distribution); Caldwell and
Caldwell 1964:16 (Colombia and Aruba); Randall
1968:73, 76, fig. 86 (description, Caribbean); Anony-
mous 1971:63-65 (popular account); Dahl 1971:225
(Colombia); Dahlberg 1975:58 (Georgia).

Haliperca bivitata Diaz 1893:52 (name only, mis-
spelled).

Diplectrum bivittatum Longley and Hildebrand
1941:104-105 (field data, Florida); Ginsburg 1948:266-
270 (systematics); Hildebrand 1955:208, 222-223, table
7 (systematics, Campeche); Bailey et al. 1960:25
(check list); Bullis and Thompson 1965:44 (field data);
Roessler 1965:314, table 2 (field data, Florida); Cald-
well 1966:40 (Jamaica); Moe et al. 1966:32 (Florida);
Starck and Davis 1966:320 (Florida); Starck 1968:19
(Florida); Bailey et al. 1970:34 (check list); Gallaway
et al. 1972:89 (key); Miller and Jorgenson 1973:310
(meristics); Dahlberg 1975:55 (Georgia); Walls
1975:168-169 (northern Gulf of Mexico).

Diplectrum arcuarium Ginsburg 1948:266-270 (original

description. Gulf of Mexico); Baughman 1950:249
(Texas); Pearse 1952b (parasites); Hildebrand
1954:304 (Texas); Hildebrand 1955:224, table 7 (sys-
tematics, Gulf of Mexico); Briggs 1958:272 (dis-
tribution); Hoese 1958:331 (Texas); Bullis and
Thompson 1965:44 (field data); Gunter and Hall
1965:31 (field data, Florida); Cervigon 1966:324 (name
only); Swingle 1971:33 (Alabama); Franks et al.
1972:84 (Mississippi).

Diplectrum spp. Lowe 1962:695 (in part, Guyana).

Diplectrum bivitatum Cervigon 1966:324 (name only,
misspelled).

The species was described originally as Serranus bivit-
tatus by Valenciennes in Cuvier and Valenciennes
(1828:241-242) from two specimens (syntypes). In the ab-
sence of any designated holotype, MNHN 6988 (Mar-
tinique) is designated the lectotype. MNHN 5889
(Havana, Cuba) was not chosen as the lectotype as it is
from the area of the intermediate population (see below)
and is here considered the only paralectotype.

Early confusion in the literature regarding D. bivit-
tatum probably was due to inclusion of its specific name
under the synonomy of D. radiale by Jordan and Eigen-
mann ( 1890:399) because they considered Serranus bivit-
tatus to be the young of D. radiale. It was not until Long-
ley and Hildebrand (1941:104-105) that D. bivittatum
was restored to specific status.

Ginsburg (1948:266-270) considered the Gulf of Mexico
population to be a distinct subspecies. However, desig-
nation by Ginsburg (1948:268) as D. arcuarium and not
as D. b. arcuarium led later authors incorrectly to ac-
cord specific status to Ginsburg's subspecies.

Figures by Meek and Hildebrand (1925:pl. 46, fig. 1)
and Randall (1968:fig. 86) are labeled as D. radiale but
actually depict D. bivittatum.

Diagnosis. — Diplectrum bivittatum in the Atlantic can
be confused only with D. radiale. The problem becomes
especially important in the area of sympatry along the
northern and northeastern coasts of South America.
Diplectrum bivittatum has a larger eye, lower number of
pectoral rays, a longer second and shorter third dorsal
spine (1.2-1.6 vs. 1.7-2.1, the second into the third) than
D. radiale. Also, the preopercular structure is more
pointed in D. bivittatum but definitely squared in D.
radiale. The upper caudal lobe is often extended as a fila-
ment in D. bivittatum but never in D. radiale. Coloration
and cheek scale rows A (mean 8.06 for D. bivittatum vs.
9.95) also separate the species.

Description. — Because of the systematic problems pre-
sent in D. bivittatum, the two geographic forms from the
Gulf of Mexico and Caribbean and an intermediate
group from south Florida will each be described separate-
ly for morphometric characters. Meristic characters ex-
cept numbers of pectoral rays, predorsal scale rows, and
gill rakers are virtually identical for all three groups and
are presented under the description for the Gulf of Mex-
ico specimens.

22

Gulf of Mexico.— D, X, 12; A, III, 7 (6-8); P,, 15.16(14-
16); pored lateral-line scales 48.54 (44-51); scale rows in
lateral line, 64.69 (54-75); scales above lateral line, 7.28
(6-9); scales below lateral line, 18.69 (16-22); predorsal
scale rows, 12.76 (10-15); cheek scale rows A, 8.06 (7-9);
cheek scale rows B, 11.86 (9-14); gill rakers, 22.03 (20-25).

Standard length, 91.38 (11.28, 75-125, 53). Total
length, 1,304 (47.77, 1,231-1,440, 45); fork length, 1,169
(13.07, 1,143-1,203, 51); body depth, 267 (12.34, 237-296,
53); head length, 347 (8.13, 324-369, 53); postorbital
length, 179 (5.76, 169-191, 53); snout length, 89 (6.01, 76-
106, 53); lacrimal width, 40 (4.53, 29-51, 53); maxillary
width, 28 (1.82, 25-34, 53); orbit length, 92 (4.74, 83-103,
53); orbit width, 72 (4.98, 60-83, 53); interorbital width,
71 (3.50, 64-80, 53); upper jaw length, 156 (3.80, 148-165,
53); lower jaw length, 176 (5.78, 158-188, 52); cheek
height, 110 (5.70, 96-123, 53); cheek length, 117 (4.35,
109-128, 53); caudal peduncle depth, 123 (4.32, 111-131,
52); pectoral fin length, 267 (10.38, 250-283, 51); pelvic
fin length, 222 (10.29, 203-249, 53); predorsal length, 344
(11.62, 316-372, 53); pelvic origin to lower jaw, 339 (17.59,
313-398, 52); pelvic origin to anus, 311 (11.93, 259-349,
53); postanal length, 398 (12.84, 364-425, 52). Dorsal
spine height: first, 52 (6.21, 38-64, 52); second, 78 (8.27,
59-95, 50); third, 105 (11.60, 79-134, 50); fourth, 124
(10.46, 101-147, 49); fifth, 132 (8.91, 103-152, 51). Anal
spine height: first, 36 (5.32, 20-49, 53); second, 63 (6.93,
33-73, 52); third, 86 (7.90, 69-105, 52).

Caribbean and South America. — Px , 15.45 (14-16);
predorsal scale rows, 14.18 (12-17); gill rakers, 19.67 (16-
23). Standard length, 94.00 (11.43, 76-122, 51). Total
length, 1,309 (60.74, 1,226-1,477, 48); fork length, 1.145
(24.56, 1,040-1,251, 51); body depth, 265 (14.63, 234-296,
51); head length, 353 (10.75, 328-376, 51); postorbital
length, 173 (5.05, 164-185, 51); snout length, 96 (7.15, 82-
110, 51); lacrimal width, 39 (3.99, 30-47, 39); maxillary
width, 28 (2.07, 24-33, 39); orbit length, 98 (5.44, 87-114,
51); orbit width, 78 (5.29, 69-90, 51); interorbital width,
74 (3.93, 63-82, 38); upper jaw length, 167 (5.48, 158-177,
51); lower jaw length, 187 (5.10, 175-197, 51); cheek
height, 108 (5.45, 100-119, 51); cheek length, 119 (5.73,
108-135, 51); caudal peduncle depth, 115 (4.29, 105-122,
51); pectoral fin length, 240 (11.25, 219-273, 48); pelvic
fin length, 204 (9.94, 186-226, 49); predorsal length, 352
(10.74, 329-377, 33); pelvic origin to lower jaw, 351 (24.85,
322-402, 46); pelvic origin to anus, 312 (18.27, 273-346,
51); postanal length, 389 (13.81, 360-416, 51). Dorsal
spine height: first, 49 (4.99, 39-62, 48); second, 73 (6.92,
58-87, 48); third, 99 (9.59, 84-120, 47); fourth, 116 (8.22,
102-131, 48); fifth, 124 (7.20, 113-137, 45). Anal spine
height: first, 32 (3.97, 24-41, 50); second, 61 (5.85, 47-76,
50); third, 81, (5.26, 69-93, 51).

Intermediate population from South Florida, Florida
Keys, Tortugas, and Cuba. — P[ , 15.39 (14-16); predor-
sal scale rows, 12.57 (10-15); gill rakers, 20.68 (18-23).
Standard length, 97.00 (13.54, 77-118, 36). Body depth,
257 (8.99, 242-269, 13); head length, 342 (9.84, 324-355,
18); postorbital length, 174 (7.48, 158-184, 17); snout

length, 90 (7.02, 74-108, 33); lacrimal width, 39 (4.15, 30-
50, 32); maxillary width, 27 (1.89, 24-30, 13); orbit
length, 91 (3.94, 84-102, 34); orbit width, 71 (3.60, 65-82,
34); upper jaw length, 161 (4.80, 147-171, 22); lower jaw
length, 179 (4.27, 173-185, 22); caudal peduncle depth,
120 (4.04, 113-130, 22); pectoral fin length, 258 (13.89,
229-294, 22); pelvic fin length, 222 (10.85, 198-244, 22).
Dorsal spine height: first, 51 (4.11, 45-61, 18); second, 74
(6.55, 64-86, 18); third, 100 (9.52, 85-119, 18).

Diplectrum bivittatum with narrow preopercular spur
slightly pointed at dorsal angle, rounded at ventral
angle, four to nine spines. Vertical and horizontal
preopercular limbs serrate. Anterior nostril with short
flap. Chest scales slightly smaller than lateral body
scales, not embedded. Gill rakers moderate in length; for
Gulf of Mexico and south Florida specimens first gill
raker at angle reaching to base of the fourth below; for
Caribbean and South American specimens first gill raker
only reaching to base of the third. Upper caudal lobe
on most specimens with elongate filament. Row of
small scales occasionally along base portion of third anal
spine.

Color. — The following color description is based on five
live adult D. bivittatum collected at Pensacola, Fla. in
the northern Gulf of Mexico in May 1973. Dorsum brown
with copper tinge. Sides tan with evidence of two broad
brown lateral stripes, the upper appearing to pass
through eye and continuing as a brown bar on snout. A
tendency for five or six blotches of more intensified pig-
ment along lateral stripes. Lateral surface tan, ventral
and lower lateral surface white, but each scale bearing a
small yellow dot at base. Chest and throat white. A dis-
tinct caudal spot (equal to eye in size) appearing as an
extension of midlateral stripe but separated from it. Pec-
toral fin pale orange, tips black. Pelvic fins pale yellow-
orange with white anterior and posterior edges. Anal fin
white at base and distal edge, pale yellow through medial
fin portion. Snout with narrow brown bar, and a narrow
blue one beneath, below this a blue cheek stripe running
upward from middle anterior lacrimal edge posteriorly to
eye, passing under the orbit and across upper cheek.
Another blue stripe below this on central cheek area.
Opercle with pale greenish-yellow blotch. A thin blue
line present on lower jaw and along upper maxillary. Pale
blue spots on head with two pale blue chevrons above.
Spinous dorsal fin with wavy blue bar at base, above this
an orange bar, above this a blue bar, and distally an
orange bar. Soft dorsal fin with irregular yellow spots
along base. Above are two parallel longitudinal rows of
pale blue spots on an orange field. Distal portion of fin
with orange bar. Caudal fin with five or six orange spots
in vertical rows on a pale blue field, upper lobe and edge
orange, lower caudal blue. Upper pharyngeal region
bright yellow. Posterior branchial surface with yellow
and black blotches, medial opercular surface black.

An 80-mm-SL specimen from Galeta Point, Panama,
agreed with the above color description with the fol-
lowing emendations: yellow spots and blotches on dorsal
fin paler and the distal area of dorsal membranes with

23

wedge-shaped white spots behind anterior spines.
Anterior dorsal spine orange. Dorsal caudal peduncle
with two dark brown spots bordered by white spots for-
ming a longitudinal line. Dorsum brown-copper with a
trace of olive-green pigment.

Orange and yellow pigments are pale to clear in
preserved specimens, blue appears dusky. Upper medial
opercular surface dusky, paler below; posterior branchial
surface dusky.

Size. — Diplectrum biuittatum is a small species, the
largest specimen recorded in the present study is 126 mm

SL.

Habits. — Diplectrum biuittatum has been captured at a
wide range of depths from 2 to 110 m but most frequently
at 15-80 m. In January-February 1973 I observed the
species off Galeta Point, Panama, in 2-3 m of water. It
appeared to be solitary and occurred over a clean coral-
shell bottom at the base of reefs. This species is the most
insular of all Diplectrum, frequently taken in the Antilles.
Starck (1968:19) listed D. biuittatum as being common
at Alligator Reef, Fla. Gulf of Mexico specimens are
found more often over a more mud-silt bottom than
Caribbean and intermediate populations.

Distribution. — Diplectrum biuittatum is found in the
western Gulf of Mexico but terminates its northern Gulf
of Mexico distribution near Pensacola, Fla. (Fig. 11). It
occurs in the area of the Dry Tortugas, northern Cuba,
southern Florida, and northward along the Florida east
coast to lat. 29°43'N, long. 80°49'W near St. Augustine,

Fla. Beebe and Tee-Van (1933:135-136) reported D.
biuittatum (as D. radiale) from Bermuda. This record is
not plotted as it could not be verified (W. Smith-Vaniz,
Acad. Nat. Sci. Philadelphia, pers. commun.). The
species is found at Jamaica, Puerto Rico, and throughout
the Lesser Antilles. Diplectrum biuittatum is also found
off Central America, along the northern and southern
coasts of South America, and terminates its southern
distribution off Brazil at lat. 2°23'S, long. 40°31'W.
The type-specimens are from Martinique and Havana,
Cuba.

Variation. — Meristic and morphometric data confirm
the existence of two reasonably distinct geographical
subspecies. One subspecies is in the Gulf of Mexico and
the other is found in the Caribbean and along the
northern South American coast; an intermediate popula-
tion occurs in the vicinity of Florida Keys, southern
Florida, Tortugas, and northern Cuba (this general area
referred to hereafter as S. Florida).

The Gulf of Mexico subspecies has a lower mean num-
ber of pectoral rays and predorsal scale rows, but a much
higher mean gill raker count than the Caribbean and
South American subspecies. The S. Florida population is
intermediate for mean number of pectoral rays and gill
raker count. The Gulf of Mexico subspecies has several
morphometric characters differing from the Caribbean
and South American subspecies (Table 9). The S.
Florida population is intermediate in several characters,
notably snout length, postorbital length, upper- and
lower-jaw lengths, caudal peduncle depth, pectoral fin
length, and dorsal spine height.

30°-

o°-

30"-

Figure 11.— Distribution map for Diplectrum bivit-
tatum (circles); D. maximum (triangles); andD. eury-
plectrum (squares). [Localities are based on the
materials examined in this study.]

24

VWOdOJJVVJ Q

wnjawnd q

Uituvqv] q

wruiodjd.und q

srumos q

luniuixuw (j
wnwixDiu Q

dUOljddDUOJ Q

wrujsoj (]

wnoiJiJDd q

djDlpDJ Q

umpmxmq q

Bpuoy "g
wniDHiajq q

JInO
lunjvwiaiq q

uj^qjnos
tunsoiujo) q

uinsouiuo) q

CO t> CM tO i— I iC t>

O (DO 05 t CO QO
CO CO w

C^COCMC^C)COO>iOtDCNaQtDr- I
COCOCCHHCOCCOlCO-tQH

HHHHHCNHCOCOCJtt

CO [•■ O CN Tf

Tf CO CO Tf Tf

CNO)t<(Ot>HQ}COCO(Q(NOH^O}HCN^HO>0)^

ior-ooaooocNcxi'tcooot~-r-r-aocNC)0)coo(X)co^

t-h CN i-h CN CO CM t-h.— ir-tr-tr-HCNCNCOCOCO

t^r-ia»HHTtO)WCOlOCC^CC'tCOHhOWC005CDCC
h W H CN CO (N

r-Hi— (.-Hi— <CNCNCOCOCOCO

NC3«HeOHP3tOCON^003JCOHOtOtOtNU5Hfll^
HtOOi01CNTtO'TWOOOCOC0011'COO»0 i— < c^ to o c-
rH IN -H CN ^t CN -H r-< HCNHHHCNc^iconnn

lOeOCNOiCDCOOlOCOCNCIOffiCN

aico^toiOGCoocNcoair-^Tf

CN »— i CM CO •— ■ i—<

IN (M CO CO CO CO

iniocoiocoHNTj'O^ONaicoor-'t'tNiN

Tf'tt^COCOHOiOCOCfliOt-iOhCOlXNmO"
N IN H IN CO (N h ,_< ,_| -

h CM Ol CO CO CO Tf

io,*oi(--'-iair-icoioaiixicocoHCCHCcococor-ccco

CDiMXiOCOCOaiCOCOaiht-^CCOHHCOfllCOCOQOO
CN i— i CM CO t— I r-l .— . HHiN--iCOCON-<f

mhhOico'tr'toinaiaiMHOccocDiflinccr-H

hQO^t-HffiCOCOOlhCCiOQOHC'-iiOHCOCOHO
N H N CO N Hr-iHr-irtNNCOCOCO'*

OOiCCOO^COCOi-iI>0

NNCOCCCOOlOlCCNai

rH CO M N CO r- rH

ON "«t **
t- t— CO C—

00301-— i t- -T O CD ^ C-

HNOliOHCCcONOi

rHrHHNNCO^COCO

ooixcoairH^Q^aicoocotoooccaj^ioaiTrcoci)

iOCOt>t-CO^O)TtCN[--CCr-CCr-CNrH(N'taiuOCOr-iO

H (N H N CO N' HHHHHtNHCOCOCO't

t- cx> ai ■<* c— -— i co » ^ r- <LCO^roo^coiccor-c~-cM^tcT3

C0)^l>CXiCCrHa)TjCNCX)CI)r-'XiC0iN^CNC0C0C0'trH01

H [N H CN CO CS HHHHHNHCOCOCOCO

CO O O

tT CO oo to

o ao ex.

CO rH CN CN

co m o

aiioiococotocnooaoco

O^tOUDOCJlCOCNCXlO
CO ^ CM CO ^

U0> Tj» o

(N CO H

■* Oi t> t> OS en

o to to -v t- CO

CO h O) CO h

Ot-OOCXiuOOCOCN^HCNaJ
tOCOOHHTj<plOiOHCO

^H^h,_i^-.^hCNCNCOCOCOCO

O X CN
CM UO CN
t— i CN CN

COCOO^COt^N^OlHCO
HHHHHiMiMCOCOCOCrj

0)01C0O(MClH(NhC0C0C0OHai(IiiO00OON'tH
rHCCQOhCD^OiONOOCDr-CDQOrHCNNCOHiO'l'rHO

-— ■ CN >— I CN CO H H HHHHHNNCOCOCO't

O O •— i iO Ol h CO O) CD
^■"tOlhiOCjlHiniN

H CO H N CO r-l rH

CTiTj-OOiiOCO^LOtO'Ti'— itOCM

m r- tor-CMCNcocorHTj'Troo

HHHHHNNCOCOCO'I'

CN CO UO Tf CM

m r— o cm co

CO O) O O CO

i— ( ao m io o
lo r- CD o •— i

CN Tf t>

r

J3

tu

H.

C

TJ

-

"a

-C

tit

C

TJ

TJ

>

X
be
C

3

TJ

■s-

"ca

c

"5

c

a*

a*
TJ

z

3

E

2

-I

■

"

C

—

c

X

S-

U.

X

x;

a,

f.

—

0

a

mm y

c 0 « _n S

Cu CD r-; "7"! 3

T "T M OC TJ

* i* '53 a Si

■" a o -c j= «

— » s —

~ ~ CO J

■o oo«

DJOOOO.DhO.S.0,

S c

<D

— Q.

c

CO M

a.

ca ro

CO fe ,— . CM

o c - J

CO

Tf

ir:

g-«N

CU C

<

25

Systematic comments. — Ginsburg (1948:266-270)
described the Gulf of Mexico form as the subspecies D. b.
arcuarium. Ginsburg (1948:268) and later Hildebrand
(1955:208) suggested that further studies should be con-
ducted on the correct status of D. b. arcuarium es-
pecially with regard to proper identification of D. bivit-
tatum from South America (for comparative purposes).
Confusion of D. bivittatum with D. radiale had occurred
for many years. In the present study only "good" D.
bivittatum from the Caribbean and South America were
compared with D. b. arcuarium. Although the question
of the "true" status of D. b. arcuarium may never be
completely answered, on the basis of meristic counts,
morphological data, and geographical distribution, it is
suggested that D. b. arcuarium Ginsburg 1948 be recog-
nized as a subspecies. The Caribbean and South
American form of D. bivittatum should hereafter be con-
sidered subspecifically as D. b. bivittatum (Valen-
ciennes 1828). The S. Florida population of D. bivit-
tatum represents an intermediate of the two subspecies
both morphologically and geographically. Future studies
should be conducted to examine further the nature of the
S. Florida population.

Specimens examined. — A total of 568 specimens from 22
to 126 mm SL. S. Florida and Keys: UMML 3804. l (63);

UMML 10170, 1 (68); UMML 2959, 1 (98); UMML 3957, 6 (86-109);
UMML 2081. 1 (109); UMML 10150, 5 (22-84); USNM 117201, 20 (73-
119); TABL [Siluer Bay Stn. 4329), 2 (33-53); TABL (Silver Bay Stn.
2354). 1 (47); UMML 12324, 2 (90-104); FMNH 61315, 5 (105-118);
USNM 4869, 3 (90-110); FSBC 799, 1 (103); FSBC 1583, 3 (57-90); FSBC
336, 11 (55-90); MNHN 5889, paralectotype, 1 (80); MCZ 10242, 1; UPR
2086, 1 (96). NORTHERN GULF OF MEXICO: FMNH 50726, 6 (62-90);
GCRL 4208, 1 (81); GCRL 533, 2 (98-115); GCRL 35, 2 (81-99); GCRL
671. 1 (33); GCRL 2318, 1 (91, deformed snout); UW 16899, 3 (62-83); UW
16569, 3 (95-110); USNM 155177. 2 (81-100); USNM 129817, 1 (74);
TABL 101472. 4 (69-97); TABL (Silver Bay Stn. .5004), 1 (105); FSM
5207, 7 (73-1021; FMNH 59807, 5 (66-112); FMNH 16187, 1 (78); FMNH
16188, 1 (78); GCRL 276, 2 (87-91); USNM 144214, 1 (83); USNM 144213,
1 (117); USNM 155175, 2 (72-107). TEXAS: FSM 5177, 4 (96-103);
FMNH 40263, 3 (76-105); FMNH 45144, 2 (90-91); INP LB3241, 1 (113);
UMML 28700, 3 (93-108); UTIMS 926, 2 (30-37); UTIMS 1629, 2 (88-
100); UTIMS 920. 1 (97); UTIMS 918, 1 (111); UTIMS 925. 4 (63-76);
UTIMS 1484. 3 (30-81); UTIMS 1627, 2 (88-88); UTIMS 922, 37 (56-112);
UTIMS 1024, 2 (101-105); CAS SU 50185, 1 (93); USNM 155176, 2 80-
85); USNM 144215, 9 (91-112); USNM 144216, 1 (80); USNM 155173, 1
(52); USNM 155174, 1 (96); USNM 118536, 1 (88); USNM 118535, 11 (90-
97); UMML 15398, 2 (82-90); UMML 7749, 4 (33-51). YUCATAN: FSM
5209. 1 (97); UTIMS 917, 2 (90-96); UTIMS 921, 6 (54-125); USNM
188298, 30 (88-107). JAMAICA: LACM 5433, 1 (70); LACM 5434, 3 (99-
108); LACM 6203, 8 (87-110); LACM 6204, 1 (72); LACM 6201, 16 (62-
100); LACM 6200, 10 (62-113); LACM 6199, 23 (62-113); UMML 28699, 1
(90) PUERTO RICO AND ANTILLES: UMML 7324. 1 (49); USNM
128262, 1 (109); UMML 7110. 1 (88); FMNH 64989, 2 (86-97); FMNH
64982. 4 (110-114); MNHN 6988, lectotype, 1 (108); BMNH 1904-7.26:67,

1 (97); LACM 6741-14. 7 (46-88); LACM 6768-1, 22 (37-98); TABL
(Oregon Stn. 2631). 1 (105); UPR 2713, 5 (67-81); UPR 2158, 1 (93); UPR
239. 1 (94l; UPR 1329, 2 (76-84); UPR 1328, 1 (42); UPR 2692, 1 (92); UPR
2055, 1 (65); UPR 238, 1 (90); UPR 1330, 1 (44). BELIZE: TABL 105165,

2 (94-99); TABL 105164, 10 (86-112). HONDURAS: FMNH 64956, 4 (95-
106); USNM 185143, 2 (96-99); TABL 105155, 11 (94-107); TABL 105156,
- (86-103); TABL 105157, 7 (82-102); TABL 105158, 1 (96); TABL 105160,
30 (74-lD'), COSTA RICA: UCR 253-31, 2 (88-92); LACM 30729-5, 1
(103). PANAMA: UCR 226-4, 3 (103-114); FMNH 8392, 9 (44-62);
FMNH 8393. 3 (31-40); MCZ 45530, 2 (31-64); UMML 23897, 5 (69-90);
I IMML 22007, 6 (90-103). VENEZUELA: RMNH 8448, 1; RMNH 8422,
1 (120); RMNH 9377. 1 (109); USNM (Oregon Stn. 4865), 6 (37-117);

TABL (Geronimo Stn. 7), 1 (122); UMML 28736, 1 (119); UMML 28759,
1 (110); UMML 28769, 4 (93-103); UMML 28767, 6 (94-107); UMML
28694, 7 (92-112); UMML 27242, 7 (31-61); UMML 27220, 12 (76-99);
UMML 26613, 3 (49-54); UMML 17071, 1 (102); UMML 16302, 1 (117);
UMML 16283, 1 (85); UMML 15397, 10 (64-78); UMML 15390, 2 (98-
104); UMML 15368, 1 (108); UMML 15301, 5 (108-115); UMML 15081, 1
(96). TRINIDAD TOBAGO: BMNH 1920-12.22:38, 1 (110); UMML
28757, 1 (116); UMML 28670, 1 (100); UMML 16802, 2 (112-
122). GUYANA: UMML 5375, 1 (115); FMNH 67904, 5 (90-118); USNM
156792, 2 (95-99); UMML 12247. 2 (117-126). FRENCH GUIANA:
UMML 6706, 3 (105-107). BRAZIL: USNM (Ace. No. 247715), 1 (103).

Diplectrum radiale (Quoy and Gaimard 1824)

Figures 6, 8, 9, 12

Serranus radialis Quoy and Gaimard 1824:316-317 (origi-
nal description, Rio de Janeiro); Cuvier and Valen-
ciennes 1828:243-244 (description, Brazil); Cuvier
1829:139 (generic position); Boulenger 1895:297-298 (in
part, catalog).

Serranus irradians Valenciennes in Cuvier and Valen-
ciennes 1828:244-245 (in part, original description,
Brazil); Canestrini 1860:306 (systematics).

Centropristis radialis Giinther 1859:83 (catalog); Stein-
dachner 1876:6 (systematics).

Centropristis ayresi Steindachner 1868:1-3, pi. 1, fig. 1
(original description, Brazil).

Diplectrum radiale Jordan 1895:451 (Mexico); Jordan
and Eigenmann 1890:397-398 (in part, systematics);
Jordan and Evermann 1896:1203-1205 (systematics);
Fowler 1907:263 (in part, distribution); Fowler
1916a:534 (West Indies); Jordan et al. 1930:318 (in
part, check list); Longley and Hildebrand 1941:104
(systematics); Fowler 1942:158 (Brazil); Ginsburg
1948:268-270 (systematics); Schultz 1949:119-120
(Venezuela); Briggs 1958:272 (in part, distribution);
Durand 1961:46, 50 (field data, Guyana); Brandao
1964:19 (Brazil); Bullis and Thompson 1965:44
(southeast coast of United States); Cervigon 1966:323-
324 (systematics, distribution, and biology,
Venezuela); Mago Leccia 1970:94 (Venezuela); Ewald
et al. 1971:54 (Venezuela); Miller and Jorgenson
1973:310 (meristics).

Haliperca radialis Miranda-Ribeiro 1915:29-30 (Brazil);
Miranda-Ribeiro 1918:93 (synonomy, Brazil). Diplec-
trum spp. Lowe 1962:695 (in part, Guyana).

Valenciennes in Cuvier and Valenciennes (1828:244-
245) described Serranus irradians and that description is
clearly of D. formosum. However, examination of the
four syntypes for S. irradians shows that MNHN A461,
A464, and 7018 are D. formosum but the other syntype,
MNHN A462, is actually D. radiale.

Many references in the synonomy are listed "in part"
owing to confusion when Gulf of Mexico Diplectrum,
other than D. formosum, were called D. radiale while in
the Pacific Ocean D. pacificum was referred to as D.
radiale until the description of D. pacificum by Meek
and Hildebrand (1925).

Centropristis ayresi was described by Steindachner

26

(1868:1-3) but later it was placed in the synonymy of D.
radiale [Centropristis (- Diplectrum) radiale, Stein-
dachner 1876:6].

The species was included under the genus Haliperca
by Miranda-Ribeiro (1915:29-30) at the inference of
Gill's (1863) generic interpretation.

Diagnosis. — Diplectrum radiale is a species with a nar-
row preopercular spur and in the Atlantic can only be
confused with D. bivittatum. Separation of these species
is possible when one considers the relative height of the
dorsal spines, especially the number of times the second
spine fits into the third (1.7-2.1 for D. radiale, 1.2-1.6 for
D. bivittatum). The eye is smaller for D. radiale. Also the
mean number of pectoral rays and of cheek scale rows A
is higher; the preopercle generally is more squared off in
D. radiale and definitely more pointed in D. bivittatum
at its upper angle.

Diplectrum radiale in the Atlantic is a "geminate
species" with D. pacificum in the Pacific as defined by
Jordan (1908). The species can be discriminated by
several characters: notably the gill raker length, color
pattern, and other characters considered under the diag-
nosis for D. pacificum.

Description.— D, X, 12 (13); A, III, 7; Plf 16.79 (16-18);
pored lateral-line scales, 48.82 (47-52); scale rows in
lateral line, 63.36 (54-71); scales above lateral line, 7.28
(6-9); scales below lateral line, 18.57 (15-21); predorsal
scale rows, 15.25 (12-19); cheek scale rows A, 9.95 (9-12);
cheek scale rows B, 11.73 (10-14); gill rakers, 18.25 (15-
21).

Standard length, 136.97 (23.12, 84.4-168, 42). Total
length, 1,248 (10.70, 1,222-1,276, 42); fork length, 1,179
(9.75, 1,160-1,207, 42); body depth, 264 (20.10, 218-302,
40); head length, 367 (19.54, 335-406, 42); postorbital
length, 211 (16.61, 188-254, 42); snout length, 93 (8.33,
74-114, 42); lacrimal width, 48 (5.15, 40-57, 32); maxil-
lary width, 27 (2.72, 22-30, 32); orbit length, 81 (5.55, 71-
95, 40); orbit width, 65 (5.95, 54-79, 42); interorbital
width, 70 (8.56, 54-83, 32); upper jaw length, 164 (12.41,
144-193, 42); lower jaw length, 180 (9.44, 160-208, 42);
cheek height, 129 (11.98, 105-149, 42); cheek length, 118
(6.50, 107-132, 42); caudal peduncle depth, 125 (6.29,
110-137, 39); pectoral fin length, 236 (6.64, 213-250, 42);
pelvic fin length, 187 (10.63, 152-204, 42); predorsal
length, 367 (11.98, 348-397, 32); pelvic origin to lower
jaw, 342 (12.82, 318-372, 39); pelvic origin to anus, 314
(19.29, 264-355, 41); postanal length, 399 (11.45, 377-420,
42). Dorsal spine height: first, 43 (5.14, 33-54, 42); sec-
ond, 64 (5.12, 55-72, 40); third, 118 (8.25, 104-136, 41);
fourth, 128 (7.32, 114-144, 40); fifth, 126 (7.03, 110-140,
38). Anal spine height: first, 30 (4.06, 21-38, 42); second,
58 (5.87, 47-76, 42); third, 79 (7.35, 64-97, 42).

Preopercular spur square but narrow, bearing 5-10
elongate spines. Vertical and horizontal preopercular
arms serrate. Anterior nostril with elongate nostril flap.
Gill rakers short, the first at angle not reaching to base of
third below. Scales present in a row along second anal
spine. Chest scales not noticeably smaller and not em-

bedded. Caudal lobes nearly equal in length, upper only
slightly longer and never extended as a filament. Caudal
fin margin nearly straight.

Color. — Color notes are from preserved specimens.
Dorsum and lateral body surface light to medium brown,
five to seven irregular dark vertical bars on lateral sur-
face terminating before ventral region. Caudal spot dis-
tinct, equal to eye size. Ventral area white. Snout, head,
and cheek gray-brown. Lower preopercle pale below.
Maxillary gray-brown and paler distally. Lower jaw
dusky at midline. Pectoral, anal, and pelvic fins pale.
Caudal fin with three to four pale broad vertical bars bor-
dered by darker bars of equal breadth. Two indistinct
dark spots on dorsum of caudal peduncle, separated by a
smaller pale spot. First and second dorsal spines dark.
Spinous dorsal fin dusky, black at spine tips. Soft dorsal
with two regular rows of pale spots on a dusky field. Soft
dorsal ray tips and caudal ray tips black. Medial oper-
cular surface black or dusky above, white below.
Posterior branchial surface dusky.

Size. — The species attains a reasonably large size; 197
mm SL is the maximum size examined in the present
study.

Habits. — Depth data indicate that D. radiale occurs in
depths ranging from 20 to 80 m. Durand (1961) con-
firmed this depth distribution off the coast of Guyana.
He took the species from sand-mud, and shell and sand-
bottom types. Cervigon (1966) stated the species prin-
cipally consumes crustaceans and small fish.

Distribution. — Diplectrum radiale is found in coastal
areas of eastern and northeastern coasts of South
America (Fig. 12). Its westernmost distribution along the
Venezuelian coast is the Gulf of Venezuela. The species
extends southward to Sao Paulo, Brazil. Literature
records indicate the species occurring southward to
Uruguay but this has not been confirmed. One specimen
(BMNH 1904-3.15:24-5) was accompanied by a label
which stated "St. Thomas." It was presumed that this
meant St. Thomas Island in the Virgin Islands. Other
than this record the species has not been taken from the
Caribbean Antilles. It is possible that the locality is in
error or that "St. Thomas" refers to a city on continen-
tal South America. The type-locality is Rio de Janeiro.

Variation. — No discernible geographic variation in
meristic characters was observed for this species. There
appears to be a slight (but statistically untested) trend of
morphological variation in D. radiale. Northern
specimens appear to have slightly larger heads and jaws
than specimens from the southern parts of South
America.

Specimens examined. — A total of 103 specimens from 64

to 197 mm SL. ST. THOMAS: BMNH 1904-3.15:24-5. 1
(174). VENEZUELA - TRINIDAD: UPR 1830. 1 (89); BMNH 1922-2.3:29,
1 (128); UMML 28757, 1 (166-166); UMML 28749, 2 (195-197); UMML

27

60°

Figure 12.— Distribution map for Diplectrum radiale
(circles); and D. pacificum (squares). [Localities are
based on the materials examined in this study.]

21851, 3 (170-173); UMML 16302, 1 (107); USNM 123132, 1 (100); TABL
(Gulf of Cariaco, 14 Apr. 1966). GUYANA - NE BRAZIL: FMNH 64967,
2 (166-167); FMNH 64980, 4 (151-158); UMML 12237, 1 (136); USNM
185497, 4 (151-164); USNM 156806, 1 (184); USNM 185484, 5 (155-168);
UMML 11536, 1 (159); UMML 11581, 1 (186); UMML 11588, 1 (178);
UMML 11553, 1 (153). SE BRAZIL: BMNH (Bahia), 1 (142); MZUSP
9007, 1 (102); MZUSP 9008, 1 (96); MZUSP 9009, 1 (90); MZUSP 9010, 1
(84); MZUSP 9011, 1 (86); MZUSP 9002, 1 (141); MZUSP 9003, 1 (125)
MZUSP 9004, 1 (113); MZUSP 9005, 1 (119); MZUSP 9006, 1 (119)
MZUSP 8997, 1 (136); MZUSP 8998, 1 (131); MZUSP 8999, 1 (131)
MZUSP 9000, 1 (107); MZUSP 9001, 1 (151); USNM 100858, 5 (119-127)
MNHN 7015, holotype, 1 (125); MNHN 7015, paratype, 1 (140); MCZ
44166, 23 (90-170); MCZ 4504, 4 (153-160); BMNH 1906-6.9:14-15, 2 (174-
177); BMNH 1923-7.30:77-9, 1 (142); CAS SU 54082, 1 (64); MCZ 10245,
1 (124); BMNH 1892-6.29:4, 1 (127); RMNH 166, 1 (140); MZUSP 8990, 1
(113); MZUSP 8987. 1 (178); MZUSP 5222, 1 (165); MZUSP 8991, 1
(115); MZUSP 8988, 1 (124); USNM 100858, 1 (131); MZUSP 8995, 1
(135); MZUSP 8993, 1 (122); MZUSP 8994, 1 (131); MZUSP 8992, 1
(157); MZUSP 8996. 1 (112); MZUSP 10244, 2 (82-109); MNHN A462, 1
(166).

Diplectrum pacificum
Meek and Hildebrand 1925

Figures 6, 8, 9, 15

kins 1892:127-128, 142 (Gulf of California); Jordan
1895:451 (Sinaloa, Mexico); Jordan and Evermann
1896:1203-1205 (systematics); Gilbert and Starks
1904:97 (Panama); Fowler 1916b:409 (Panama); Os-
burn and Nichols 1916:163 (Baja California); Kumada
1937:39, pi. 6, fig. C (Mexico).

Diplectrum pacificum Meek and Hildebrand 1925:472,
477-478, pi. 46, fig. 2 (original description, Panama);
Jordan et al. 1930:318 (check list); Breder 1936:24
(Pacific coast); Walford 1937:123 (general account);
Seale 1940:20, 23 (Panama); Fowler 1944:262-263
(color notes); Hildebrand 1946:185 (key); Grey,
1947:140 (type catalog); Berdegue Anzar 1956:53, 264-
265, fig. 183 (key and fishery data, Mexico); Orces
1959:81 (systematics, Ecuador); Gosline 1966:91-112
(osteology); Rosenblatt and Johnson 1974:178-191
(key, systematics); Chirichigno F. 1974:294-298 (key,
Peru); Bortone in press a (gonad morphology); Bor-
tone in press b (life history, Panama).

Diplectrum radialis Streets 1877:52-53 (systematics, Ba-
ja California).

Centropristis radiale Steindachner 1876:6 (in part, Pan-
ama).

Serranus radialis Jordan and Gilbert 1881:274 (Baja
California); Jordan 1886:376 (Panama and Mazal-
tan); Jordan 1887:82 (in part, catalog); Boulenger
1895:297-298 (in part, catalog).

Diplectrum radiale Jordan and Bollman 1890:157-158,
181 (Panama); Jordan and Eigenmann 1890:398-399
(in part, systematics account); Evermann and Jen-

Earlier authors considered D. pacificum as a Pacific
Ocean representative of D. radiale. Meek and
Hildebrand (1925) described the Pacific form as a new
species and considered it distinct from D. radiale as they
knew it. However, the specimen of D. radiale figured by
them (Meek and Hildebrand 1925:pl. 46, fig. 2) is ac-
tually D. bivittatum. Nevertheless, D. pacificum and D.
radiale are distinct species, qualifying as "geminate
species" as defined by Jordan (1908).

28

Diagnosis. — Diplectrum pacificum is distinguished from
other Pacific Diplectrum by a combination of characters
which include: less than 30 total gill rakers; 7 anal rays;
lack of extended tips on second, third, or fourth spines;
lack of an enlarged second anal spine; a relatively high
number of cheek scale rows A (8-12) and B (10-14); and a
narrow preopercular spur.

Diplectrum pacificum could readily be confused with
D. rostrum, another Pacific species. It can be differen-
tiated from D: rostrum by the following characters: snout
spot present in D. pacificum but not a distinct bar as in
D. rostrum; the soft-dorsal-fin color patterns, total gill
raker counts; orbit length and width, preopercle shape
and other characters listed under diagnosis for D. ros-
trum.

It might readily be confused with its geminate Atlan-
tic form, D. radiale. Separation of these species ad-
mittedly is difficult but several discernible differences
are evident. Meristically the number of gill rakers is
higher for D. pacificum (20.25 vs 18.25), but other
characters show a high degree of overlap and do not allow
discrimination. Morphometrically there are no charac-
ter differences reliably separating these species but in
general the eye is smaller for D. pacificum, the caudal
peduncle deeper, the upper caudal lobe more elongate,
and fins and spines tend to be longer in D. pacificum.
Gill rakers are moderate in length in D. pacificum (the
first at the angle reaching to the origin of the fourth
below) and are short in D. radiale (the first not reaching
to the base of the third, usually reaching only to the base
of the second below).

In preservative there are color differences between the
species which surely are more distinctive in fresh
specimens but no satisfactory color data for living D.
radiale exist. Caudal fin of D. pacificum with five or six
thin, light caudal bands bordered by wider dusky bands;
D. radiale bears four light bands equal in width to the
dusky bands which border them. The soft dorsal fin bears
several irregular rows of spots in D. pacificum but D.
radiale has one or two rows of regularly spaced, larger
spots.

Description— D, X, 12 (13); A, III, 7 (8); P, , 17.02 (16-
18); pored lateral-line scales, 49.05 (47-51); scale rows in
lateral line, 65.48 (58-74); scales above lateral line, 7.64
(6-9); scales below lateral line, 18.88 (16-21); predorsal
scale rows, 14.85 (11-19); cheek scale rows A, 9.70 (8-12);
cheek scale rows B, 12.21 (10-14); gill rakers, 20.25 (17-
24).

Standard length, 150 (29.12, 101-211, 40). Total
length, 1,289 (24.57, 1,242-1,354, 40); fork length, 1,178
(17.23, 1,128-1,229, 40); body depth, 278 (16.66, 240-327,
37); head length, 369 (12.13, 339-392, 39); postorbital
length, 211 (9.54, 188-231, 40); snout length, 97 (6.82, 84-
108, 40); lacrimal width, 49 (3.98, 39-56, 39); maxillary
width, 27 (1.72, 24-31, 39); orbit length, 79 (6.66, 66-90,
40); orbit width, 63 (5.07, 53-71, 40); interorbital width,
70 (4.27, 63-77, 39); upper jaw length, 163 (6.15, 149-176,
38); lower jaw length, 176 (5.54, 163-189, 39); cheek
height, 128 (6.77, 114-146, 40); cheek length, 118 (5.90,

108-133, 40); caudal peduncle depth, 129 (6.18, 115-142,
40); pectoral fin length, 244 (10.04, 218-268, 40); pelvic
fin length, 195 (8.05, 176-212, 39); predorsal length, 359
(9.22, 341-378, 39); pelvic origin to lower jaw, 334 (10.93,
307-358, 28); pelvic origin to anus, 313 (20.23, 273-363,
40); postanal length, 406 (13.95, 364-428, 40). Dorsal
spine height: first, 49 (5.83, 36-60, 39); second, 70 (5.80,
60-84, 37); third, 122 (6.42, 106-134, 38); fourth, 135
(7.33, 120-150, 40); fifth, 136 (6.98, 122-152, 39). Anal
spine height: first, 33 (4.58, 21-44, 40); second, 60 (6.33,
47-72, 40); third, 80 5.62, 68-92, 39).

Preopercle with narrow spur bearing five to eight
elongate spines (an occasional spine missing). Vertical
and horizontal preopercular arms serrate. Anterior nos-
tril with short flap. Upper caudal lobe noticeably longer
than the lower. Caudal fin only slightly forked. Second
anal spine as stout or more stout than third. Gill rakers
moderate in length, first at angle reaching to base of
fourth below. Chest scales smaller than lateral scales and
slightly embedded.

Color. — Color notes were taken from live specimens
from the Bay of Panama in January-February 1973. Dor-
sum green-brown with a copper cast. Ventral surface
white. Sides of juveniles with two distinct lateral stripes,
the lower passing through eye into snout. In adults
lateral bars becoming disrupted by five or six vertical
brown bars. Caudal spots distinct and equal to eye size.
Lower lateral surface (area just above anus to midanal
fin) with pink-orange blotch, about one-fourth body
depth in height. Anal fin with white bars distally and
proximally, the central portion of the fin yellow. Anterior
and posterior edges of pelvic fin white, the central por-
tion yellow. Pectoral fin pink-orange, occasionally a blue
bar at its base. First and second dorsal spines dark blue-
black, remainder of spines with orange tips. Spinous dor-
sal fin membrane clear at base, dusky brown through its
central portion and pale blue above. Distal edge of soft
dorsal fin orange. Remainder of membrane a pale blue
field with irregular series of orange spots and blotches.
These spots variable in shape and pattern between in-
dividuals; usually the anterior spots more rounded,
posteriorly the spots elongate or fuse with other spots
presenting a wavy-bar appearance. Upper caudal lobe
bright orange, lower white. Two parallel orange stripes
just below upper caudal lobe. Distal caudal ray tips
black. Body of caudal fin with five to six pale orange bars
separated by equally wide pale blue bars. Caudal fin
base with small orange spots on pale blue background.
Snout region with irregular elongate pale orange-yellow
bar running from anterior snout to anterior orbit rim.
Two or three pale orange-yellow spots below this, a thin
similarly colored bar just below orbit rim, and a pale
orange-yellow wavy cheek bar. The first snout bar ir-
regular and not well delineated. Occasional spots of
orange about preopercle and on the opercular-interoper-
cular area bordering preopercular spur. Upper pharyn-
geal region bright yellow in adults. The medial oper-
cular surface black above and white below. Posterior
branchial region variable, black to clear. Dentary and

29

maxillary pale blue, small spots of orange present on
these bones.

Color is faded in preserved specimens but above pat-
tern may be discernible with orange appearing pale and
blue appearing dusky.

Size. — The largest specimen observed in the present
study is 224 mm SL, however, Rosenblatt and Johnson
(1974) recorded a specimen 226 mm SL.

Habits. — Diplectrum pacificum was collected from the
Bay of Panama at many different localities. Juveniles
(30-70 mm SL) were common in shallow water (2 m) over
mud bottom. Adults were collected in deeper water (20-
30 m) over mud-sand bottom. The species was abundant
at certain localities and was usually taken with D. mac-
ropoma (Bortone in press b). Field-collection data from
museum specimens indicate the species has been taken
from water 0.1-90 m deep but appearing more frequently
at 15-30 m. The shallowest depth record is about 0.1 m
(UA66-69) from a tide pool near Puerto Pehasco, Gulf of
California.

Distribution. — Diplectrum pacificum is found in the
coastal areas of the Gulf of California (Fig. 12), north-
ward along the outer Baja California coast of Bahia San
Juanico (lat. 26°15'N, long. 112°28'W), along the Cen-
tral American coast, and southward to Ecuador-Peru
(lat. 3°15'S, long. 80°25'W). The type-locality is Naos Is-
land in the Bay of Panama.

Variation. — There is noticeable variation in the posterior
branchial surface pigmentation. Southern populations
tend to be very heavily pigmented in this body region.
Northward along the extensive Mexican coast the degree
of pigmentation decreases. Finally, populations of D.
pacificum in the Gulf of California and outer Baja
California are generally devoid of pigment. Geographical
variation occurs with the number of scales in lateral line
(Table 2); a trend from higher numbers to lower num-
bers occurs from north to south. It would appear that
more concentrated studies on variation in D. pacificum
are in order.

Specimens examined. — A total of 156 specimens exam-
ined from 25 to 224 mm SL. Outer Baja California:

LACM W55-92, 1 (131); USNM 125347, 2 (101-119); CAS SU 19107, 1
(160); SIO 60-296, 1 (142); LACM W52-195, 1 (223); USNM 167563, 2
(154-177); LACM 3667, 1 (115); SIO 65-175, 7 (97-108); SIO 64-78, 1
(182). GULF OF CALIFORNIA: USNM 165509, 2 (130-133); FMNH
62751. 4 (119-139); GCRL 2593. 5 (155-202); FMNH 63062, 2 (131-154);
FMNH 63063, 2 (109-150); FMNH 57546, 1 (140); USNM 167562, 9 (122-
190); LACM 9602-1, 1 (110); SIO 62-236, 10 (94-224); SIO 72-60, 8 (122-
145); SIO 70-90, 4 (169-217); SIO 70-70, 1 (141); SIO 65-304, 2 (171-173);
USNM 216384, 1 (134); USNM 211361, 1 (150); USNM 211411, 1 (185);
USNM 183980. 13 (118-140); UA 66-69, 1 (81). CENTRAL MEXICAN
COAST: LACM 30121-10, 1 (118); FMNH 70807, 6 (166-183); CAS (Ace.
No. 1951-111:7), 1 (184); CAS 2327, 1 (68); INP LB-1341, 1 (164); SIO 65-
164.1i69k GUATEMALA: UCR 463-48, 6 (125-173). COSTA RICA:
LACM 9754-24, 5 (105-156); LACM 6917-12, 1 (133); LACM 30710-22, 1
(127): LACM 30713-6, 1 (144); LACM 30728-7, 1 (164); LACM 30717-3, 1
(156). PANAMA: SIO 64-365, 2 (103-109); SIO 63-295, 7 (76-189); SIO
69-386, 2 (113-120); LACM 31310-20, 2 (134-137); MCZ 46148, 6 (157-

211); UMML 26229, 1 (197); UMML 25069. 4 (160-168); UMML 25068, 1
(160); UMML 26248, 1 (174); USNM 81756, holotype, 1 (72); USNM
80228, paratypes, 3 (33-46); LACM 6509-23, 17 (25-148); UP (6 Dec.
1971), 1 (165); USNM (Ace. No. 247715), 7 (130-149). COLOMBIA:
USNM 211392, 1 (149); USNM 211370 1 (125); USNM 211353, 1 (150);
USNM 211382, 1 (148); USNM 211395, 1 (202); USNM 211400, 4 (124-
162); USNM 211356, 1 (142); USNM 211380, 3 (180-189).

Diplectrum rostrum Bortone 1974

Figures 6, 8, 9, 13

Diplectrum rostrum Bortone 1974:61-65, fig. 1-2 (original
description, eastern Pacific); Rosenblatt and Johnson
1974:191 (systematics).

The species is not common within its range and per-
haps some references pertaining to D. pacificum may in
fact refer to D. rostrum.

Diagnosis. — Diplectrum rostrum is distinct from other
species of Diplectrum in that it has two light bars on each
side of the snout and by the unique marking on the soft
dorsal fin which consists of a transverse row of seven to
nine light spots on a dark field. Confusion with D.
pacificum might arise because of the similar preopercle
shape. However, these two species can easily be
separated by the already mentioned color patterns; gill
raker count, 20.25 (17-24) in D. pacificum, 23.44 (22-25)
in D. rostrum; orbit length, 78.52 (66-90) in D. pacificum
and 90.68 (81-100) in D. rostrum ; cheek scale rows A, 9.70
(8-12) in D. pacificum and 8.52 (8-10) inD. rostrum. Also
the caudal pigmentation is a series of dark and light
bands in D. pacificum and a spotted (light on dark) pat-
tern in D. rostrum . The preopercular spur tends to have a
more rounded lower edge in D. rostrum. Other notable
differences between D. rostrum and D. pacificum are:
the caudal lobe tends to be more elongate in D. rostrum
as evidenced by its greater total length (1,320 for D.
rostrum. 1,289 for D. pacificum); D. rostrum has a
shorter postorbital length (193.35 vs. 211.05); a larger or-
bital width (71.50 vs. 62.58) and a shorter caudal pedun-
cle depth (120.92 vs. 129.32) than D. pacificum.

Description.— D, X, 12 (13); A, III, 7 (8); P, , 16.94 (16-
18); pored lateral-line scales, 49.00 (48-51); scale rows in
lateral line, 64.62 (60-70); scales above lateral line, 7.54
(7-9); scales below lateral line, 16.76 (15-18); predorsal
scales, 14.16 (12-16); cheek scale rows A, 8.52 (8-10);
cheek scale rows B, 11.18 (10-12); gill rakers, 23.44 (22-
25).

Standard length, 120.50 (21.78, 78.8-166, 26). Total
length, 1,320 (23.08, 1,277-1,360, 25); fork length, 1,165
(13.13, 1,133-1,186, 26); body depth, 288 (16.15, 251-310,
26); head length, 369 (9.49, 354-388, 26); postorbital
length, 193 (7.22, 183-206, 26); snout length, 103 (6.98,
88-117, 26); lacrimal width, 51 (4.14, 43-58, 26); maxil-
lary width, 27 (2.21, 24-33, 26); orbit length, 91 (5.41, 81-
100, 25); orbit width, 72 (3.92, 66-78, 25); interorbital
width, 74 (6.89, 66-102, 26); upper jaw length, 164 (5.66,
150-173, 26); lower jaw length, 175 (5.75, 162-187, 26);

30

cheek height, 119 (6.24, 110-129, 26); cheek length, 122
(4.96, 112-129, 26); caudal peduncle depth, 121 (5.19,
109-130, 26); pectoral fin length, 257 (14.30, 233-292, 26);
pelvic fin length, 214 (11.56, 186-237, 26); predorsal
length, 360 (10.17, 342-385, 21); pelvic origin to lower
jaw, 336 (14.46, 314-367, 19); pelvic origin to anus, 321
(20.32, 291-382, 24); postanal length, 397 (14.83, 369-421,
26). Dorsal spine height: first, 57 (6.81, 37-68, 26); sec-
ond, 83 (7.80, 70-94, 24); third, 129 (9.12, 113-150, 25);
fourth, 143 (8.05, 130-159, 23); fifth, 145 (8.92, 130-162,
25). Anal spine height: first, 32 (4.02, 19-37, 26); second,
59 (5.57, 46-66, 26); third, 80 (5.85, 66-89, 26).

Preopercle with narrow spur bearing 7-11 spines at
angle. Upper spur edge somewhat squared off, lower edge
more rounded. Vertical and horizontal preopercular
limbs serrate. Caudal fin slightly forked, the upper lobe
extending further posteriorly than lower lobe, almost as a
filament. Chest scales not embedded but reduced in size
anteriorly. Anterior nostril with elongate flap. Gill rakers
moderately long, the first at the angle reaching to fifth
below.

Color. — Characteristic color patterns remain dis-
cernible in preserved specimens. Snout gray-brown with
two lighter bands, perhaps pinkish in fresh specimens (J.
E. McCosker, pers. commun.), passing from anterior or-
bital edge forward to anterior lacrimal edge. Cheeks with
somewhat less distinct wavy bar of lighter pigment. Soft-
dorsal-fin membrane bearing color pattern consisting of
seven to nine light spots on dark field which traverses fin
laterally. Distal edge of fin not pigmented but a lightly
pigmented band near base of fin. Dorsal spines tipped

with black. Caudal peduncle blotch equal to eye size, but
irregular in outline. Dorsal surface of caudal peduncle
with two dark spots separated by a lighter spot. Caudal
fin bearing series of small light spots on dark back-
ground. Lateral body surface with five to seven indis-
tinct darkened vertical bars appearing darker dorsally.
The lateral body surface also traversed by an indistinct
midlateral stripe.

Size. — The species is of medium size; a maximum of
166 mm SL recorded in the present study.

Habits. — Virtually nothing is known of the habits of D.
rostrum except what could be ascertained from collec-
tion data. It has been collected at depths ranging from 13
to 80 m and does not appear to be abundant anywhere.

Distribution. — Collection data indicate that D. rostrum
occurs on the western side of the Gulf of California; at
Bahia de Banderas, Mexico; Puntarenas, Costa Rica;
Gulf of Panama; the Galapagos Islands; and along the
northwest coast of South America (Fig. 13). The species
ranges from lat. 23°34.3'N, long. 110°23.9'W along the
outer Baja California Sur to lat. 3°39'S, long. 80°41'W in
the south.

Variation. — No significant morphological or meristic
variation was noted for this species except for number of
scale rows above the lateral line. A group of northern
specimens (from Bahia de Banderas northward) had a
mean value of 63.10 for scale rows above the lateral line
while the southern group had a mean value of 67.08 for

30"

o°-

Figure 13.— Distribution map for Diplectrum rostrum
(closed circles); and D. macropoma (open
circles). [Localities are based on the materials ex-
amined in this studv.l

30°~

31

this same character. Insufficient material was available
throughout the range of the species to permit speculation
about the nature of this variation.

Specimens examined. — A total of 36 specimens from 60
to 166 mm SL. Outer Baja California: SIO 64-867, paratype,

1 (132). GULF OF CALIFORNIA: LACM 64-64, paratypes, 4 (62-128);
SIO 65-349. paratypes, 2 (98-137); SIO 65-257. paratype, 1 (126); SIO 65-
285. paratype. 1 (103); SIO 65-304, paratype, 1 (113); UA 67-65-3, 1 (142);
INP LB1S16. 1 (148); USNM 125347. 1 (71); UA 67-121-1, 2 (63-72); SIO
70-62, paratype. 1 (71). BAHIA DE BANDERAS. MEXICO: SIO 65-160, 6
(60-104). COSTA RICA: LACM 30714-12, 2 (134-142). PANAMA:)
USNM llat. 7°33'N. long. 78°11'W), 1. COLOMBIA: USNM 208199,
paratype, 1 (87): USNM 208198, paratype. 1 (144); CAS 15296, paratype,
1 (145). ECUADOR: USNM 208196, holotype, 1 (131); USNM 208197,
paratypes, 5 (100-130); CAS 15295. paratype. 1 (117). GALAPAGOS:
FMNH 41433, 1 (166).

Diplectrum conceptione (Valenciennes 1828)

Figures 6, 8, 9, 12

Serranus conceptionis Valenciennes in Cuvier and Valen-
ciennes 1828:246 (original description, Conception,
Chile); Guichenot 1848:148-149 (description and color
notes, Chile); Boulenger 1895:296-297 (catalog); Porter
1909:8 (Chile).

^Plectropoma paytensis Lesson 1830:233-244 (original
description, Peru); Giinther 1859:165 (catalog).

Serranus conceptionis Lesson 1830:236 (Chile).

Centropristis conceptionis Giinther 1859:84 (catalog).

Diplectrum conceptione Jordan and Eigenmann
1890:397-399 (systematics); Abbott 1900:349 (Callao,
Peru); Delfin 1901:62-63 (Chile and Peru); Evermann
and Radcliffe 1917:75-76 (in part, Peru); Fowler
1945b:217 (Peru); Fowler 1945c:78 (Chile); Hildebrand
1946:185, 186-187 (description, Peru); Orces 1951:304
(Ecuador, as cited by Koepke 1963); Mann F. 1954:65,
81, 223 (description, economics, Chile); Koepke 1963:
241 (synonomy, Peru); Chirichigno F 1974:294-298, fig.
576 (key, Peru); Rosenblatt and Johnson 1974:178-191
(key, systematics).
Hemilutjanus paytensis Jordan and Eigenmann
1890:345 (systematics).

Diplectrum conceptione was described from a single
specimen by Valenciennes in 1828. Since that time only
slight confusion has resulted regarding the correct iden-
tity of the species. Lesson (1830:233-244) described a new
species, Plectropoma paytensis, from a drawing. He did
not believe that Valenciennes (Cuvier in Lesson's inter-
pretation) had described the species. It would appear
from the species description of Lesson that he may have
been referring to D. conceptione but several doubts exist
concerning the inclusion of Lesson's species in the syn-
onymy. First, Lesson was aware of the Serranus concep-
tionis description by Valenciennes (Lesson 1830:236).
The collection label indicates Lesson collected the
specimen so therefore why did he state that P. paytensis
had been overlooked by Cuvier? Second, Lesson stated
that the species is common near Paita, Peru, but others
(Mann F. 1954:223; Hildebrand 1946:187) imply D. con-

ceptione is infrequently seen. The color description of P.
paytensis does not closely fit other color descriptions for
D. conceptione (e.g., Evermann and Radcliffe 1917:75-
76). The inclusion of Plectropoma paytensis, Giinther
(1859:165) and Hemilutjanus paytensis Jordan and
Eigenmann (1890:345) in the synonomy for D. concep-
tione may in fact not be justified because Plectropoma
paytensis may be a nomen nudum.

Fowler (1945c:78-79) listed Pristipoma conceptione as
a synonym of D. conceptione. This action was not jus-
tified in that characters given by Steindachner (1875:6-8)
and Guichenot (1848:195-196) as referred to in Fowler's
citation of Schmeltz (1869:13) and Cuvier and Valen-
ciennes (1828:268) exclude this species from the genus
Diplectrum on the basis of dorsal- and anal-fin ele-
ments.

Diagnosis. — Diplectrum conceptione is readily distin-
guishable from other Diplectrum in having two to four
strong spines at the angle of the preopercle and a high
number of pectoral rays (modally 18). The rather long sec-
ond anal spine can confuse this species only with D.
maximum from which it can be distinguished by the
above characters, color patterns, number of gill rakers,
and number of cheek scale rows A.

Description.— D, X, 12 (11-13); A, III, 7; P: , 18.16 (17-
19); pored lateral-line scales, 49.84 (48-51); scale rows in
lateral line, 68.82 (61-75); scales above lateral line, 7.92
(7-9); scales below lateral line, 19.93 (17-23); predorsal
scales, 13.98 (12-18); cheek scale rows A, 8.49 (7-10);
cheek scale rows B, 11.60 (9-14); gill rakers, 25.28 (23-28).

Standard length, 107.77 (23.00, 80-161, 39). Total
length, 1,275 (17.45, 1,236-1,318, 37); fork length, 1,187
(13.78, 1,160-1,214, 39); body depth, 277 (12.72, 254-305,
34); head length, 379 (10.74, 361-408, 39); postorbital
length, 213 (7.48, 202-237, 39); snout length, 94 (7.07,
79-106, 39); lacrimal width, 37 (3.48, 32-46, 39); maxil-
lary width, 34 (1.89, 30-37, 39); orbit length, 90 (6.05, 82-
101, 39); orbit width, 75 (5.81, 62-87, 39); interorbital
width, 69 (3.18, 63-76, 38); upper jaw length, 159 (6.12,
145-173, 39); lower jaw length, 182 (5.15, 169-196, 39);
cheek height, 111 (6.01, 98-127, 39); cheek length, 108
(5.17, 96-119, 39); caudal peduncle depth, 116 (4.32, 108-
128, 39); pectoral fin length, 256 (9.98, 235-279, 39); pel-
vic fin length, 215 (8.18, 196-230, 39); predorsal length,
365 (10.34, 342-399, 38); pelvic origin to lower jaw, 336
(11.96, 317-363, 31); pelvic origin to anus, 317 (19.88, 282-
369, 37); postanal length, 401 (12.10, 381-438, 37). Dor-
sal spine height: first, 46 (5.23, 38-57, 37); second, 72
(7.20, 57-87, 39); third, 147 (10.17, 127-165, 35); fourth,
158 (10.56, 132-183, 32); fifth, 150 (9.23, 126-168, 37).
Anal spine height: first, 45 (4.70, 37-54, 39); second, 84
(8.69, 66-100, 39); third, 95 (8.19, 76-113, 39).

Preopercle with a group of two to four strong spines at
angle. Few serrae present on ventral edge, many on ver-
tical edge of preopercle. Gill rakers long and slender, the
first reaching the base of fifth below. Anterior nostril
without distinct flap. Scales on operculum in definite
rows. Small scales running down membrane between

32

third spine and first soft anal ray. Also, small scales
along base of anterior soft dorsal rays. Second anal spine
more stout than third. Caudal fin nearly straight.

Color. — Color description is from preserved specimens.
Pelvic fin, ventral edge light, pale along medial base and
edge, body of fin appearing dusky, with dark pigment
mainly on interradial portions. Pectorals with line of
dark pigment along the dorsal edge of each ray, no pig-
ment interradially. Anal fin pale. Caudal fin dusky to
dark. Spinous dorsal membrane dusky, distal edge of
membrane white with black at extreme distal edge. Base
of spinous dorsal less dusky than body of membrane. Soft
dorsal fin with a broad pale distal margin, a darker band
below, and below this a dusky field bearing a group of ir-
regular light spots fusing into diagonal bars. This pat-
tern of light spots and bars quite variable and oc-
casionally appearing only as spots or only as horizontal
light bars on the soft dorsal. Caudal spot present but in-
distinct and smaller than orbit. Ventral portion of body
light brown. Dorsum medium brown, bases of soft dorsal
rays black. Lateral body surface with nine indistinct
dusky vertical bars terminating above midventral line.
Evidence of interconnecting midlateral stripes on body.
Lateral and medial opercular surfaces black. Posterior
branchial cavity black. Slight evidence of small indis-
tinct spots of light pigment on lacrimal area, about five
in number, and may interconnect. Lower premaxillary
border with dark edge, remainder of premaxillary dusky.
Center tip of lower jaw dusky. No evidence of pigment in
upper pharyngeal area.

The following color description of live D. conceptione
is quoted from Evermann and Radcliffe (1917:75-76):
"Color in life: Back and upper part of sides dark olive
green, but with three pale horizontal stripes; one, from
just above pectoral to just beneath lateral line on pedun-
cle; a second, from level of eye, and beginning a short dis-
tance posterior to eye, to upper part of peduncle; a third,
at a corresponding distance above the second; a darker
mottling on the sides gives an indistinct effect of cross-
barring, especially on upper part of sides; sides, below
lowest stripe, dusky, greenish with much gold; below
white with a tinge of orange along median line of belly;
head above olive with reddish spots, orange on lower part
of sides; under side of opercle with jet black and gold;
skin posterior to fourth gill blue-black; caudal mostly
reddish orange, ventral margin pale; dorsal light olive
with large spots of bright orange; a little black on tips of
membrane between dorsal spines; pectorals olivaceous;
ventrals mixed dusky olive and orange; anal mostly
white, but with some orange on membrane between con-
secutive rays."

Size. — Maximum size of specimens examined is 169
mm SL.

Habits. — The species seems to prefer fairly deep water
with only one collection of specimens from 13 m and the
remainder from 33 to 80 m deep. Little is known of the
habits of this species.

Distribution. — Diplectrum conceptione is limited in its
distribution to the coasts of Ecuador, Peru, and Chile in
the eastern Pacific (Fig. 12). The northernmost locality is
at lat. 3°15'S and its southern limit is also the type-
locality at Conception, Chile (lat. 37°S).

Variation. — Most material examined of this species is
from the same general area off the coast of Peru; there-
fore, no variation was discernible over the species range.

Specimens examined. — A total of 230 specimens from 69
to 169 mm SL. Ecuador: USNM 213574, 72 (96-136); USNM

211418, 1 (135); USNM 211414, 1 (96). PERU: USNM 213583, 60 (69-
161); USNM 213582, 15 (92-138); USNM 211371, 1 (144); USNM 211390,
2 (119-129); USNM 213578 + 211385, 69 (89-169); USNM 77549, 1 (94);
USNM 77541, 1 (111); USNM 127969, 1 (137); CAS SU 6293, 5 (130-
159). CHILE: MNHN 7025, holotype, 1 (102).

Diplectrum maximum Hildebrand 1946

Figures 6, 8, 9, 11

Diplectrum conceptione Evermann and Radcliff 1917:75-
76, pi. 7, fig. 2 (in part, figure and one text description
is of D. maximum, Peru).

Diplectrum maximum Hildebrand 1946:185-187, fig. 42
(original description, Paita, Peru); Orces 1951:304 (not
seen, Ecuador); Orces 1959:81 (D. maximum possibly
a subspecies of D. pacificum); Rosenblatt and John-
son 1974:178-191 (description of juveniles, Baja
California); Chirichigno F. 1974:294-298, fig. 577 (key,
Peru).

Diplectrum pacificum maximum Koepke 1963:241
(Peru).

The larger specimen described and figured by Ever-
mann and Radcliffe (1917:75-76, pi. 7, fig. 2) is USNM
77698. This specimen was examined and is most cer-
tainly D. maximum. One of the other specimens to which
they referred (Evermann and Radcliffe field no. 09529)
was designated as the holotype as figured by Hildebrand
(1946:185-187, fig. 42). The other specimens they referred
to (Evermann and Radcliffe field nos. 09543 and 09547)
were examined and are D. conceptione bearing USNM
numbers 77549 and 77541, respectively. Koepke
(1963:241) appears to have accepted the suggestion of
Orces (1959:81) that D. maximum may be a subspecies of
D. pacificum. Orces (1959:81) indicated they were closely
aligned because of similar number of scales on the
"preopercle" (i.e., cheek) and the caudal fin shape. The
indiscriminate acceptance of the suggestion by Orces
seems to have led to the recognition of an invalid sub-
species of D. pacificum.

Diagnosis. — Diplectrum maximum is the largest of all
known Diplectrum, reaching 302 mm SL. It can be dis-
tinguished from D. conceptione by its lower mean total
gill raker count, 19.28 (17-21), as opposed to 25.28 (23-28)
for D. conceptione; cheek scale rows A tend to be higher,
10.07 (9-11) than those of D. conceptione, 8.49 (7-10);
and the preopercle of D. conceptione bears two to four

33

strong spines while that of D. maximum is in the shape of
an elongate bony flap bearing four to eight longer and
thinner spines. The species also can be separated on the
basis of color patterns described below. Confusion with
D. pacificum should not occur due to the distinctive
preopercular shape of D. maximum. Also the second anal
spine is notably stout and nearly equal (longer in
juveniles) in length to the third in D. maximum, but the
second anal spine is much shorter than the third in D.
pacificum.

Description.— D. X, 12; A, III, 7; P! , 16.93 (16-17); pored
lateral-line scales, 49.36 (48-50); scale rows in lateral
line. 66.33 (61-72); scales above lateral line, 8.47 (7-10);
scales below lateral line, 21.13 (18-25); predorsal scale
rows. 16.43 (14-20); cheek scale rows A, 10.07 (9-11);
cheek scale rows B, 11.47 (10-13); gill rakers, 19.28 (17-
21).

Owing to the unusual nature of the size disparity in-
volved with this species and because so few specimens
were available for examination it was decided to present
the data for each character in the following manner:
character data for the eight larger specimens of D. max-
imum. 245.38 mm SL (32.56, 205-302, 8) followed im-
mediately by character data for the seven smaller
specimens, 64.43 mm SL (17.31, 45.9-93.2, 7); data for
larger specimens are separated from data for smaller
specimens by a hyphen. Total length, 1,245 (16.77, 1,221-
1,270, 8) - 1,224 (20-21, 1, 191-1,248, 6); fork length, 1,178
(13.09, 1,162-1,200, 8) - 1,189 (14.15, 1,168-1,207, 7); body
depth, 285 (19.38, 257-319, 7) - 257 (13.99, 241-276, 7);
head length, 363 (4.74, 354-369, 8) - 361 (7.20, 350-373, 7);
postorbital length. 211 (3.73, 203-214, 8) - 189 (2.16, 187-
192. 7); snout length. 102 (2.82, 98-106, 8) - 91 (6.34, 80-
98. 7); lacrimal width, 54 (3.01, 50-59, 8) - 33 (3.56, 28-39,
7); maxillary width, 30 (2.25, 26-32, 8) -'35 (2.45, 31-38,
7); orbit length, 67 (3.36, 62-72, 8) - 99 (7.03, 87-108, 7);
orbit width, 50 (4.10, 46-57, 8) - 76 (6.85, 66-87, 7); in-
terorbital width, 72 (3.31, 67-76, 8) - 73 (3.30, 70-78, 7);
upper jaw length, 159 (6.27, 149-167, 8) - 158 (5.19, 152-
168, 7); lower jaw length, 173 (6.75, 164-180, 8) - 181
(1.91, 179-184, 7); cheek height, 130(4.78, 121-134, 8) - 98
(5.71, 87-105, 7); cheek length, 97 (5.18, 87-103, 8) - 111
(5.59, 106-122, 7); caudal peduncle depth, 124 (4.10, 120-
130, 8) - 118 (1.80, 116-121, 7); pectoral fin length, 234
(6.58, 225-242, 8) - 230 (7.21, 222-239, 7); pelvic fin
length, 202 (7.77, 198-212, 8) - 196 (13.86, 175-209, 7);
predorsal length, 352 (10.50, 336-370, 8) - 366 (10.05, 349-
374, 6); pelvic origin to lower jaw, 340 (14.21, 326-371, 8)
- 337 (9.88, 320-344, 5); pelvic origin to anus, 322 (22.94,
295-352, 8) - 286 (15.14, 255-301, 7); postanal length, 403
(13.54. 380-419, 8) - 406 (11.07, 389-420, 7). Dorsal spine
height: first, 66 (7.49, 54-76, 8) - 77 (4.80, 73-82, 5); sec-
ond, 83 (9.85, 67-96, 8) - 95 (7.98, 84-105, 5); third, 128
(9.01, 117-144, 7) - 132 (6.56, 125-143, 6); fourth, 137
(11.63, 115-154, 8) - 146 (6.68, 133-153, 6); fifth, 135
(10.08, 118-150, 8) - 140 (2.88, 137-144, 6). Anal spine
height: first, 41 (3.74, 36-46, 8) - 56 (8.10, 45-70, 7); sec-
ond, 68 (2.83, 64-72, 8) - 91 (4.30, 85-98, 7); third, 74
(4.07, 68-80, 8) - 88 (5.21, 81-94, 7).

Preopercle with a notable elongate shape bearing four
to eight elongate spines at its angle. Vertical and
horizontal propercular arms serrate. Second anal spine
very strong, much stronger than third. First gill raker at
the angle reaching to the base of the fourth below.
Anterior nostril with elongate flap. Rows of small scales
along third anal spine, first anal ray, and first and second
membranous portions of soft-dorsal -fin base. Thoracic
scales smaller, slightly embedded. Caudal fin nearly
truncate.

Color. — Color description for larger preserved
specimens: anterior lateral edges of pelvic fins light, fin
dusky but paler at base and on medial rays. Pectoral fin
rays dusky, membrane pale. Anal fin pale. Distal edges
of caudal fin black, fin brown with a dusky dorsal edge.
Spinous dorsal dusky between first and second spines,
tips of all spines black, membrane dusky, paler distally.
Soft dorsal with black edge distally, small black spots at
interradial bases. Remainder of membrane pale except
between last two dusky interradials. Body brown, paler
ventrally, slight evidence of indefinite blotches on dor-
solateral surface. Posterior branchial area dusky to
black, medial opercular surface black. Dorsum brown,
slightly darker at anterior base of spinous dorsal. Snout
with a pale wedge-shaped spot anteriorly. Anterior tip of
lower jaw dusky. An indistinct bar of light pigment at
lateral pectoral base.

Color description for smaller preserved specimens:
lateral body surface with two wide brown horizontal
stripes, a narrower stripe of light pigment between these
brown stripes. Below the midlateral brown stripe, a row
of 8-10 pale or white blotches. White spots above and
below the lateral band on caudal peduncle. White
predorsal stripe extending to interorbital area. Ventral
body surface white. Dark pigment on first through fifth
spinous dorsal membranes. Anterior base of pectoral
dusky. Dorsal spines with black tips. Upper pharyngeal
area slightly pigmented.

Habits. — Nothing is known of the habits of D. max-
imum. The larger specimens came from deeper water
(70-128 m) but the smaller specimens were from shallow
water (1-7 m). The species attains considerable size (302
mm SL) and may prefer cooler waters of the subtropics.

Distribution. — Two areas of distribution are noted (Fig.
11): the northern group of smaller specimens with
northernmost distribution lat. 24°44.35'N, long.
111°58.4'W in Bahia de Magdalena; the southern group
of larger specimens found along the Peruvian-Ecuador
coast having as its southernmost point of distribution
Lobos de Tierra, Peru. The type-specimens are from
Paita and Lobos de Tierra, Peru.

Variation. — The small sample sizes and the disjunct dis-
tribution of specimen size did not allow for satisfactory
studies on variation for this species. However, the few
meristic data available do not indicate any obvious
variations between the two known distribution centers.

34

Rosenblatt and Johnson (1974) noted that juveniles
are found only off the outer Baja California coast and
possess a jet black blotch on the anterior portion of the
spinous dorsal fin membrane. While this character is
lacking in adults of D. maximum, there is evidence of
some, but less intensified, pigmentation on this fin.
Rosenblatt and Johnson (1974) suggested there can be no
further speculation as to an ontogenetic or genetic ex-
planation for this noted variability until additional
material is available.

There is some question concerning the species integrity
of D. maximum because of the extreme geographical and
size differences in the two populations considered. Ex-
amination of additional material (presently unavailable)
of intermediate size and parts of its range may warrant
reevaluation of the present scope of D. maximum.

Specimens examined. — A total of 15 specimens from 46
to 302 mm SL. Outer Baja California: SIO 62-726, 2 (46-50);

SIO 62-106, 1 (69); SIO 62-706, 2 (74-75); SIO 64-63, 1 (93); SIO 64-869, 1
(50). ECUADOR: USNM 213581, 2 (260-275). PERU: USNM 77624,
holotype, 1 (241); USNM 77698, paratype, 1 (222); USNM 200373, 1
(302); USNM 213809, 2 (205-243); USNM 211407, 1 (215).

Diplectrum sciurus Gilbert 1892

Figures 6, 8, 9, 10

Diplectrum sciurus Gilbert 1892:550-551 (original de-
scription. Gulf of California; Jordan and Evermann
1896:1203-1204 (systematics); Osburn and Nichols
1916:162 (Baja California); Jordan et al. 1930:318
(check list); Walford 1937:123 (general account);
Rosenblatt and Johnson 1974:178-191 (key, sys-
tematics).

Serranus sciurus Boulenger 1895:298 (catalog).

The USNM has three syntypes listed as designated by
Gilbert: USNM 46630 (an 87-mm-SL specimen in good
condition) is here designated the lectotype in the ab-
sence of any known designated holotype; USNM 46571
(an incomplete skeleton) and USNM 125348 now become
paralectotypes.

(16-18); pored lateral-line scales, 49.18 (47-52); scale
rows in lateral line, 70.76 (63-79); scales above lateral
line, 8.17 (7-10); scales below lateral line, 19.18 (17-21);
predorsal scale rows, 15.48 (12-19); cheek scale rows A,
7.66 (6-9); cheek scale rows B, 13.44 (12-17); gill rakers,
36.64 (32-41).

Standard length, 95.04 (10.40, 76.8-116, 33). Total
length, 1,263 (16.78, 1,228-1,298, 33); fork length, 1,172
(11.94, 1,149-1,201, 32); body depth, 269 (13.15, 245-296,
33); head length, 356 (9.87, 333-376, 33); postorbital
length, 186 (5.99, 171-198, 33); snout length, 89 (5.08, 79-
97, 33); lacrimal width, 27 (2.43, 23-33, 33); maxillary
width, 36 (2.58, 30-41, 33); orbit length, 92 (4.31, 84-100,
33); orbit width, 78 (4.28, 71-90, 33); interorbital width,
79 (3.70, 70-84, 33); upper jaw length, 142 (5.69, 130-159,
33); lower jaw length, 172 (6.68, 159-191, 33); cheek
height, 95 (3.40, 86-103, 33); cheek length, 122 (3.47, 116-
129, 33); caudal peduncle depth, 105 (3.39, 98-112, 33);
pectoral length, 254 (10.90, 234-275, 33); pelvic length,
211 (6.31, 194-224, 33); predorsal length, 343 (7.73, 330-
361, 32); pelvic origin to lower jaw, 351 (9.20, 331-366,
30); pelvic origin to anus, 336 (16.79, 302-363, 32);
postanal length, 372 (8.82, 355-388, 33). Dorsal spine
height: first, 62 (5.61, 53-74, 32); second, 88 (5.95, 80-101,
31); third, 125 (9.93, 110-157, 32); fourth, 142 (9.54, 125-
167, 32); fifth, 146 (9.42, 129-166, 33). Anal spine height:
first, 31 (3.75, 24-40, 32); second, 59 (6.06, 47-73, 33);
third, 86 (6.44, 76-99, 32).

Diplectrum sciurus is a highly specialized member of
the genus with a high number of elongate gill rakers,
broad maxillary, narrow lacrimal, and short upper and
lower jaws. Distal tip of maxillary arm not exceeding
midpoint of eye on a vertical plane. Preopercle somewhat
square in outline bearing a single cluster of 5-10 en-
larged spines on spur. Vertical and horizontal preoper-
cular arms serrate. Second anal spine as strong as third.
Distally the maxillary very broad and spatulate. Lac-
rimal wedge-shaped, broadest anteriorly. Anterior nos-
tril with low flap perhaps absent in some specimens.
Opercular scales small. Chest scales present and well
developed. Caudal fin slightly forked, upper lobe ex-
tending further posteriorly than lower.

Diagnosis. — Diplectrum sciurus is immediately recog-
nized by its numerous and elongate gill rakers (the first
reaching almost to the base of the 12th below). The
species also has the least caudal peduncle depth, least
lacrimal width, and greatest maxillary width of any
Diplectrum. It is the only Diplectrum whose maxillary
width is consistently greater than its lacrimal width as
an adult. The upper- and lower-jaw lengths are also the
least of any Diplectrum. The species has the least cheek
height and least postanal length. The species has eight
anal rays which separates it from all other Diplectrum
except D. euryplectrum and outer Baja populations of D.
labarum. Again the high number of gill rakers for D.
sciurus facilitates identification.

Description.— D, X, 12 (11-13); A, III, 8 (7); Pj , 16.96

Color. — Color notes are based on preserved specimens.
Dorsum and lateral body surfaces light brown. Lateral
body surface with two wide brown longitudinal stripes ir-
regularly interrupted by light brown flecks. Eight to ten
brown indistinct vertical bars along sides, extending
about four-fifths the body depth down the sides. Ventral
surface pale. Premaxillary dusky, like lower jaw at mid-
line. Top of head dark brown. Caudal fin with irregular
pattern of small, pale blotches on dusky background.
Upper caudal edge pale, the lower tending to be dusky.
Anal, pectoral, and ventral fins pale. Spinous dorsal fin
with three rows of large, irregular, pale blotches on
darker field. Blotches present along the distal edge, mid-
laterally, and along base of spinous dorsal fin mem-
brane. Soft dorsal with two midlateral rows of pale
blotches on darker field. Often blotches fusing together

35

to form a wavy, pale bar. Distal, soft dorsal edge clear.
No apparent cheek or snout bars on preserved specimens.
Caudal peduncle spot present but small, irregular in out-
line, appearing as intensified portion of midlateral body
bar. Upper pharyngeal, medial opercular, and posterior
branchial surfaces lacking pigment.

Gilbert (1892:550-551) stated that a light blue line ran
just below the orbit and that the soft dorsal and caudal
fins bore yellow spots surrounded by pale blue.

Size. — The species is small in size, reaching no longer
than 137 mm SL in the present study.

Habits. — The extreme modifications of the jaws, gill
rakers, and lacrimal area suggest that D. sciurus may
have different feeding habits than other Diplectrum. Al-
though no data are yet available, the species probably
feeds on small crustaceans and generally may live just off
the bottom as opposed to the benthic habits known for
the other species. The species has been taken at depths
ranging from 25 to 100 m. The few data available do not
indicate any depth preference.

Distribution. — Diplectrum sciurus is found almost ex-
clusively along the coastal areas of the Gulf of California
(Fig. 10). It does occur along the outer Baja California to
lat. 24°35'50"N, long. 112°05'00"W near Bahia de Mag-
dalena. The type-locality is lat. 31°22'00"N, long.
114°07'00"W from the upper Gulf of California.

Variation. — The species does not show any obvious
geographical variation due to its limited distribution.

Specimens examined. — A total of 201 specimens from 67
to 137 mm SL. Gulf of California: USNM 46630, lectotype, l

(87); USNM 125348, paralectotype, 1 (92); USNM 46571, paralectotype,
1 (incomplete, disarticulated skeleton); LACM 8825-7, 79 (68-98); LACM
8824-7, 22 (84-116); LACM 8339-4, 5 (94-99); LACM 8834-1, 1 (98); CAS
W 53-93. 1 (104); CAS SU 53100, 1 (105); FMNH 62747, 1 (90); CAS W
59-17. 4 (77-103); SIO 60-120, 17 (120-137); SIO 70-225, 28 (100-123);
SIO 68-71, 36 (67-110). OUTER BAJA CALIFORNIA: CAS SU 243. 3 (92-
97).

Diplectrum euryplectrum
Jordan and Bollman 1890

Figures 6, 8, 9, 11

Diplectrum euryplectrum Jordan and Bollman 1890:157-
158 (original description, Galapagos); Jordan and
Eigenmann 1890:397-398 (review); Jordan and Ever-
mann 1896:1203, 1206 (systematic account); Gilbert
and Starks 1904:98 (Panama); Meek and Hildebrand
1925:472, 474 (Panama); Jordan et al. 1930:318 (check
list); Walford 1937:124 (general account); Hildebrand
1946:184 (key); Chirichigno F. 1974:294-298 (key,
Peru); Rosenblatt and Johnson 1974:178-191 (key, sys-
tematics).

Diplectrum macropoma Gilbert and Starks 1904:97-98
(in part, misidentified, Gulf of California).

Diagnosis. — This species is easily recognized by the
broad spur at the preopercular angle, eight anal rays
(rarely seven or nine) and a dark band of pigment on the
soft-dorsal-fin base. It has the greatest fork length, head
length, postorbital length, eye diameter, interorbital
width, upper- and lower-jaw length, cheek height and
length, and pelvic-origin-to-lower-jaw dimensions of any
Diplectrum. It is a distinct species but juveniles of D.
euryplectrum might possibly be confused with juveniles
of D. labarum. Except for a population of D. labarum
from outer Baja California which has a high proportion of
individuals with eight anal rays they could normally be
separated on this character alone. Scales tend to be
smaller in D. euryplectrum than in D. labarum (Tables
2-7). Also, total number of gill rakers is higher in D. eury-
plectrum (23.68, 21-27) and lower in D. labarum (21.52,
19-24). The only other species having eight anal rays isD.
sciurus but this species is easily distinguished by the
high total gill raker count.

Description.— D, X, 12 (13); A, III, 8 (7-9); Pj , 17.62 (16-
18); pored lateral-line scales, 48.77 (46-52); scale rows in
lateral line, 64.12 (56-73); scales above lateral line, 7.71
(7-10); scales below lateral line, 17.81 (15-20); predorsal
scales, 13.73 (10-18); cheek scale rows A, 7.97 (5-10);
cheek scale row B, 10.49 (8-13); gill rakers, 23.68 (21-27).

Standard length, 111.62 (13.75, 76.8-143, 39). Total
length, 1,260 (15.56, 1,217-1,293, 36); fork length, 1,193
(12.92, 1,155-1,222, 38); body depth, 291 (1,278, 261-313,
38); head length, 423 (12.99, 384-449, 39); postorbital
length, 241 (10.47, 215-264, 39); snout length, 103 (7.96,
91-117, 39); lacrimal width, 46 (3.89, 39-53, 20); maxil-
lary width, 36 (1.96, 32-39, 20); orbit length, 102 (4.74,
90-111, 39); orbit width, 84 (4.78, 72-93, 39); interorbital
width, 88 (5.89, 78-98, 20); upper jaw length, 189 (6.00,
174-198, 39); lower jaw length, 206 (5.95, 198-222, 39);
cheek height, 141 (6.15, 127-158, 39); cheek length, 130
(5.69, 117-143, 30); caudal peduncle depth, 106 (4.65, 98-
116, 37); pectoral fin length, 256 (9.98, 235-276, 39); pel-
vic fin length, 212 (9.41, 188-224, 22); predorsal length,
375 (9.10, 363-392, 20); pelvic origin to lower jaw, 361
(11.21, 345-397, 35); pelvic origin to anus, 309 (23.32, 268-
368, 38); postanal length, 374 (10.22, 356-397, 38). Dorsal
spine height: first, 60 (8.90, 42-80, 37); second, 87 (7.34,
71-104, 34); third, 118 (9.36, 102-138, 34); fourth, 132
(9.29, 117-162, 32); fifth, 137 (7.63, 124-154, 31). Anal
spine height: first, 32 (4.77, 25-44, 37); second, 62 (6.94,
49-77, 37); third, 82 (7.24, 70-100, 36).

Preopercle with wide broad spur, bearing 6-12 spines.
Vertical preopercular arm generally lacking serrae with
only a few near base at dorsal point of spur. Head large,
caudal peduncle shallow. Gill rakers moderately long
and slender, the first at the angle reaching to the fifth
below. Gill filaments long. Anterior nostril bears
elongated flap. Scales absent on anal fin base. Chest
scales not noticeably reduced in size. Caudal fin straight
without elongated filament. Second anal spine as strong
as third. Scales somewhat deciduous.

Color. — The following color description is based on

36

three adults and two juveniles obtained live from the
Gulf of Nicoya, Costa Rica, 29 June 1973. Membrane
between first and third dorsal spines pale blue, slightly
darker on spines and distal margin. Remainder of
spinous dorsal fin dusky blue on distal half. Spine tips
orange. Pale brown blotch on middle portion of spinous
dorsal. Soft dorsal fin pale gray, a dark lateral band on
base about one-sixth to one-fifth fin height. Distal, soft
dorsal membrane pale blue. Soft dorsal ray tips pale
orange. Caudal fin dusky with traces of pale yellow on
membranes. Lower caudal edge pale orange. Ventrals
dark gray with same pale yellow on membrane. Pectoral
fin dusky. A dusky midventral stripe present from the
posterior ventral fin base to the anus. Remainder of ven-
tral area pale gray. Dorsum gray-brown interspersed with
small gold flecks in two lateral rows, one on upper dor-
sum, the other more lateral. Snout with indistinct yel-
low bar between anterior edge of orbit to anterior lac-
rimal edge. Another indistinct yellow bar passes from
lacrimal area to just below and along lower orbital edge
diverging into two to four indistinct bars on cheek. Dis-
tal edge of premaxillary yellow. Distal maxillary tip pale
yellow. Lower jaw dusky at midline. Upper pharyngeal
region white. Branchiostegal membrane white. Caudal
peduncle spot and vertical body bars present but not dis-
tinct in specimens examined.

The following color description is from preserved
specimens. Pelvic fin dusky, but paler medially. Pec-
toral fin pale with some dark pigment on dorsal side of
rays. Anal fin pale. Caudal fin dusky, pale dorsal edge,
broad pale ventral edge, posterior border black. Spinous
dorsal with pale base, becoming darker distally. Tips of
first and second spines black. Base of soft dorsal fin with
distinct dusky band about one-fifth fin height. Above the
band soft dorsal dusky with pale distal edge. Dorsum
brown, sides becoming pale to white ventrally. Lateral
body surface with five wide, indistinct, vertical bars with
pale narrow bars between. Caudal spot distinct, equal to
eye size. Lateral and medial opercular surfaces black
with white area below blotch on medial opercular sur-
face. Posterior branchial area pale or slightly pig-
mented. Chest white. Pectoral fin base white with pale
orange bar distal to base. Head brown above, pale below.
An indistinct pattern of pale orange bars on head; some
pale orange on distal maxillary, cheeks, preopercle spur,
and operculum. Indistinct pale orange markings on
snout. Upper pharyngeal area pale. Upper jaw dusky,
lower jaw somewhat paler.

Size. — The largest specimen examined was 163 mm

SL.

Habits. — Diplectrum eury plectrum is found in waters of
the eastern Pacific from 15 to 150 m deep. However, the
species appears to be more commonly taken from 60 to 90
m depth and is abundant at some localities. The species
does not seem to prefer shallow coastal areas.

Distribution. — The species is found along the tropical-
subtropical eastern Pacific coast, in the Gulf of Califor-

nia, and at the Galapagos Islands (Fig. 11). The northern
range of the species ends along the outer Baja California
coast at lat. 24°27'N, long. 111°59'W and extends south
to Ecuador at lat. 01°03'S, long. 80°59'W. The type-
locality is the Galapagos Islands.

Variation. — There is little evidence of geographical
variation in this species except for the number of predor-
sal scales. The differences in this character among three
areas (i.e., Mexico, Panama, and Colombia; Table 5) ap-
pear distinct and may reflect real population dif-
ferences.

Specimens examined. — A total of 152 specimens from 32
to 163 mm SL. Baja California: CAS SU 47, l (43)-, SIO 70-257,

3 (116-122). MEXICO: CAS 4780, 1 (68); CAS 4781, 1 (71); CAS 4782, 1
(69); CAS 4783, 1 (71); USNM 131405, 36 (35-77); GCRL 2495, 1 (94);
SIO 63-513, 25 (62-137); SIO 63-518, 13 (43-86); SIO 63-515, 2 (82-123);
LACM 30123-12, 1 (anterior half only, head length 20 mm). COSTA
RICA: UCR 323-15, 1 (93); UCR 139-14. 1 (120); UCR 323-27, 5 (74-112);
LACM 30718-2, 2 (93-122). PANAMA: USNM uncat. (Pelican 685), 1
(98); USNM 41148, paratype, 1 (108); UMML 26084, 4 (100-137); UMML
26041, 13 (98-126); CAS SU 1458, paratypes, 4 (108-143); CAS SU 1172,
paratypes, 4 (70-137); UMML 26268, 13 (59-117); USNM uncat. (Pelican
760), 5 (115-135); CAS SU 1409, 1 (104); UMML 26255, 16 (57-111);
UMML 26267, 11 (74-112); UMML 26267, 18 (49-66); UMML 28621, 9
(53-115); UMML 26269, 10 (32-55); SIO 71-57, 10 (65-93). GALAPAGOS
ISLANDS: USNM 41141 Holotype, 1 (130); USNM 41141, paratype, 1
(128). ECUADOR: USNM 211389, 1 (113); USNM 213598, 1 (110);
USNM 213572, 30 (123-163); USNM 211372, 1 (126); USNM 211415, 2
(92-95).

Diplectrum labarum
Rosenblatt and Johnson 1974

Figures 6, 8, 9, 14

Diplectrum macropoma Gilbert and Starks 1904:97-98

(in part, misidentification).
Diplectrum labarum Rosenblatt and Johnson 1974:178-

191, fig. 1A. (original description, coastal eastern

Pacific).

Gilbert and Starks (1904:97-98) stated they obtained
D. macropoma from outer Baja California in 1889 at Al-
batross Stn. 3039 and this collection was cataloged as
CAS SU 47 under the identification of D. macropoma.
This collection, in fact, contained D. euryplectrum, D.
eumelum, D. labarum, and D. macropoma.

Diagnosis. — Diplectrum labarum is distinguishable as
an adult from other species of Diplectrum by the
elongate, filamentous tips on the second, third, and
fourth (occasionally the fifth) dorsal spines. Juveniles of
this species might be confused with juvenile D. eury-
plectrum but may be separated by the number of anal
rays (eight in D. euryplectrum, seven in most pop-
ulations of D. labarum). Juvenile D. labarum might also
be confused with juvenile D. eumelum. Diplectrum
labarum tends to have higher scale meristics (Tables 3-
7), especially in number of cheek scale rows B (10.61, 9-
12 in species C; 8.41, 7-10 in D. eumelum).

37

Description.— D, X. 12 (11-13); A, III, 7(8); ^ , 16.61 (16-
18); pored lateral-line scales, 48.14 (47-49); scale rows in
lateral line. 58.94 (51-66); scales above lateral line, 6.98
(6-9); scales below lateral line, 16.93 (14-19); predorsal
scale rows. 13.62 (11-17); cheek scale rows A, 7.34 (6-9);
cheek scale rows B, 10.61 (9-12); gill rakers, 21.52 (19-
24).

Standard length, 116.64 (33.50, 78.4-188, 37). Total
length. 1.271 (21.25, 1,232-1,311, 35); fork length, 1.179
(15.76. 1.142-1.211. 37); body depth, 291 (23.43, 257-341,
35); head length. 381 (11.36, 364-410, 37); postorbital
length. 204 (10.24, 184-227, 37); snout length, 99 (6.95,
85-115. 37); lacrimal width, 42 (5.47, 33-54, 37); maxil-
lary width, 33 (1.82, 28-37, 36); orbital length, 95 (7.34,
78-110. 37); orbit width, 76 (5.58, 67-90, 37); interorbital
width, 77 (4.38. 69-86, 36); upper jaw length, 176 (8.03,
157-197. 37); lower jaw length, 194 (7.47, 176-209, 36);
cheek height, 128 (10.09, 108-147, 37); cheek length, 121
(5.13. 111-132, 37); caudal peduncle depth, 117 (4.50,
110-131, 37); pectoral fin length, 250 (13.42, 221-270, 36);
pelvic fin length, 203 (9.70, 175-221, 37); predorsal
length, 363 (9.43, 347-390, 36); pelvic origin to lower jaw,
349 (8.87, 326-365, 32); pelvic origin to anus, 346 (18.53,
318-376, 34); postanal length, 378 (12.72, 351-401, 34).
Dorsal spine height: first, 61 (10.87, 36-84, 35); second,
140 (22.52, 99-186, 29); third. 170 (20.34, 129-204, 30);
fourth, 165 (11.60, 147-187, 30); fifth, 151 (11.18, 127-175,
32). Anal spine height: first, 39 (4.92, 28-51, 37); second,
69 (7.70, 56-82, 37); third, 91 (7.96, 75-106, 37).

Second, third, and fourth (occasionally the fifth) dor-
sal spines with elongate filamentous tips. Tips develop-
ing gradually during ontogeny beginning about 60 mm
SL and increasing in proportion to standard length
thereafter. Elongate filaments occasionally broken but
spines bearing dark pigment their entire length. Preoper-
cle squared, broader than in D. sciurus but not quite as
broad as in D. eury 'plectrum. Enlarged spines, usually
about 8-13 in number, occurring on spur. Vertical and
horizontal preopercle arms serrate. Anterior nostril with
distinct flap. Caudal fin slightly forked, upper lobe a lit-
tle longer than lower but never extending as elongated
filament.

Color. — Color description is based on preserved
specimens. Dorsum, light brown, becoming darker on
head. Lateral body surface with two indistinct wide
horizontal stripes interrupted by five to seven wide, in-
distinct vertical bars. Bars not extending to paler ven-
tral area. Caudal spot distinct and equal to orbit. Dor-
sum of caudal peduncle with two dark spots with a white
spot of equal size bordering these anteriorly and
posteriorly. Caudal fin dusky, becoming darker distally.
Evidence of pale caudal fin spots present on some
specimens. Anal, pelvic, and pectoral fins pale. Spinous-
dorsal-fin membrane pale, becoming dusky distally. Tips
of spines, extended filaments, and first three spines dark.
Soft dorsal fin dusky with three to four pale elongate
blotches on basal portion. Cheek and branchiostegal
membranes white. Premaxillary and midline of lower jaw
dusky. No apparent snout or cheek markings. A dark

blotch present on lateral and medial opercular surfaces.
Posterior branchial surface devoid of pigment. Upper
pharyngeal area lacking pigment.

The following live color description is taken from
Rosenblatt and Johnson (1974): "... a dull orange line
running longitudinally along the membrane of the
spinous dorsal, slightly above its base; on the soft dorsal
5 dull orange diagonal bars (sloping backwards), and a
very narrow line of brighter orange running just at the
tips of the rays; 2 yellow stripes on the soft anal joining
between the 2nd and 3rd rays; 3 dark orange vertical
wavy lines on the caudal fin; the upper and lower caudal
margin tipped posteriorly with reddish orange; pectorals
transparent; ventrals pale to dusky."

Size. — The species attains a maximum size of 212 mm
SL.

Habits. — Depth records from collection data indicate
that D. labarum has been captured at depths ranging
from 25 to 160 m. However, the species seems to be taken
more often at depths of 40-120 m. Nothing is known of
the life history aspects of D. labarum.

Distribution. — Diplectrum labarum occurs in the Gulf of
California and northward along outer Baja California to
a limit at lat. 27°07.3'N, long. 114°08.3'W and south-
ward along the Central American coast to the Perlas Is-
lands in the Gulf of Panama (Fig. 14). The type-locality
is Bahia de Banderas, Jalisco, Mexico.

Variation. — Rosenblatt and Johnson (1974) noted a dis-
cernible variation in the number of anal rays. They ob-
served that populations of D. labarum along the outer
Baja California (from Almejas Bay northward) had a sig-
nificantly greater number of individuals with eight anal
rays (mean 7.12) than those populations southward of
Almejas Bay along the Baja (7.04) or through the
remainder of its range (7.02). In this study 18% of the
specimens examined (94) from the outer Baja California
coast had eight anal rays. Only 6^0 of all other D.
labarum examined from other localities (50) had eight
anal rays. Rosenblatt and Johnson (1974) were unable to
substantiate their hypothesis that this variation was in-
duced by environmental variables.

Other recognizable differences are seen in the mean
number of scale rows in lateral line for D. labarum (61.27
for Gulf of California and 55.87 for Panama and Costa
Rica specimens). Slight but recognizable differences are
observed in the mean number of pectoral rays in the pop-
ulation from outer Baja (17.00), Gulf of California
(16.24), and Panama and Costa Rica (16.97).

Specimens examined. — In addition to those listed by
Rosenblatt and Johnson (1974); a total of 630 specimens
from 34 to 212 mm SL. Outer Baja California: lacm

33329-1, 1 (82); LACM 32094-1, 1 (135); CAS SU 47, 4 (70-88). GULF OF
CALIFORNIA: LACM 8841-4, 1 (107); LACM 8834-4, 9 (84-139); LACM
8839-5, 15 (88-188); LACM 8834-3, 3 (106-173); LACM 8824-6, 9 (79-172);
UA 67-59-6, 2 (66-76); UA 70-29-7, 1 (126): UA 67-58-3, 3 (65-
88). MEXICO: CAS 4853, 1 (57).

38

Figure 14.— Distribution map for Diplectrum
eumelum (open circles); and D. labarum (closed
circles). [Localities are based on the materials ex-
amined in this study.]

Diplectrum eumelum
Rosenblatt and Johnson 1974

Figures 6, 8, 9, 14, 15

Diplectrum macropoma Jordan and Eigenmann 1890:
397-398 (key, description); Jordan and Evermann
1896:1203, 1205-1206 (systematics); Gilbert and Starks
1904:97-98 (in part, Panama and Gulf of California);
Walford 1937:123-124 (key, systematics); Meek and
Hildebrand 1925:472, 475 (key, systematics, Panama);
Orces 1959:81 (description, Ecuador).

Diplectrum eumelum Rosenblatt and Johnson 1974:178-
191, fig. IB (original description, coastal eastern
Pacific).

The description given by Jordan and Eigenmann
(1890:397-398), although vague in many respects, can
nevertheless be attributed to D. eumelum and not to D.
macropoma. Meristic and proportional data presented
could be assigned to either species, however, the color
description depicting the snout with "... four or five
pale blotches . . . ." (Jordan and Eigenmann 1890:397)
clearly indicates D. eumelum. Later publications, it ap-
pears, merely followed the error of Jordan and Eigen-
mann (1890).

Gilbert and Starks (1904:97-98) recorded this species
from Magdalena Bay, outer Baja California, from Al-
batross Stn. 3039. These specimens are now cataloged as
D. macropoma, CAS SU 47, but examination of this lot
revealed that D. macropoma, D. labarum, D. euryplec-
trum, and D. eumelum were taken at that station. Wal-

ford (1937:123-124) and Orces (1959:81) presented data
for size and color of D. macropoma which can only be at-
tributed to D. eumelum.

Diagnosis. — Diplectrum eumelum is immediately recog-
nized by the distinct series of three to five pale (orange in
live specimens) spots on the snout and lacrimal area. It
has large scales which separate it meristically (Tables 2-
7) from other Diplectrum except D. macropoma. Diplec-
trum eumelum differs from D. macropoma on the basis
of its color pattern, and size (D. macropoma is usually no
greater than 140 mm SL). Caudal-peduncle-depth-to-
head ratio is 1:3.2 for D. eumelum and 1:2.7 for D. mac-
ropoma. The posteriormost dorsal and anal rays are as
long as each fin's respective anterior rays in D. eumelum
and shorter than each fin's respective rays for D. mac-
ropoma. The snout is blunt for D. eumelum and more
pointed for D. macropoma. The number of pseudobranch
filaments is tentatively a character which may separate
these species (Fig. 15) but consideration must be given to
the standard length of specimens.

Juvenile D. eumelum might be confused with juveniles
of D. labarum but color patterns and meristics (es-
pecially cheek scale rows B) will separate these forms.

Description.— D, X, 12 (13); A, III, 7 (8); P, , 17.02 (16-
18); pored lateral-line scales, 48.63 (45-52); scale rows in
lateral line, 57.42 (48-64); scales above lateral line, 6.39
(5-9); scales below lateral line, 15.54 (13-17); predorsal
scale rows, 11.46 (10-15); cheek scale rows A, 5.98 (5-7);
cheek scale rows B, 8.41 (7-10); gill rakers, 19.55 (18.22).

39

40 -

30

10 -

• •

•

• • • • m •

• «• o •• •••

• • a* • • •»•»• • •

m •»• •

-

• •> •• • • • •

mt •••com m

• • • » m

• • O CDCO 0 O

909CMO OCCD O

• •• ••CXDCDCOD CO O O

• •• c»o ocdxsxd o •

o • *o crocacccD

O • CKDCD a> o

• B»ca>ooco oo

-

OO CD O

O • O CO

CD O O

o oooo o

OOO ODO

OOCDOO

oo

o oo oo

cm

o

1 1 1 1

100 150

Standard Length (mm)

200

Figure 15.— Scatter diagram of number of
pseudobranch Filaments versus standard length for
Diplectrum macropoma (open circles) and D.
eumelum (closed circles). [Localities are based on
the materials examined in this study.)

Standard length, 151.63 (47.92, 81.0-233, 41); total
length, 1,279 (15.74, 1,241-1,310, 39); fork length, 1,187
(14.00, 1.156-1,212, 40); body depth, 286 (18.54, 232-321,
35); head length, 387 (11.96, 369-412, 41); postorbital
length, 221 (12.21, 200-266, 41); snout length, 99 (7.29,
87-118, 39); lacrimal width, 48 (4.05, 40-56, 40); maxil-
lary width, 33 (1.85, 28-36, 41); orbit length, 86 (7.73, 72-
102, 40); orbit width, 72 (7.93, 56-93, 40); interorbital
width, 70 (3.80, 64-79, 36); upper jaw length, 171 (5.52,
160-180, 41); lower jaw length, 184 (6.15, 166-196, 40);
cheek height, 129 (7.87, 110-143, 41); cheek length, 121
(5.31, 110-134, 41); caudal peduncle depth, 122 (5.10,
110-133, 40); pectoral fin length, 267 (7.89, 244-285, 40);
pelvic fin length, 201 (10.09, 175-223, 41); predorsal
length, 369 (8.44, 353-382, 36); pelvic origin to lower jaw,
339 (9.47, 316-361, 33); pelvic origin to anus, 314 (13.11,
286-343, 36); postanal length, 402 (10.38, 380-421, 36).
Dorsal spine height: first, 50 (5.69, 39-66, 36); second, 74
(5.80, 61-86, 36); third, 121 (8.45, 107-142, 36); fourth,
132 (20.37, 118-147, 38); fifth, 133 (7.54, 119-146, 36).
Anal spine height: first, 29 (4.88, 21-36, 41); second, 60
(9.38, 48-83, 41); third, 83 (9.21, 69-107, 41).

Diplectrum eumelum has broad spur at preopercular
angle bearing 8-14 elongate spines. Vertical and horizon-
tal preopercular arms serrate. Anterior nostril with elon-
gate flap. Single row of scales along posterior edge of
second anal spine. Caudal fin without elongate filament
but upper lobe noticeably longer than lower. Chest scales
not reduced or embedded. Gill rakers short, the first at
the angle not reaching past base of the third below.
Pseudobranch filaments numerous, 28-35 in adults.

Color. — Color notes are from two adult specimens
taken from the Gulf of Nicoya, Costa Rica, 28 June 1973.
The specimens had been on ice for 24 h before examina-
tion. Dorsum gray-brown, paler ventrally. Ventral sur-
face white with copper tinge more laterally. Lateral body
surface with gold flecks. Some evidence of five to six in-
distinct, thin, dusky ventral bars becoming paler ven-
trally. Somewhat intensified cluster of black pigment in
series of large midlateral blotches. Spinous dorsal fin
pale blue along base, body of fin pale orange. Distally
each interspinous membrane bears a scalloped-shaped
pale orange wedge. Distal portion of spines tipped with
black and orange. Soft dorsal with blue field bearing
series of wavy, orange diagonal lines. Posteriorly, the
wavy bars interrupted to present spotted pattern, pale
orange on pale blue. Anal white at base becoming blotch-
ed with pale orange-yellow distally. Snout with three to
five orange spots in row. These becoming more irregular
posteriorly, the last in series as elongate bar traversing
cheek in a wavy pattern terminating at lower distal por-
tion of preopercular spur. Irregular arrangement of
orange spots also present on preopercles and opercles.
Distal premaxillary edge with orange pigment line. Dis-
tal maxillary with small orange blotch.

Color notes from preserved specimens: preserved
specimens show similar pattern but pale blue appearing
as gray and orange as pale or clear. Caudal peduncle spot
distinct, dark, and as large as eye but oriented ver-
tically. Caudal fin with numerous pale spots in vertical
row on dusky field. Distal caudal ray tips black. Medial
opercular surface black like the posterior branchial cavi-

40

ty. Chest and branchiostegal region pale. Anterior nasal
flap pale. Each lateral body scale with dark pigment at
base giving lateral surface appearance of being marked
by X's. Upper pharyngeal area with dark pigment. Pel-
vic and pectoral fins dusky, pigment running along ray
edge.

Size. — The species is large, reaching at least 234 mm
SL as observed in the present study.

Habits- -The two adults (209-228 mm SL) from the Gulf
of Nicoya on 28 July 1973 had enlarged gonads but the
ovaries did not contain ripe eggs. Stomachs were empty.
Available collection depth data show that the species is
taken from 15 to 100 m but most often from depths of 50
to 90 m.

Distribution. — Diplectrum eumelum is infrequently
taken within its range which extends northward off the
outer Baja California at Bahia Magdalena (Fig. 14), into
the Gulf of California, and southward along the eastern
Pacific coast to Ecuador (lat. 2°29'S, long. 80°59'W). It is
also found at the Galapagos Islands.

Variation. — No obvious variation in characters was dis-
cernible for D. eumelum throughout its distribution.

Specimens examined. — In addition to those listed by
Rosenblatt and Johnson (1974), a total of 130 specimens
from 40 to 234 mm SL. Outer Baja California: UCLAW55-

78. 1 (87); CAS SU 47, 6 (47-121); LACM W52-195, 1 (216). GULF OF
CALIFORNIA: UCLA W62-61, 8 (72-107). SAN SALVADOR: CAS 4761, 1
(54). COSTA RICA: LACM 30719-3, 2 (160-188); UCR 323-15, 1
(165). PANAMA: USNM (lat. 6°44'N, long. 77°33'W, 8 Dec. 1967), 2
(146-149); USNM (Gulf of Chiriqui. 29 Mar. 1963), 4 (136-188); CAS SU
1409, 2 (98-145); UMML 25094, 2 (96-156); UMML 26255, 2 (150-163);
USNM 41481, 5 (40-93); UMML 26150, 1 (142); UMML 26252, 1
(130). COLOMBIA: USNM 211393, 12 (75-123); USNM 211391, 1 (150);
USNM 211381, 1 (205); USNM 213813, 7 (173-213); USNM 213588, 19
(100-170). ECUADOR: USNM 213808, 10 (191-234); USNM 211396, 1

(186). Galapagos Islands: UCLA W53-12, 2 (42-78).

Diplectrum macropoma (Giinther 1864)

Figures G, 8, 7, 13, 15

Centropristis macropoma Giinther 1864:145 (original
description, Panama); Giinther 1869:409, pi. 65, fig. 1
(redescription and figure, Panama).

Diplectrum macropoma. Jordan and Bollman 1890:157-
158, 181 (Panama); Gilbert and Starks 1904:97 (in
part, Panama and Gulf of California); Jordan et al.
1930:318 (check list); Hildebrand 1946:185 (key);
Berdegue A. 1956:53, 264-265 (key, fishery data, Mex-
ico); Chirichigno F. 1974:294-298, fig. 574 (key, Peru);
Rosenblatt and Johnson 1974:178-191, fig. 1C (key,
systematics).

Diplectrum mexicanum Hildebrand 1948:13-15, fig. 6
(original description, Gulf of California); Bortone in
press b (life history, Panama).

The taxonomic confusion of D. macropoma with D.

eumelum has already been discussed under the heading
of the latter. References by Jordan and Bollman (1890),
Jordan et al. (1930), and Hildebrand (1946) may have ac-
tually referred to D. eumelum, but there is no way to
determine this positively from data presented by those
authors.

Giinther (1864:145) described Centropristis mac-
ropoma from three specimens collected by Dow and Sal-
vin from the Pacific coast of Panama. The same descrip-
tion reappeared accompanied by a figure (Giinther
1869:409, pi. 65, fig. 1). Hildebrand (1948:13-15, fig. 6)
described Diplectrum mexicanum based on one
specimen from the Gulf of California. Examination of the
syntype (designated as the lectotype in the absence of a
holotype) of Centropristis macropoma (BMNH 1864-
1.26:426, labeled as Serranus macropoma and hereafter
referred to as a Diplectrum) and a comparison with the
holotype of Diplectrum mexicanum (USNM 46518)
assures me that the two species are synonymous.

The most notable charaters used by Giinther to desig-
nate D. macropoma were: the long spines at the preoper-
cular angle graduating to a serrated preopercular arm;
six rows of cheek scales; and a distinct notch between the
spinous and soft portions of the dorsal fin. Other charac-
ters more clearly noted from his figure and from the syn-
type include: dark pigment (not discernible in a pat-
tern) on the distal half of the dorsal fin; deep body (3.7 in
SL); and an angular snout. The characteristics of D.
mexicanum presented by Hildebrand (1948) include:
deep body (3.2 in SL); angular snout; and six cheek scale
rows. The figure also depicts a preopercular structure
similar to that described and figured by Giinther (1864,
1869).

Hildebrand's specimen was collected in 1889 and the
fact that he did not describe any dorsal fin pigmentation
could be owing to its loss during prolonged preservation.
Current examination also shows a lack of dorsal fin pig-
mentation. A comparison of other meristic and morpho-
metric data indicate no apparent specific differences.

Jordan and Eigenmann (1890:397-398) and Meek and
Hildebrand (1925:475) gave accounts of D. macropoma
which are apparently not of this species but of D.
eumelum. Later, Hildebrand considered D. mexicanum
as distinct from D. macropoma as understood by Meek
and Hildebrand.

Diagnosis. — Diplectrum macropoma has large scales
and, therefore, scale counts tend to be low for this species
(Tables 2-7). These characters should separate it from
other species of Diplectrum except D. eumelum which it
closely resembles. Diplectrum macropoma has pale yel-
low markings on its snout but these are inconspicuous in
live material and nonexistent in preserved specimens.
Diplectrum macropoma has a more pointed snout, a
proportionally lower number of pseudobranch filaments
(Fig. 15), and the posterior dorsal and anal rays are con-
siderably shorter than the anterior rays. These charac-
ters all differ for D. eumelum. Other differences are
noted under the diagnosis for D. eumelum.

41

Confusion with D. labarum might occur but D. mac-
ropoma has none of its anterior spines elongated and the
mean caudal peduncle depth is considerably greater than
that of D. labarum (138.51 as opposed to 117.11 for D.
labarum). Diplectrum macropoma has the greatest body
depth, caudal peduncle depth, predorsal length, and
postanal length of any Diplectrum.

Description.— D, X. 12 (11-13); A, III, 7; P, , 17.12 (16-
18); pored lateral-line scales, 48.09 (44-51); scale rows in
lateral line. 53.86 (48-63); scales above lateral line, 6.32
(5-7); scales below lateral line, 14.60 (11-18); predorsal
scale rows, 13.96 (10-18); cheek scale rows A, 6.27 (5-8);
cheek scale rows B, 8.53 (7-10); gill rakers, 19.93 (17-24).

Standard length, 107.39 (13.74, 90.6-140, 41). Total
length, 1,308 (28.13, 1,260-1,350, 26); fork length, 1,190
(9.30. 1,175-1,207, 25); body depth, 306 (14.64, 273-338,
39); head length, 367 (8.30, 353-391, 41); postorbital
length. 202 (6.12, 194-216, 41); snout length, 96 (6.39, 82-
106, 41); lacrimal width, 41 (2.28, 36-47, 36); maxillary
width, 35 (3.05, 30-43, 36); orbit length, 87 (4.43, 79-97,
41); orbit width, 73 (4.94, 64-85, 41); interorbital width,
67 (4.03, 59-77, 36); upper jaw length, 163 (6.10, 148-177,
41); lower jaw length, 182 (5.85, 165-191, 41); cheek
height, 119 (4.56, 112-135, 41); cheek length, 118 (4.25,
107-140. 41); caudal peduncle depth, 139 (4.31, 128-147,
41); pectoral fin length, 265 (8.93, 246-281, 41); pelvic
fin length, 196 (9.06, 175-210, 36); predorsal length, 382
(9.90, 366-410, 33); pelvic origin to lower jaw, 348 (11.77,
333-373, 27); pelvic origin to anus, 296 (13.04, 269-318,
37); postanal length, 411 (15.51, 387-456, 40). Dorsal
spine height: first, 46 (5.33, 35-63, 40); second, 67 (5.62,
54-79, 39); third, 130 (9.87, 109-154, 40); fourth, 142
(10.47, 122-158, 21); fifth, 144 (10.31, 126-155, 20). Anal
spine height: first, 32 (3.78, 23-38, 41); second, 59 (5.70,
47-69. 41); third, 90 (6.29, 73-101, 40). *

Preopercular spur broad, with 8-13 elongate spines.
Vertical and horizontal arms serrate. Distinct flap pres-
ent on anterior nostril. Gill rakers moderate in length,
the first at angle not reaching past the fourth below.
Scales large but notably smaller in predorsal area. Scales
present along third anal spine and first anal ray. Caudal
lobes unequal, upper being longer but tip not produced
as a filament. Caudal fin only slightly forked. Chest
scales not embedded. Posteriormost soft dorsal and anal
rays short, nearly 1.5 into longest respective soft dorsal
and anal rays. Second anal spine slightly more stout than
third. Pseudobranch filaments few, 20-30 in adults (Fig.
15).

Color. — Color notes were taken from numerous D.
macropoma from the Bay of Panama, January-February
1973. Dorsum pale copper to pale brown. Venter white,
almost silvery. Sides in smaller specimens (60 mm SL)
with two dark brown stripes, uppermost running through
eye onto snout. Larger specimens with two lateral bands
interrupted by pale pigment and appearing indistinct in
outline. Dark snout bar yellow in larger specimens. Pec-
toral fin pale reddish-orange, pelvic fin pale orange. Anal
fin white at base and distally; a wide pale yellow lateral

band through the center of fin. Caudal fin bright reddish
orange on upper lobe, pale gray or white on lower. Body
of caudal fin with five to six vertical rows of small orange
spots on pale blue field. Caudal spot distinct; as large as
eye, oval shaped, oriented laterally (an extension of
lateral stripe) in small specimens; vertical in orientation
in larger specimens. Spinous dorsal fin white, becoming
pale gray distally. Spine tips orange. Soft dorsal rays
tipped with orange. Body of fin with two rows of distinct
round orange spots on pale blue field. Below this, fin
clear to base. Chest and branchiostegal membranes
white. Upper medial opercular surface black, area below
yellow. Posterior branchial area black with yellow lining.
Upper pharyngeal region yellow with sparse gray-blue
flecks. Dorsal surface of eye blue-black, yellow laterally,
and white below. Smaller specimens with dark spot on
caudal peduncle dorsum disappearing in larger speci-
mens. Maxillary gray-brown, dentary white. Lateral
opercular surface metallic blue-green. Three to four
pale, indistinct yellow spots on snout, the uppermost
elongate. Snout spots fade quickly, barely discernible.

Color in preserved specimens shows a similar pattern
as above but all colors have faded. The venter appears
silvery, dorsum tan with evidence of brown lateral indis-
tinct bars. Caudal spot larger than eye and vertical in
orientation. Soft dorsal fin with appearance of wide dark
band above and pale or clear bar below.

Size. — Diplectrum macropoma is a small species not
exceeding 144 mm SL (Rosenblatt and Johnson 1974).

Habits. — In the Bay of Panama (10-15 km east of Naos
Island) I collected the species from about 20 to 30 m over
an open mud and sand bottom along with D. pacificum
(Bortone in press b). Diplectrum macropoma was abun-
dant at this locality with about 50 being obtained in each
20-min tow with a 10-m otter trawl at night. The species
has been recorded to occur at depths ranging from 9 to 80
m but is more frequently taken at depths of 20-40 m.

Distribution. — Diplectrum macropoma is found in the
Gulf of California; northward on the outer Baja Califor-
nia to at least lat. 24°27'N, long. 111°59'W; and south-
ward along the eastern Pacific coast to the Ecuador-
Peruvian border at lat. 3°39'S, long. 80°41'W (Fig. 13).
The species was also taken at the Galapagos Islands. The
type-locality is the Pacific coast of Panama (Giinther
1864).

Variation. — Examination of meristic data indicates
some variation in D. macropoma with regard to the mean
number of predorsal scale rows (Table 5) . Gulf of Califor-
nia (13.81), Panama (14.96), and Colombia (12.92) pop-
ulations all show different means for this character.

Specimens examined. — Outer Baja California: CASSU47,

2 (113-113); SIO 64-807, 1 (29). GULF OF CALIFORNIA: USNM 46518. 1
(95); USNM 183977, 2 (128-140); SOSC (Choya Bay, Mexico, 14 Feb.
1969), 1 (141); SIO 68-72, 24 (54-114); CAS W 59-14, 26 (36-102); INP
6601, 7 (97-129); FMNH 62743, 8 (38-135); UA 67-74-3, 3 (45-50); UA 67-
72-2, 1 (120); SIO 62-75, 1 (128); USLA W52-45, 12 (110-138); SIO 70-141,

42

4; SIO W62-60, 8 (72-139). MEXICO: SIO 62-37, 3 (95-108); SIO 62-46,
1; SIO 62-36, 2 (97-100). COSTA RICA: SIO W63-148, 2. PANAMA:
BMNH 1864-1.26:426, lectotype, 1 (91); SOSC (lat. 8°08'N. long.
80°20'W, 23 July 1967), 22 (86-118); UMML 25094, 21 (25-85); UMML
25075, 5 (33-74); UMML 26094, 2 (111-118); UMML 26253, 4 (30-115);
UMML 25041. 1 (106); UMML 25067, 8 (63-120); UMML 2646, 13 (52-
109); CAS SU 1641, 1 (126); UCR 223-8, 1 (109); UMML 26247, 1
(122). COLOMBIA: USNM 211369, 2 (96-97); USNM 216385, 2 (83-90);
USNM 211421, 1 (104); USNM 211416, 2 (117-122); USNM 211410, 5 (90-
129); USNM 211394, 13 (81-136); USNM 211398, 18 (97-121); USNM
213590, 8 (91-122); USNM 211402, 15 (91-119); USNM 213591, 152 (62-
128); USNM 211397, 12 (100-131). GALAPAGOS ISLANDS: CAS SU
1569, 1 (116).

ZOOGEOGRAPHY

The genus Diplectrum is restricted to the western
hemisphere (Rosenblatt 1967) and, in general, is found in
coastal areas of tropical-subtropical regions. Ekman
(1953) has termed groups with this distribution pattern
as "endemic amphi-American thermophiles." The
generic distribution closely parallels other groups such as
the decapod crustacean genus Mithrax, the pen-
natularian genus Renilla (Ekman 1953) and numerous
fish genera such as Centropomus (Fraser 1968), Calamus
(Randall and Caldwell 1966), Haemulon (Courtenay
1961), and Mycteroperca (Smith 1959). Diplectrum has
affinities for continental coastal areas but D. bivittatum
is commonly present in the Antillean Archipelago.

Six of the nine Pacific species have a fairly continuous
coastal distribution which encompasses nearly the en-
tire Panamanian province (i.e., the Eastern Pacific
Zoogeographic Region of Briggs (1974) ). Of these six, only
D. euryplectrum and D. maximum are not found in the
Gulf of California. Diplectrum maximum is unique (Fig.
11) in that its northern population is separated by a long
geographical range from its southern population. As
stated previously, only juveniles were taken at the
northern locality. Although the larval life history of some
serranids may be relatively long (Smith 1959), the vast
distance involved does not allow for transport of larvae or
juveniles from the southern population (Rosenblatt and
Johnson 1974). Most assuredly this deepwater species
will be found as adults much further north of its southern
population.

Diplectrum sciurus is almost restricted to the Gulf of
California. Although its movement out of the Gulf may
be hindered by geographical and hydrological barriers
(Walker 1960), its extrme morphological adaptations
related to feeding (i.e., long and numerous gill rakers)
give credence to the hypothesis of a restricted dis-
tribution owing to competitive interaction.

Diplectrum conceptione is outside the Panamanian
province in the Peru-Chilean province of Briggs (1974). It
is the only nonthermophilic species in the genus. This
species' southernmost limit at Conception, Chile (the
type-locality) could be explained by possible mis-
labeling by early collectors. Its northern limit is governed
by rapid temperature changes caused by a westward turn
of the Peru Current near Punta Aguja, Paita, Peru. This
temperature block is formidable during "El Nino" years
(Rosenblatt and Walker 1963).

Four species of Diplectrum have been recorded from
the Galapagos Islands (i.e., D. euryplectrum, D. ros-
trum, D. macropoma, and D. eumelum). The fauna of
the Galapagos Archipelago represents a unique
zoogeographical situation (Rosenblatt and Walker 1963).
Although comparative material is scarce, there seem to
be no significant taxonomic differences between
Galapagos and mainland conspecific representatives.
This suggests a nonisolated gene pool for Galapagos
Diplectrum. Larvae may periodically invade the islands
from the Panamanian province during "El Nino" years
when current patterns seem to favor this transport
mechanism (Rosenblatt and Walker 1963). Briggs (1974)
has also indicated the currents which flow toward the
Galapagos Islands from the Panama area during Feb-
ruary through April may be important in explaining the
similarities in marine inshore fauna between the
Galapagos and Panamanian province. It may be sig-
nificant to note the absence of D. conceptione, a Peru-
Chilean province species. Currents would seem to favor
its movement to the Galapagos (Rosenblatt and Walker
1963), yet its absence may be due to competitive ex-
clusion or simply inadequate sampling.

Hubbs (1960) has stated that the northern boundary
for the Panamanian province is not abrupt. Northern
limits for Pacific Diplectrum support this statement.
Magdalena Bay (lat. 24.5°N) is the northern limit for
five of eight Pacific species recorded from the outer Baja
California. Magdalena Bay is considered a northern
boundary for the eastern Pacific tropical fish fauna
(Rosenblatt 1967). Hubbs (1960) indicated that rapid
temperature changes in this area are due to insolation
and upwelling. This is a plausible reason for Diplectrum
northern distribution limits in this area. Two species go
beyond the above faunal boundary and one of these (D.
labarum) extends to Bahia Asuncion (lat. 27°N). It
would be reasonable to expect Diplectrum to extend
further north of the Panama-San Diegan province boun-
dary, especially when local weather conditions favor
northern movement of warm-water species.

In the western Atlantic, the northern limit for the
Carolina Province water is Cape Hatteras, N.C. (Briggs
1974). From collection data it appears Cape Lookout (110
km southwest of Cape Hatteras) is the northernmost area
of abundance for D. formosum . The one locality record
from Chincoteaque, Va. (lat. 38°N) apparently occurred
at a time during favorable weather conditions. Diplec-
trum bivittatum (from the intermediate population) is
probably limited more by favorable reef habitat than
temperature along the southeast U.S. coast. Presumably
the species occurs at deep patch reefs at higher latitudes
than it is currently known.

Southern limits for Atlantic Diplectrum have been
reported in the literature to be Montevideo, Uruguay.
This southern distribution limit cannot be confirmed as
yet. Montevideo (i.e., Rio del Plata) is certainly con-
sidered a zoogeographic boundary (Ekman 1953) and
Briggs (1974) considered this area as the southernmost
boundary of the Eastern South American Region.
However, the present study indicates a slightly more

43

northern boundary for S. Atlantic Diplectrum near Sao
Paulo and Rio de Janeiro. This area is also considered a
boundary by Ekman (1953). and Briggs (1974) attributed
the presence of this boundary to a cooling of the Brazil
current.

Diplectrum formosum, with its two subspecies, pre-
sents an interesting distribution pattern. The northern
form {D. f. formosum) is almost restricted to continental
North America. Except for a single record from the Vir-
gin Islands, the southern form is essentially restricted to
continental South America. The vast expanse of Carib-
bean Sea with its Antillean Island chain (i.e., West In-
dian Province of Briggs (1974) ), appears to have been a
formidable barrier to the species distribution which led
to considerable genetic isolation. Conceivably the species
may have lacked the competitive ability of other reef-
dwelling serranids and was therefore not able to occupy
or use the Antilles coast for its habitat.

Diplectrum bivittatum presents us with two
geographical populations herein considered subspecies.
The Gulf of Mexico form (D. b. arcuarium) seems
restricted to areas of the Gulf with characteristically
"soft" bottom (i.e., mud or sand; Lynch 1954:fig. 19).
The absence of D. b. arcuarium from the west penin-
sular Florida coast and the coast of the Yucatan Penin-
sula, both characterized by limestone and coral sub-
strates, attests to this premise. Diplectrum bivittatum
with its distribution in the Antilles is apparently capable
of surviving at reef as well as coastal localities. The inter-
mediate form of the two recognized subspecies must be
considered by itself zoogeographically. Because the ap-
parent ranges for the two subspecies do not actually over-
lap, an explanation must be offered for the existence of
the S. Florida population. Gulf currents may favor a lar-
val drift of the northern Gulf of Mexico form to the south
Florida area, especially during summer months (Leip-
per 1954; Nowlin 1971). Larval transport across the
mouth of the Gulf from Yucatan is not probable in view
of these currents (Leipper 1954; Nowlin 1971). General
circulation in the Caribbean and Antilles areas favors a
northwest drift of larvae from these areas to south
Florida. Another possible explanation for the existence of
an intermediate form of the two subspecies is that the in-
termediate form represents a third subspecies, almost
completely isolated from the Gulf and excluded by an
unknown factor from the southern form. The first view is
the most favored. Further studies on these forms will
help resolve this question.

Both D. formosum and D. radiale terminate their dis-
tribution along the northern coast of South America at or
near the Gulf of Venezuela. The significance of this dis-
tribution pattern is important when considering the
amphi-American distribution of Diplectrum and par-
ticularly the very close "geminate" relation D. radiale
has with D. pacificum.

It is reasonable to postulate that in view of the ob-
vious affinity of the two species, D. radiale may at one
time have had a distribution which included the Atlan-
tic side of Panama and Colombia. There is general agree-
ment that the latest transoceanic passage available to

marine species occurred during the late Pliocene (Simp-
son 1950). Considering the present distribution of the
geminate pair, the Colombian "gap," open from middle
Eocene to late Eocene or late Oligocene (Smith 1971),
may have served as the last area and time for passage.

If this is true, then relative selective differences
between D. radiale and D. pacificum have been small. A
test for this hypothesis, assuming the species are indeed
distinct, would be to perform comparative in situ life his-
tory studies of the two species. Rosenblatt (1967) has
suggested that morphological diversity and genetic
changes may not be clearly related.

Diplectrum thus portrays an almost classic case of a
distribution limit, approaching lat. 25°N and 5°S (ex-
cept for D. conceptione) along the coastal east Pacific,
due to cold-water current "squeeze" (Rosenblatt 1967).
The Atlantic warm-water gyres have allowed the genus
to assume the classic warm-water distribution which is
essentially from lat, 35°N to 35°S.

ACKNOWLEDGMENTS

Many persons aided me in the course of this study. I
thank the following: Allwyn C. Wheeler (BMNH), Wil-
liam N. Eschmeyer (CAS), Loren P. Woods (FMNH),
Robert W. Topp (FSBC), Carter R. Gilbert (FSM),
Ralph W. Yerger (FSU), Charles E. Dawson (GCRL),
William D. Anderson, Jr. (GMBL), Aurelio Solorzano P.
(INP), Camm C. Swift (LACM), Myvanwy Dick (CMZ),
M. L. Bauchot (MNHN), Naercio A. Menezes
(MZUSP), Maurice Boesemen (RMNH), Richard H.
Rosenblatt (SIO), Leslie W. Knapp (SOSC), George C.
Miller (TABL), Lloyd T. Findley (UA), Boyd W. Walker
(UCLA), William C. Bussing (UCR), C. Richard Robins
(UMML), Frank J. Schwanz (UNO, William H. Eger
(UPR), Victor G. Springer (USNM), Donald E. Wohl-
schlag (UTMSI), and Arthur D. Welander (UW) for
making available museum specimens from their respec-
tive institutions; Fernando Cervigon, Nucleo de Nueva
Esparta, Porlamar, Venezuela, and Richard Moore,
University of Texas, Marine Sciences Institute, for per-
sonally collecting specimens used in this study; Ben
Cooper, Costa Rica, and David Drennan, Panama, for
collecting specimens in their respective countries; Ira
Rubinoff, Smithsonian Tropical Research Institute,
Panama, for making available facilities at that in-
stitution; James K. Dooley, Richard H. Rosenblatt.
David Johnson, John E. McCosker, C. Lavett Smith,
Robert Lavenberg, C. Richard Robins, and Victor G.
Springer for contributing through helpful discussion, ad-
vice, and correspondence; Charles M. Johnson for aid in
obtaining specimens and histological advice; A. F.
Chestnut, Institute of Marine Sciences, University of
North Carolina, for making available facilities necessary
for conducting my research at that institution; Camm C.
Swift for aid in collecting specimens, loan of material,
and very able advice; the members of my graduate com-
mittee, Elizabeth A. McMahan, H. Eugene Lehman, Jan
J. Kohlmeyer, William J. Woods, University of North

14

Carolina, and Joseph R. Bailey, Duke University, for ser-
ving on that committee and editing this manuscript. Jan
J. Kohlmeyer also translated several references.

Special thanks are due to William E. Fahy, my
graduate advisor, for his editing of the manuscript and
for his direction and guidance; and to my wife Midge for
collecting specimens, typing the manuscript, and main-
taining stability in the laboratory and at home.

This work was supported by a traineeship through the
National Science Foundation, an Advanced Service Fel-
lowship from the University of North Carolina, a grant
(GB34290) from the National Science Foundation, and
my wife's salary.

LITERATURE CITED

ABBOTT. J. F.

1900. The marine fishes of Peru. Proc. Acad. Nat. Sci. Phila.
51(18991:324-364.
ADAMS, A. C, and W. C. KENDALL.

1891. Report upon an investigation of the fishing grounds off the
west coast of Florida. Bull. U.S. Fish Comm. 9(1889):289-312.
ANDERSON, W. W.

1968. Fishes taken during shrimp trawling along the South Atlan-
tic coast of the United States, 1931-35. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 570, 60 p.
ANDERSON, W. W., and J. W. GEHRINGER.

1965. Biological-statistical census of the species entering fisheries
in the Cape Canaveral area. U.S. Fish Wildl. Serv., Spec. Sci.
Rep. Fish. 514, 79 p.
ANONYMOUS.

1971. (No title) Mar y Pesca 1971(681:61-64.
BAILEY, R. M., J. E. FITCH, E. S. HERALD, E. A. LACHNER, C. C.
LINDSEY, C. R. ROBINS, and W. B. SCOTT.

1970. A list of common and scientific names of fishes from the
United States and Canada. 3rd ed. Am. Fish. Soc, Spec. Publ.
6, 150 p.

BAILEY, R. M., E. A. LACHNER, C. C. LINDSEY, C. R. ROBINS,
P. M. ROEDEL, W. B. SCOTT, and L. P. WOODS.

1960. A list of common and scientific names of fishes from the
United States and Canada. 2nd ed. Am. Fish. Soc, Spec. Publ.
2. 102 p.
BAUGHMAN, J. L.

1944. Notes on the Serranidae and Lobotidae of Texas. Copeia

1944:89-90.
1950. Random notes on Texas fishes. Part II. Tex. J. Sci. 2:242-
263.
BEARDEN, C. M., and M. D. McKENZIE.

1971. An investigation of the offshore demersal fish resources of
South Carolina. S.C. Wildl. Resour. Dep., Mar. Res. Div., Tech.
Rep. 2, 19 p.

BEAUMARIAGE, D. S.

1968. Manufacture of fish meal from Florida's fishery waste and
underexploited fishes. Fla. State Board Conserv., Tech. Ser. 54,
56 p.
BEEBE, W., and J. TEE-VAN.

1933. Field book of the shore fishes of Bermuda. G. P. Putnam's
Sons, N.Y., 337 p.
BERE, R.

1936. Parasitic copepods from Gulf of Mexico fish. Am. Midi.
Nat. 17:577-625.
BERDEGUE, A., J.

1956. Peces de importancia comercial en la costa nor-occidental de
Mexico. Com. Fomento Piscicultura Rural, Mexico, 345 p.
BERG, C.

1895. Enumeracion sistematica y sinonimica de los peces de las
costas Argentina y Uruguaya. An. Mus. Nac. Buenos Aires 4
(Sect. 2), 120 p.

BOHLKE, J. E., and C. C. G. CHAPLIN.

1968. Fishes of the Bahamas and adjacent tropical waters. Living-
ston Publ. Co., Wynnewood, Pa., 771 p.
BORODIN, N. A.

1934. Scientific results of the yacht "Alva" Mediterranean cruise,
1933, in command of William K. Vanderbilt. Fishes. Bull. Van-
derbilt Oceanogr. Mus. 1:103-123.
BORTONE, S. A.

1970. Studies on the biology of the sand perch, Diplectrum formo-
sum (Linnaeus), (Perciformes: Serranidae). M.S. Thesis, Florida
State Univ., Tallahassee, 56 p.

1971a. Studies on the biology of the sand perch, Diplectrum formo-
sum (Perciformes: Serranidae). Fla. Dep. Nat. Resour., Tech.
Ser. 65, 27 p.

1971b. Ontogenetic and behavioral color patterns in the sand perch,
Diplectrum formosum (Pisces: Serranidae). (Abstr.) Assoc.
Southeast. Biol. Bull. 18:28.

1972. Age and growth of the sand perch, Diplectrum formosum
(Pisces: Serranidae) from North Carolina. (Abstr.) Assoc.
Southeast. Biol. Bull. 19:55.

1973. A systematic revision of the genus Diplectrum Holbrook, 1855
(Pisces: Serranidae: Serraninae). Ph.D. Thesis, Univ. North
Carolina, Chapel Hill, 203 p.

1974. Diplectrum rostrum, a hermaphroditic new species (Pisces:
Serranidae) from the eastern Pacific coast. Copeia 1974:61-65.

In press a. Gonad morphology of the hermaphroditic fish Diplec-
trum pacificum (Serranidae). Copeia.
In press b. Observations on the life history of Diplectrum pacificum
and D. macropoma from the Bay of Panama. Pac. Sci.
BOULENGER, G. A.

1895. Catalogue of the perciform fishes in the British Museum (2nd
ed). Vol. 1.: Centrarchidae, Percidae, and Serranidae (part).
Bri. Mus. (Nat. Hist.). Lond., 394 p.
BRANDAO, J. M.

1964. Glossario de nomes dos peixes; Portugues, Ingles, sistema-
tico. Bol. Estud. Pesca 4(4 and 51:1-59.

BREDER, C. M., JR.

1936. Scientific results of the second oceanographic expedition of
the "Pawnee," 1926. Heterosomata to Pediculati from Panama to
Lower California. Bull. Bingham Oceanogr. Collect., Yale Univ.
2(3), 56 p.
1948. Field book of marine fishes of the Atlantic coast, from Labra-
dor to Texas. G. P. Putnam's Sons, N.Y., 332 p.
BRIGGS, J. C.

1958. A list of Florida fishes and their distribution. Bull. Fla.

State Mus. 2:223-318.
1974. Marine zoogeography. McGraw-Hill, Inc., N.Y., 475 p.
BULLER, R. J.

1951. A fishery survey of southern coastal waters. U.S. Fish Wildl.
Serv., Spec. Sci. Rep. Fish. 58, 21 p.
BULLIS, H. R., JR., and J. R. THOMPSON.

1965. Collections by the exploratory fishing vessels Oregon, Silver
Bay, Combat, and Pelican made during 1956 to 1960 in the south-
western North Atlantic. U.S. Fish Wildl. Serv., Spec. Sci. Rep.
Fish. 510, 130 p.

CALDWELL, D. K.

1966. Marine and freshwater fishes of Jamaica. Bull. Inst. Ja-
maica, Sci. Ser. 17, 120 p.

CALDWELL, D. K., and M. C. CALDWELL.

1964. Fishes from the southern Caribbean collected by Velero III in
1939. Allan Hancock Atlantic Exped., Rep. 10. 61 p.
CANESTRINI, J.

1860. Zur Systematik der Percoiden. Verh. Zool.-Bot., Ges. Wien
10:291-314.
CASTELNAU. F. de.

1855. Poissons. In Animaux nouveaux ou rares recueillis pendant
l'expedition dans les parties centrales de l'Amerique du Sud, de
Rio de Janeiro a Lima, et de Lima au Para; executee par ordre
du Gouvernement Francais pendant les annees 1843 a 1847, sous
la direction du Comte Francis de Castelnau. (Septieme partie.
Zoologie). P. Bertrand, Paris, Tome second, deuxieme volume,
122 p.

45

CERVIGON" M., F.

1966. Los peces marinos de Venezuela. Fondo de Cultura Cienti-
fica. Caracas. Venezuela, Tome I, 436 p.
CHIRICHIGXO F.. X.

1974. Clave para identificar los peces marinos del Peru. Inst, del
Mar del Peru. Callao. Informe No. 44:294-298.

CHRISTENSEN. R. F.

1965. An ichthyological survey of Jupiter Inlet and Loxahatchee
River, Florida. M.S. Thesis. Florida State Univ., Tallahassee,
318 p.
CLEM. L. W.. L. MOEWUS. and M. M. SIGEL.

1961. Studies with cells from marine fish in tissue culture. Proc.
Soc. Exp. Biol. Med. 108:762-766.
CLIBL'RX. J. W.

L973. A key to the fishes of Mississippi Sound and adjacent waters.
Gulf Coast Res. Lab., Ocean Springs, Miss., 35 p.
COURTEXAV. \V. R.. JR.

1961. Western Atlantic fishes of the genus Haemulon (Pomadasyi-
dae); systematic status and juvenile pigmentation. Bull. Mar.
Sci. Gulf Caribb. 11:66-149.
CUVIER. G.

1829. Le regne animal. Nouvelle ed., Vol. 2, Deturville, Paris,
406 p.
CUVIER. G., and A. VALEXCIENNES.

1828. Histoire naturelle des poissons. F. G. Levrault, Paris, vol. 2,
372 p.
DAHL, G.

1971. Los peces del norte de Colombia. Inst. Desarrollo Recursos
Xaturales Renovables, Inderena, Bogota, Colombia, 391 p.
DAHLBERG, M. D.

1975. Guide to coastal fishes of Georgia and nearby states. Univ.
Georgia Press. Athens, 186 p.

D'AXCONA, U.

1952. Territorial sexualization in the gonads of teleosteans. Anat.
Rec. 114:1.

DeKAY, J. E.

1842. Zoology of New York or the New York fauna. Part 4. Fishes.
Albany, 415 p.
DELFIN. F.T.

1901. Catalogo de los peces de Chile. Revista Chilena de Historia
Natural. Valparaisio, Tomos III + IV, 133 p.

DEVIXCENZI, G. J.

1924. Peces del Uruguay. Anal. Mus. Nac. Montevideo. 5(Ser.
21:222-223.
DIAZ, J. V.

1893. Algo sobre peces de Cuba. Havana, 176 p.
DURAND, J.

1961. Notes sur le Plateau Guyanais. Les elements principeaux
de la faune et leurs relations avec le fond. Cahiers Off. Rech. Sci.
Tech. Outre-Mer, Paris 3, 93 p.
EKMAN, S.

1953. Zoogeography of the sea. [Transl. from Swedish.] 1967 ed.
Sidgwick and Jackson, Ltd., Lond., 417 p.

EVERMANN, B. W., and 0. P. JENKINS.

1892. Report upon a collection of fishes made at Guaymas, Sonora,
Mexico, with descriptions of new species. Proc. U.S. Natl. Mus.
14:121-165.
EYERMANN, B. W„ and W. C. KENDALL.

1900. Check-list of the fishes of Florida. Rep. U.S. Comm. Fish
Fish. 25(1899):35-103.
EVERMAXX, B. W., and M. C. MARSH.

1902. The fishes of Porto Rico. Bull. U.S. Fish Comm. 20(1900),
part 1:49-350.

EVERMAXX, B. W., and L. RADCLIFFE.

1917. The fishes of the west coast of Peru and the Titicaca Basin.
Bull. U.S. Natl. Mus. 95, 166 p.
EWALD. J. J., W. BRANDHORST, F. H. DURANT, V. De ESPINOSA,
and W. DIAZ V.

1971. Cruceros de pesca exploratoria del arrastrero "Carmelina" en
la zona occidental de Venezuela. Proyecto de Investigation y
Desarrollo Pesquero, Caracas, Venezuela, Informe Tecnico 25,
58 p.

FOWLER. H. W.

1903. Description of a new gurnard from Florida, with notes on the
colors of some other Florida fishes. Proc. Acad. Nat. Sci. Phila.
55:328-336.

1907. Notes on Serranidae. Proc. Acad. Nat. Sci. Phila. 59:249-

269.
1916a. Fishes of Trinidad. Grenada, and St. Lucia, British West

Indies. Proc. Acad. Nat. Sci. Phila. 67(1915):520-546.
1916b. Cold-blooded vertebrates from Costa Rica and the Canal

Zone. Proc. Acad. Nat. Sci. Phila. 68:389-414.
1927. Fishes from Florida, Brazil, Bolivia, Argentina, and Chile.

Proc. Acad. Nat. Sci. Phila. 78(19261:249-285.
1929. Fishes from Florida and the West Indies. Proc. Acad. Nat.

Sci. Phila. 80(19281:451-473.

1940. The fishes obtained by the Wilkes Expedition, 1838-1842.
Proc. Am. Philos. Soc. 82:733-800.

1941a. A collection of fishes obtained on the west coast of Florida
by Mr. and Mrs. C. G. Chaplin. Proc. Acad. Nat. Sci. Phila.
92(19401:1-22.
1941b. Notes on Florida fishes with descriptions of seven new spe-
cies. Proc. Acad. Nat. Sci. Phila. 93:81-106.
1942. A list of the fishes known from the coast of Brazil. Arquiv.

Zool. Estado de Sao Paulo 3:115-184.
1944. The fishes. Monogr. Acad. Nat. Sci. Phila. 6:57-529.
1945a. A study of the fishes of the southern piedmont and coastal

plain. Monogr. Acad. Nat. Sci. Phila. 7, 408 p.
1945b. Los peces del Peru: catalogo sistematico de los peces que
habitan en aguas peruanas. Mus. Hist. Nat. "Javier Prado",
Univ. Nac. Mayor de San Marcos. Lima, Peru, 298 p.
1945c. Fishes of Chile. Systematic catalog. Rev. Chilena Hist.
Nat. Santiago, 171 p.
FRANKS, J. S.. J. Y. CHRISTMAS, W. L. SILER, R. COMBS, R.
WALLER, and C. BURNS.

1972. A study of nektonic and benthic faunas of the shallow Gulf of
Mexico off the state of Mississippi as related to some physical,
chemical and geological factors. Gulf Res. Rep. 4:1-148.
FRASER. T. H.

1968. Comparative osteology of the Atlantic snooks (Pisces, Cen-
tropomus). Copeia 1968:433-460.
FREIHOFER, W. C.

1963. Patterns of the ramus lateralis accessorius and their systema-
tic significance to teleostean fishes. Stanford Ichthyol. Bull.
8:79-189.
GABRIELSON, I. N., and F. LA MONTE (editors).

1963. The fisherman's encyclopedia. (2nd rev. ed.) Stackpole
Co., Harrisburg, Pa., 759 p.
GALLAWAY. B. J., J. C. PARKER, and D. MOORE.

1972. Key to the estuarine and marine fishes of Texas. 2nd ed.
TAMU-SG-72-402, Texas A&M Univ., College Station, 177 p.
GALLOWAY, J. C.

1941. Lethal effect of the cold winter of 1939-40 on marine fishes at
Key West, Florida. Copeia 1941:118-119.

GILBERT, C. H.

1892. Descriptions of thirty-four new species of fishes collected in
1888 and 1889, principally among the Santa Barbara Islands and
in the Gulf of California. Proc. U.S. Natl. Mus. 14:539-566.

GILBERT, C. H.. and E. C. STARKS.

1904. The fishes of Panama Bay. Mem. Calif. Acad. Sci. 4, 304 p.
GILL, T. N.

1862. Catalogue of the fishes of the eastern coast of North America,
from Greenland to Georgia. Proc. Acad. Nat. Sci. Phila. 13
(1861):l-63.

1863. Remarks on the relations of the Genera and other groups of
CUBAN FISHES. Proc. Acad. Nat. Sci. Phila. 14(1862):235-242.

1864. Review of Holbrook's Ichthyology of South Carolina. Am. J.
Sci. Arts (Silliman's), 2nd ser., 87:89-94.

1873. Catalogue of the fishes of the east coast of North America.
Smithson. Misc. Collect. 14(283), 50 p.
GINSBURG. I.

1948. Some Atlantic populations related to Diplectrum radiate
(Serranidae), with description of a new subspecies from the Gulf
coast of the United States. Copeia 1948:266-270.

■Hi

GMELIN, J. F.

1788. Pisces. In Caroli a Linne Systema Naturae per regna tria
naturae, secundum classes, ordines, genera, species; cum charac-
teribus, differentiis, synonymis, locis. Tom I. Pars III. Edito de-
cimo tertia, aucta, reformata. Cura J. Fred Gmelin. Lipsiae,
1126-1516 p.

GODCHARLES, M. F.

1970. Exploratory fishing for southern sea bass, Centropristes
striatus melanus, in the northeastern Gulf of Mexico. Fla. Dep.
Nat. Resour., Tech. Ser. 63, 26 p.

1971. A study of the effects of a commercial hydraulic clam dredge
on benthic communities in estuarine areas. Fla. Dep. Nat. Re-
sour., Tech. Ser. 64, 51 p.

GOODE, G. B.

1880. A preliminary catalogue of the fishes of the St. John's River
and the east coast of Florida, with descriptions of a new genus and
three new species. Proc. U.S. Natl. Mus. 2:108-121.
1884. The food fishes of the United States. In The fisheries and
fishery industries of the United States, Sect. 1, part 3, p. 163-
682.
1887. American fishes; a popular treatise upon the game and food
fishes of North America, with especial reference to habits and me-
thods of capture. Estes and Lauriat, Boston, 496 p.
GOODE, G. B.. and T. H. BEAN.

1886. On the American fishes in the Linnaean collection. Proc.
U.S. Natl. Mus. 8:193-208.
GOSLINE, W. A.

1966. The limits of the fish family Serranidae, with notes on other
lower percoids. Proc. Calif. Acad. Sci., Ser. 4, 33:91-111.
GREY, M.

1947. Catalogue of type specimens of fishes in Chicago Natural His-
tory Museum. Fieldiana:Zoology 32:109-205.
GRIMES, C. B., and J. A. MOUNTAIN.

1971. Effects of thermal effluent upon marine fishes near the Crys-
tal River stream electric station. Fla. Dep. Nat. Resour., Prof.
Pap. Ser. 17, 65 p.
GUICHENOT, A.

1848. Fauna Chilena. Peces. In C. Gay, Historia fisica y politica
de Chile. Paris ("En casa del Autor"). Zoologia, Tomo Secundo,
p. 137-372.
GUNTER. G.

1935. Records of fishes rarely caught in shrimp trawls in Louisiana.

Copeia 1935:39-40.
1941. Relative numbers of shallow water fishes of the northern Gulf
of Mexico, with some records of rare fishes from the Texas coast.
Am. Midi. Nat. 26:194-200.
GUNTER, G„ and G. E. HALL.

1965. A biological investigation of the Caloosahatchee Estuary of
Florida. Gulf Res. Rep. 2:1-71.
GUNTHER. A.

1859. Catalogue of the acanthopterygian fishes in the collection of

the British Museum. Vol. I. Br. Mus., Lond., 524 p.
1864. Report of a collection of fishes made by Messrs. Dow, God-
man, and Salvin in Guatemala. Proc. Zool. Soc. Lond. 1864:144-
154.
1869. An account of the fishes of the states of Central America,
based on collections made by Capt. J. M. Dow, F. Godman, Esq.,
and O. Salvin, Esq. Trans. Zool. Soc. Lond. 6:377-494.
HENSHALL, J. A.

1891. Report upon a collection of fishes made in southern Florida

during 1889. Bull. U.S. Fish Comm. 9(1889):371-389.
1895. Notes on fishes collected in Florida in 1892. Bull. U.S. Fish
Comm. 14(1894):209-221.
HILDEBRAND, H. H.

1954. A study of the fauna of the brown shrimp (Penaeus aztecus
Ives) grounds in the western Gulf of Mexico. Publ. Inst. Mar.
Sci., Univ. Tex. 3:233-366.

1955. A study of the fauna of the pink shrimp (Penaeus duorarum
Burkenroad) grounds in the Gulf of Campeche. Publ. Inst. Mar.
Sci., Univ. Tex. 4:169-232.

HILDEBRAND, S. F.

1946. A descriptive catalog of the shore fishes of Peru. Bull. U.S.
Natl. Mus. 189:1-530.

1948. A new genus and five new species of American fishes. Smith-
son. Misc. Collect. 110(90), 15 p.
HOESE, H. D.

1958. A partially annotated checklist of the marine fishes of Texas.
Publ. Inst. Mar. Sci., Univ. Tex. 5:312-352.
HOLBROOK, J. E.

1855. Ichthyology of South Carolina. John Russell, Charleston,

S.C., 182 p.
i860. Ichthyology of South Carolina. 2nd ed. Russell and Jones,
Charleston, S.C., 205 p.
HUBBS, C. L.

1960. The marine vertebrates of the outer coast. Syst. Zool. 9:134-
147.
HUBBS, C. L., and K. F. LAGLER.

1964. Fishes of the Great Lakes region. Univ. Michigan Press,
Ann Arbor, 213 p.
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLA-
TURE.

1964. International code of zoological nomenclature adopted by the
XV International Congress of Zoology. [In Engl, and French.]
Int. Trust Zool. Nomencl., Lond., 176 p.
JORDAN, D. S.

1885. List of fishes at Key West, Florida, with notes and descrip-
tions. Proc. U.S. Natl. Mus. 7:103-150.

1886. A list of the fishes known from the Pacific coast of tropical
America, from the Tropic of Cancer to Panama. Proc. U.S. Natl.
Mus. 8:361-394.

1887. A catalogue of the fishes known to inhabit the waters of North
America, north of the Tropic of Cancer, with notes on the species
discovered in 1883 and 1884. Rep. U.S. Comm. Fish. 13(1885):
789-973. [Preprint U.S. Fish. Comm., Doc. No. 94(1885), 185 p.]

1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci., Ser. 2, 5:377-
514.

1908. The law of geminate species. Am. Nat. 42:73-80.
JORDAN, D. S., and C. H. BOLLMAN.

1890. Description of new species of fishes collected at the Galapagos
Islands and along the coast of the United States of Colombia, 1887-
88. Proc. U.S. Natl. Mus. 12(1889):149-183.
JORDAN, D. S.. and C. H. EIGENMANN.

1890. A review of the genera and species of Serranidae found in the
waters of America and Europe. Bull. U.S. Fish Comm. 8(1888):
329-441.
JORDAN, D. S., and B. W. EVERMANN.

1896. The fishes of North and Middle America. Bull. U.S. Natl.
Mus. 47, Part I, 1240 p.

1923. American food and game fishes. Dover Publications, Inc.,
N.Y. [1969 ed. Doubleday, Page, and Company], 574 p.
JORDAN, D. S„ B. VV. EVERMANN, and H. W. CLARK.

1930. Check list of the fishes and fishlike vertebrates of North and
Middle America north of the northern boundary of Venezuela and
Colombia. Rep. U.S. Comm. Fish, for the year 1928, Part 2, 670 p.
JORDAN. D. S., and C. H. GILBERT.

1881. List of fishes collected by Lieut. Henry E. Nichols, U.S.N., in
the Gulf of California and on the west coast of Lower California,
with description of four new species. Proc. U.S. Natl. Mus. 4:273-
279.

1882. Synopsis of the fishes of North America. Bull. U.S. Natl.
Mus. 16, 1018 p.

1883. Notes on fishes observed about Pensacola, Florida, and Gal-
veston, Texas, with description of new species. Proc. U.S. Natl.
Mus. 5:241-307.

JORDAN, D. S.. and J. C. THOMPSON.

1905. The fish fauna of the Tortugas Archipelago. Bull. U.S. Bur.
Fish. 24(19041:229-256.
JOSEPH, E. B., and R. W. YERGER.

1956. The fishes of Alligator Harbor, Florida, with notes on their
natural history. Fla. State Univ. Stud. 22:111-156.
KATAYAMA, M.

1960. Fauna Japonica. Serranidae (Pisces). Tokyo News Serv.,
Ltd., Tokyo, 189 p.
KOEPKE, H. W.

1963. Lista de los peces marinos conocidos del Peru. Biota (4)
Colegio Solesiana Lima 34:237-258.

47

KONNERTH, A.

1965. Preparation of ligamentary articulated fish skeletons. Cur-
ator 8:325-332.

KUMADA, T. (editor).

1937. Marine fishes of the Pacific coast of Mexico. Nissan Fish.
Inst.. Odawara. Japan. 75 p.
LA MOXTE. F.

1952. Marine game fishes of the world. Doubleday & Co., Inc.,
Garden City. N.Y., 190 p.
LEIPPER. D. F.

1954. Physical oceanography of the Gulf of Mexico. In P. S. Galt-
soff (coordinator). Gulf of Mexico its origin, waters, and marine
life. p. 119-137. U.S. Fish Wildl. Serv., Fish Bull. 55.
LESSON. R. P.

1930. Voyage autour de monde sur la corvette de la mageste. La
Coquille. Poissons, 66-238 p., 37 pis. Zoologie vol. 2, part. I,
471 p.
LINNAEUS. C.

1766. Systema naturae per regna tria naturae, secundum classes,
ordines. genera, species, cum characteribus, differentiis, synony-
mis, locis. 12th ed. Laurentii Salvii, Holmiae(Part 1), 532 p.
1789. Systema naturae. Tome I, pars. Ill, p. 1033-1516.
LONGLEY. W. H.

1932. Observations upon Tortugas fishes. Carnegie Inst. Wash.
Publ. 31:285-286.
LONGLEY. W. H., and S. F. HILDEBRAND.

1941. Systematic catalogue of the fishes of Tortugas, Florida. Car-
negie Inst. Wash. Publ. 535, 331 p.
LOWE. R. H.

1962. The fishes of the British Guiana continental shelf, Atlantic
coast of South America, with notes on their natural history. J.
Linn. Soc. Lond., Zool. 44:669-700.
LYNCH, S. A.

1954. Geology of the Gulf of Mexico. In P. S. Galtsoff (coordina-
tor). Gulf of Mexico its origin, waters, and marine life, p. 67-86.
U.S. Fish Wildl. Ser., Fish. Bull. 55.
MAGO LECCIA, F.

1970. Lista de los peces de Venezuela, in clayendo un estudio pre-
liminar sobre la ichtiogeografia del pais. Minist. Agric. Cria. Of.
Nac. Pesca, Caracas, 283 p.
MANN F.. G.

1954. Vida de los PECES en aguas chilenas. Minist. Agric, Santi-
ago, 339 p.

McAllister, d. e.

1968. Evolution of branchiostegals and classification of teleostome
fishes. Bull. Natl. Mus. Can. 221, 239 p.
McCULLY, H. H.

1961. The comparative anatomy of the scales of the serranid fishes.
Ph.D. Thesis, Stanford Univ., 258 p.
MEEK, S. E., and S. F. HILDEBRAND.

1925. The marine fishes of Panama. Field Museum of Natural
History. Publ. 226, Zool. Ser. 15:472-478.
METZELAAR, J.

1919. Report on the fishes collected by Dr. J. Boeke in the Dutch
West Indies, 1904-1905. Tweede Gedeelte, Bijlage 1, part 1, 314 p.
MIGDALSKI, E. C.

1958. Angler's guide to the salt water game fishes. Ronald Press
Co., N.Y., 506 p.
MILLER, G. L., and S. C. JORGENSON.

1973. Meristic characters of some marine fishes of the western At-
lantic Ocean. Fish. Bull., U.S. 71:301-312.
MIRANDA RIBEIRO, A. de.

1904. Pescas do "Annie." Imprensa Nacional, Rio de Janeiro, 53 p.
1915. Fauna Braziliense. Peixes. V. Physoclisti. Arch. Mus. Nac,

Rio de Janeiro, vol. 17 (Serranidae), 39 p.
1918. Fauna Braziliense. Peixes. V. Physoclisti. Arch. Mus. Nac,
Rio de Janeiro, vol. 21, 227 p.
MOE, M. A., JR.

1966. Tagging fishes in Florida offshore waters. Florida State
Board Conserv., Tech. Ser. 49, 40 p.

MOE, M. A.. JR., P. C. HEEMSTRA, J. E. TYLER, and H. WAHL-
QUIST.

1966. An annotated listing of the fish reference collection at the

Florida Board of Conservation Marine Laboratory. Florida State
Board Conserv., Spec. Sci. Rep. 10, 121 p.
MOE, M. A., JR., and G. T. MARTIN.

1965. Fishes taken in monthly trawl samples offshore of Pinellas
County, Florida, with new additions to the fish fauna of the Tampa
Bay area. Tulane Stud. Zool. 12:129-151.

MONOD, T.

1968. Le complexe urophore des poissons teleosteens. Mem. Inst.
Fondam. D'Afrique Noire 81, 705 p.
MOUNTAIN, J. A.

1972. Further thermal addition studies at Crystal River, Florida
with an annotated checklist of marine fishes collected 1969-1971.
Fla. Dep. Nat. Resour., Prof. Pap. Ser. 20, 103 p.
NAHHAS, F. M., and E. C. POWELL.

1971. Digenetic trematodes of marine fishes from the Floridian
northern Gulf of Mexico. Tulane Stud. Zool. 17:1-9.
NEAVE, S. A.

1939. Nomenclator Zoologicus, Vol. 2. Zool. Soc. Lond., 1025 p.
NICHOLS, J. T.

1929. The fishes of Porto Rico and the Virgin islands: Branchio-
stomidae to Sciaenidae. Scientific Survey of Porto Rico and the
Virgin Islands. New York Acad. Sci. 10(part IT):161-295.
NORMAN, J. R.

1966. A draft synopsis of the orders, families and genera of recent
fishes and fish-like vertebrates. Br. Mus. (Nat. Hist.), Lond.,
649 p.

NOWLIN, W. D., JR.

1971. Water masses and general cumulation of the Gulf of Mexico.
Oceanol. Int. 6(2):28-33.
ORCES, G.

1951. Sobre una coleccion de peces marinos obtenida en el norveste
del Ecuador. Bol. Inf. Cient. Nac. Quito 4(1951):353-369. [Not
seen.]

1959. Peces marinos del Ecuador que se conservan en las colecciones
de Quito. Cienc. Nat. 2(2):72-91.
OSBURN, R. C, and J. T. NICHOLS.

1916. Shore fishes collected by the 'Albatross' expedition in Lower
California with descriptions of new species. Bull. Am. Mus. Nat.
Hist. 35:139-181.
OUTDOOR WRITERS ASSOCIATION OF AMERICA.

1952. Standard checklist of common names for principle American
sport fishes. Monumental Print. Co., Baltimore.

PEARSE, A. S.

1952a. Parasitic crustaceans from Alligator Harbor, Florida. Q. J.

Fla. Acad. Sci. 15:187-243.
1952b. Parasitic Crustacea from the Texas coast. Publ. Inst. Mar.

Sci., Univ. Tex. 2(2):5-42.
POEY, F.

1866. Repertorio fisico-natural de la isla de Cuba. Tome I. Bar-

cina Y. Comp., Habana, 420 p.
1868. Synopsis piscium Cubensium. Catalogo razonado de los peces

de la isla de Cuba, extractado del Repertorio-fisico-natural de

Cuba, tomo secundo, pagina 279 y siguientes. Barcina Y. Comp..

Havana, 484 p.
1871. Genres des poissons de la faune de Cuba appartenant a la

famille Percidae. Ann. Lycium Nat. Hist. New York 10:1-79.
1876. Enumeratio piscium cubensium. Madrid, 224 p.
1955. Ictiologia Cubana. Ministerio de Educacion Vol. 1, No. 1 de

la Comision, Habana, Cuba.
PORTER, C. E.

1909. Breve nota de ictiologia. Revista Universitaria, Lima 3, 14 p.
QUOY, J. R. C. and P. GAIMARD.

1824. Zoologie (Poissons). In Louis de Freycinet, Voyage autour

du monde, execute sur les corvettes de L. M. YUranie et la Physi-

cienne, pendant les annees 1817, 1818, 1819 et 1820. Pillet Aine,

Paris: Text, Chapters VIII and IX, p. 183-401.
RANDALL, J. E.

1968. Caribbean reef fishes. T. F. H. Publications, Jersey City,

N.J., 318 p.
RANDALL, J. E., and D. K. CALDWELL.

1966. A review of the sparid genus Calamus, with descriptions of

four new species. Bull. Los Ang. Cty. Mus. Nat. Hist. Sci. 2,

47 p.

48

REID, G. K„ JR.

1954. An ecological study of the Gulf of Mexico fishes, in the vi-
cinity of Cedar Key, Florida. Bull. Mar. Sci. Gulf Carihb.
4:1-94.
RINGUELET, R. A., and R. H. ARAMBURU.

1960. Peces marinos de la Republica Argentina clave para el recon-
ocimiento de familias y generos, catalogo critico abreviado. Argo:
Publ. Tecnica, Ano II, No. 5:36, 61-62.

ROBINS, C. R.

1958. Check list of the Florida game and commercial marine fishes.
Fla. State Board Conserv., Educ. Ser. 12, 46 p.

ROBINS, C. R.,'and W. A. STARCK, JR.

1961. Materials for a revision of Serranus and related fish genera.
Proc. Acad. Nat. Sci. Phila. 113:259-314.

ROESSLER, M.

1965. An analysis of the variability of fish populations taken by ot-
ter trawl in Biscayne Bay, Florida. Trans. Am. Fish. Soc. 94:311-
318.
ROITHMAYR. C. M.

1965. Industrial bottomfish fishery of the northern Gulf of Mexico,
1959-63. U.S. Fish Wildl. Serv., Spec. Sci. Rep. Fish. 518, 23 p.
ROSENBLATT. R. H.

1967. The zoogeographic relationships of the marine shore fishes of
tropical America. Stud. Trop. Oceanogr. (Miami) 5:579-592.
ROSENBLATT, R. H.. and G. D. JOHNSON.

1974. Two new species of sea basses of the genus Diplectrum, with a
key to the Pacific species. Calif. Fish Game 60:178-191.
ROSENBLATT, R. H.. and B. W. WALKER.

196.3. The marine shore-fishes of the Galapagos Islands. Occas.
Pap. Calif. Acad. Sci. 44:97-106.
SCHMELTZ, J. D. E., JR.

1869. Museum Godeffroy, Catalog IV, nebst einer, Beilage, Ham-
burg, 29 p.
SCHULTZ, L. P.

1949. A further contribution to the ichthyology of Venezuela. Proc.
U.S. Natl. Mus. 99(3235), 211 p.

SEALE, A.

1940. Report on the fishes from Allan Hancock expeditions in the

California Academy of Sciences. Allan Hancock Pac. Exped. 9:1-

46.
SIMPSON, G. G.

1950. History of the fauna of Latin America. Am. Sci. 38:361-389.
SMITH. C. L.

1959. Hermaphroditism in some serranid fishes from Bermuda.
Pap. Mich. Acad. Sci., Arts Lett. 44:111-119.

1965. The patterns of sexuality and the classification of serranid

fishes. Am. Mus. Novitates No. 2207, 20 p.
1971. A revision of the American groupers: Epinephelus and allied
genera. Bull. Am. Mus. Nat. Hist. 146:67-241.
SMITH, C. L., and E. H. ATZ.

1969. The sexual mechanism of the reef bass Pseudogramma ber-
mudensis and its implications in the classification of the Pseudo-
grammidae. (Pisces: Perciformes). Z. Morph. Tiere 65:315-326.
SMITH. C. L., and R. M. BAILEY.

1961. Evolution of the dorsal-fin supports of percoid fishes. Pap.
Mich. Acad. Sci.. Arts Lett. 46:345-363.
SMITH. H. M.

1896. Notes on Biscayne Bay, Florida, with reference to its adapt-

ability as the site of a marine hatching and experiment station.
Rep. U.S. Comm. Fish Fish. 21(1895):169-191.
1907. The fishes of North Carolina. N.C. Geol. Econ. Survey,
Raleigh 2. 453 p.
SPRINGER, V. G., and K. D. WOODBURN.

1960. An ecological study of the fishes of the Tampa Bay area.
Fla. State Board Conserv., Prof. Pap. Ser. 1, 104 p.

STARCK. W. A., II.

1968. A list of fishes of Alligator Reef, Florida, with comments on
the nature of the Florida reef fish fauna. Undersea Biol. l(l):l-40.

STARCK. W. A., II, and W. P. DAVIS.

1966. Night habits of fishes of Alligator Reef, Florida. Ichthyolo-
gica 38:313-356.

STEINDACHNER. F.

1868. Ichthyologische Notizen. VII. Sitzber. Akad. Wiss. Wien,
1868, 57, 1. Abth., 965-1008.

1875. Ichthyologische Bietrage. II. Sitzber. Akad. Wiss. Wien,
1875, 71, 1. Abth., 1-38.

1876. Ichthyologische Beitrage. IV. Sitzber. Akad. Wiss. Wien,
1875. 72, 1. Abth., 551-616.

STORER. P. H.

1846. A synopsis of the fishes of North America. Metcalf and Co.,
Cambridge, Mass., 298 p.
STREETS, T. H.

1877. Contributions to the natural history of the Hawaiian and
Fanning Islands and Lower California, made in connection with
the United States North Pacific surveying expedition, 1873-'75.
Smithson. Misc. Collect. 13(7), 172 p.

SWINGLE, H. A.

1971. Biology of Alabama estuarine areas — cooperative Gulf of
Mexico estuarine inventory. Ala. Mar. Res. Bull. 5, 123 p.
TABB, D. C, and R. B. MANNING.

1961. A checklist of the flora and fauna of northern Florida Bay and
adjacent brackish waters of the Florida mainland collected during
the period July, 1957 through September, 1960. Bull. Mar. Sci.
Gulf Caribb. 11:552-649.

TAYLOR, W. R.

1967. An enzyme method of clearing and staining small vertebrates.
Proc. U.S. Natl. Mus. 122(3596), 17 p.

TROSCHEL.

1858. Arch. Naturgesch. No. 24, Abt. 2, p. 80. [Not seen.]
TYLER, J. E.

1964. Preliminary exploratory fishing on the Florida west coast.
Fla. State Board Conserv., Spec. Sci. Rep. 8, 14 p.
VIEILLOT.

1816. Analyse, 50 p. [Not seen.]
WALFORD, L. A.

1937. Marine game fishes of the Pacific coast from Alaska to the
Equator. Univ. Calif. Press, Berkeley, 205 p.
WALKER, B. W.

1960. The distribution and affinities of the marine fish fauna of the
Gulf of California. Syst. Zool. 9:123-133.
WALLS, J. G.

1975. Fishes of the northern Gulf of Mexico. T. F. H. Publ. Inc.,
Ltd., Hong Kong, 4.32 p.
WOLF, K., and M. C. QUIMBY.

1969. Fish cell and tissue culture. In W. S. Hoar and D. J. Randall
(editors). Fish physiology, 3:253-301. Academic Press, N.Y.

49

&GPO 798-634

388. Proceedings of the first U.S. -Japan meeting on aquaculture at
Tokyo, Japan, October 18-19, 1971. William N. Shaw (editor). (18
papers, 14 authors.) February 1974, iii + 133 p. For sale by the
Superintendent of Documents, U.S. Government Printing Office,
Washington, D.C. 20402.

389. Marine flora and fauna of the northeastern United States.
Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + 50 p.. Ill
figs. For sale by the Superintendent of Documents, U.S. Government
Printing Office," Washington, D.C. 20402.

390. Fishery publications, calendar year 1973: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. September 1974, iv + 14 p., 1 fig.
For sale bv the Superintendent of Documents, U.S. Government Printing
Office, Washington, D.C. 20402.

391. Calanoid copepods of the genera Spinocalanus and Mimocalanus
from the central Arctic Ocean, with a review of the Spinocalanidae. By
David M. Damkaer. June 1975, x + 88 p., 225 figs., 4 tables. For sale

bv the Superintendent of Documents, U.S. Government Printing Office,
Washington, D.C. 20402.

392. Fishery publications, calendar year 1974: Lists and indexes. By
Lee C. Thorson and Mary Ellen Engett. June 1975, iv + 27 p., 1 fig.

393. Cooperative Gulf of Mexico estuarine inventory and study — Texas:
Area description. By Richard A. Diener. September 1975, vi + 129 p.,
55 figs., 26 tables.

394. Marine Flora and Fauna of the Northeastern United States. Tar-
digrada. Bv l.eland W. Pollock. May 1976, iii + 25 p., figs. For sale
by the Superintendent of Documents, U.S. Government Printing Office,
Washington. D.C. 2(1402.

395. Report of a colloquium on larval fish mortality studies and their
relation to fishery research, January 1975. By John R. Hunter. May
1976, iii + 5 p. For sale by the Superintendent of Documents, U.S.
Government Printing Office, Washington, D.C. 20402.

UNITED STATES
DEPARTMENT OF COMMERCE

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION

NATIONAL MARINE FISHERIES SERVICE

SCIENTIFIC PUBLICATIONS STAFF

ROOM 450

1107 N E 45TH ST

SEATTLE. WA 98105

PENN STATE UNIVERSITY LIBRARIES

ADDDOTSDlflfib

OFFICIAL BUSINESS

NOAA SCIENTIFIC AND TECHNICAL PUBLICATIONS

NOAA, the National Oceanic and Atmospheric Administration, was established as part of the Department of
Commerce on October 3, 1970. The mission responsibilities of NOAA are to monitor and predict the state of the
solid Earth, the oceans and their living resources, the atmosphere, and the space environment of the Earth, and to
assess the socioeconomic impact of natural and technological changes in the environment.

The six Major Line Components of NOAA regularly produce various types of scientific and technical infor-
mation in the following kinds of publications:

PROFESSIONAL PAPERS— Important definitive
research results, major techniques, and special in-
vestigations.

TECHNICAL REPORTS— Journal quality with
extensive details, mathematical developments, or
data listings.

TECHNICAL MEMORANDUMS— Reports of
preliminary, partial, or negative research or tech-
nology results, interim instructions, and the like.

CONTRACT AND GRANT REPORTS— Reports

prepared by contractors or grantees under NOAA
sponsorship.

TECHNICAL SERVICE PUBLICATIONS—

These are publications containing data, observations,
instructions, etc. A partial listing: Data serials; Pre-
diction and outlook periodicals: Technical manuals,
training papers, planning reports, and information
serials; and Miscellaneous technical publications.

ATLAS — Analysed data generally presented in the
form of maps showing distribution of rainfall,
chemical and physical conditions of oceans and at-
mosphere, distribution of fishes and marine mam-
mals, ionospheric conditions, etc.

Information on availability ot NOAA publications can be obtained from:

ENVIRONMENTAL SCIENCE INFORMATION CENTER

ENVIRONMENTAL DATA SERVICE

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION

U.S. DEPARTMENT OF COMMERCE

3300 Whitehaven Street, N.W.
Washington, D.C. 20235