Anisochromis straussi, New Species

of Protogynous Hermaphroditic Fish,

and Synonymy of Anisochromidae,

Pseudoplesiopidae, and

Pseudochromidae

VICTOR G. SPRINGER, C. LAVETT SMITH,

and THOMAS H. FRASER

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 252
SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of "diffusing knowledge" was expressed

by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph

Henry outlined a program that included the following statement: "It is proposed to

publish a series of reports, giving an account of the new discoveries in science, and

of the changes made from year to year in all branches of knowledge." This theme

of basic research has been adhered to through the years by thousands of titles issued

in series publications under the Smithsonian imprint, commencing with Smithsonian

Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoo/ogy

Smithsonian Studies in Air and Space

Smithsonian Studies in History and Technology

In these series, the Institution publishes small papers and full-scale monographs

that report the research and collections of its various museums and bureaux or of

professional colleagues in the world cf science and scholarship. The publications are

distributed by mailing lists to libraries, universities, and similar institutions throughout

the world.

Papers or monographs submitted for series publication are received by the

Smithsonian Institution Press, subject to its own review for format and style, only

through departments of the various Smithsonian museums or bureaux, where the

manuscripts are given substantive review. Press requirements for manuscript and art

preparation are outlined on the inside back cover.

S. Dillon Ripley

Secretary

Smithsonian Institution
SMITHSONIA N CONTRIBUTION S T O ZOOLOGY • NUMBER 252

Anisochromis straussi, New Species

of Protogynous Hermaphroditic Fish,

and Synonymy of Anisochromidae,

Pseudoplesiopidae, and Pseudochromidae

Victor G. Springer, C. Lavett Smith,

and Thomas H. Fraser

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Springer, Victor G., C. Lavett Smith, and Thomas H. Fraser. Anisochromis

straussi, New Species of Protogynous Hermaphroditic Fish, and Synonymy of

Anisochromidae, Pseudoplesiopidae, and Pseudochromidae. Smithsonian Contributions to Zoology^ number 252, 15 pages, 2 figures, 4 tables, 1977.—Anisochromis

straussi is described from the isolated St. Brandon's Shoals in the southwestern

Indian Ocean. The species exhibits two types of color patterns, which are correlated to some degree with size and sex. Histological sections of the gonads indicate

that all testes have a nonfunctional cavity that represents the remnant of the

ovarian lumen, and all have remnants of oocytes among the acini. The smaller

specimens are female, the larger, males. Evidence is presented that the Anisochromidae, Pseudoplesiopidae, and Pseudochromidae form a monophyletic group.

Evidence is also presented that within this group the Pseudoplesiopidae and Anisochromidae form a monophyletic group and each of these two taxa is definable

based on autapomorphies. The Pseudochromidae appear to be a possibly polyphyletic primitive residue group that is not definable on the basis of autapomorphies. The lowest phylogenetic level for which there is evidence of monophyly

among the fishes currently assigned to the Anisochromidae, Pseudoplesiopidae,

and Pseudochromidae was chosen as the level for family recognition. Pseudochromidae is the oldest family-group name available for this level.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Springer, Victor Gruschka, 1928-

Anisochromis straussi, new species of protogynous hermaphroditic fish, and synonymy of Anisochromidae, Pseudoplesiopidae, and Pseudochromidae.

(Smithsonian contributions to zoology : no 252)

Bibliography, p.

1. Anisochromis straussi. 2. Pseudochromidae 3. Hermaphroditism. 4. Fishes—Mauritius—

Cargados C'.arajos Shoals. I. Smith, C. Lavett, 1927- , joint author. II. Fraser, Thomas H.,

joint author. III. Title: Anisochromis straussi. new species of protogynous hermaphroditic

fish. . . IV. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 252.

QL1.S54 no. 252 [QL638.P84] 591.'08s [597/58J 77-608069
Contents

Page

Introduction 1

Methods 1

Abbreviations 1

Comparative Material 1

Acknowledgments 2

Anisochromis straussi, new species 2

Reproductive Biology 7

Relationships of the Anisochromidae 7

Interrelationships of the Pseudochromoids 9

Synonymization of the Anisochromidae and Pseudoplesiopidae 11

Diagnosis of the Pseudochromidae 13

Literature Cited 14

Anisochromis straussi, New Species

of Protogynous Hermaphroditic Fish,

and Synonymy of Anisochromidae,

Pseudoplesiopidae, and Pseudochromidae

Victor G. Springer, C. Lavett Smith,

and Thomas H. Fraser

Introduction

This study was initiated by our collecting an

undescribed species of the previously monotypic

genus Anisochromis, currently assigned to the

monotypic family Anisochromidae. Except for listings or discussions based on the original descriptions (J. L. B. Smith, 1954b) of Anisochromis and

the Anisochromidae, there has been no new information presented on these two taxa. During preparation of the description of our new species, we

became impressed by the similarity of Anisochromis

to fishes of the family Pseudochromidae and Pseudoplesiopidae. We were thus prompted to investigate

the systematic relationships of Anisochromis. We

also noted that eggs were often present in the gonads of specimens exhibiting either of the two types

of color patterns shown by our new species. J. L. B.

Smith (1954b) had reported that each of the two

color patterns exhibited by his species (which are

quite similar to those of our species) was indicative

Victor G. Springer, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D.C. 20560. C. Lavett Smith, Department of

Ichthyology, American Museum of Natural History, Central

Park West at 79th Street, New York, New York 1002-f. Thomas

H. Fraser, Environmental Quality Laboratory, 590-D Olean

Boulevard, Port Charlotte, Florida 33592.

of a different sex. This apparent discrepancy caused

us to examine the gonads of the new species in

more detail. The results of our investigations are

reported below.

METHODS.—Vertebrae and dorsal and anal-fin ray

counts were taken from radiographs. The osteological description of Anisochromis is based on four

cleared and stained specimens (see "Comparative

Material").

Gonadal tissue for cross sectioning was taken

from specimens originally fixed in approximately

ten percent formalin, washed in freshwater, and

preserved in 75 percent ethanol. Sections were

stained in Mayer's hematoxylin and eosin-phloxine

solution.

ABBREVIATIONS.—ANSP = Academy of Natural

Sciences of Philadelphia; AMNH = American Museum of Natural History, New York City; BMNH =

British Museum (Natural History), London; CAS =

California Academy of Sciences, San Francisco;

RUSI = Rhodes University, J. L. B. Smith Institute of Ichthyology, Grahamstown, South Africa;

USNM = acronym for former United States National Museum, collections of which are now in

National Museum of Natural History (NMNH),

Smithsonian Institution, Washington, D.C.

COMPARATIVE MATERIAL.—A wide variety of pre-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

served whole specimens representing various families, genera, and species of fishes was examined,

often by radiography, to determine osteological and

other characters. With the exception of specimens

of Percichthys, this material is not listed here. Four

specimens of Percichthys trucha (USNM 52426)

were radiographed to obtain information on dorsal,

anal, and caudal-fin structure, and one specimen

was dissected to obtain information on the branchial and hyoid arches (poor condition of the specimens precluded clearing and staining). It was not

always possible to determine genus or species for

the specimens of Pseudochromidae and Pseudoplesiopidae. Supplementary information on these two

families was derived from the studies of Lubbock

(1975, 1976), Schultz (1967), and J. L. B. Smith

(1945a,b). The most relevant cleared and stained

specimens examined are listed as follows (numbers

in parentheses indicate number of specimens

examined):

ACANTHOCLINIDAE: Acaiithoclinus species, USNM 200546 (1);

AcanthopJestops hiatti, USNM uncataloged (2); Belonepterygion fasciolatum, USNM 211306 (2). ANISOCHROMIDAE:

Anisochromis straussi, USNM 215859 (4). CEPOLIDAE: Cepola

pauclradiata, USNM 213502 (1). GRAMMIDAE: Gramma loreto, USNM 199487 (3); Lipogramma anabantoides, USNM

216405 (2). OPISTOCNATHIDAE: Opistognathus whitehursti,

ANSP 120714 (1); Stalix moenensis, USNM 211963 (1). PERCICHTHVIDAE: Morone americana, USNM 109851 (3). PLESIOPIDAE: Assessor mcneilli, USNM uncataloged (3); Plesiops

coeruleolineatus, USNM 211304 (2); Trachinops taeniatus,

USNM uncataloged (1). PSEUDOCHRO.MIDAE: Pseudochromis

flavivertex, USNM 211305 (2). PSEUDOGRAMMIDAE: Aporops

species, USNM uncataloged (3); Pseudogramma polyacantha,

USNM 209575 (3). PSEUDOPLESIOPIDAE: Chlidichthys auratus,

USNM 211780 (2); C. ntbiceps, USNM 211777 (5); Pseudoplesiops anuae, USNM uncataloged (2). SERRANIDAE: Bostockia

porosa, AMNH uncataloged (2); Epinephelus fasciatus, USNM

uncataloged (3); Serranus cabrilla, USNM uncataloged (2).

Twenty-one specimens of Anisochromis kenyae served as

the basis for comparing that species with our new one: RUS1

824 (14), USNM 216415 (6), and ANSP 134469 (1).

ACKNOWLEDGMENTS.—We wish to thank the following individuals for providing loans of specimens

or information on holotypes: M. Boeseman, Rijksmuseum van Natuurlijke Historic Leiden; A. Kotthaus, Hamburg; H. Nijssen, Zoologisch Museum,

Amsterdam; M. M. Smith and R. Winterbottom,

RUSI; W. F. Smith-Vaniz, ANSP; P. J. P. Whitehead and A. C. Wheeler, BMNH.

A. Spreitzer, NMNH, provided laboratory assistance, and P. Melville, NMNH, prepared the histological sections of gonads for our study.

The specimens of the new species of Anisochromis

were collected during the 1976 expedition to St.

Brandon, which was conceived, produced, and

funded by L. H. Strauss, Washington, D.C.

Drafts of the manuscript were read by, and benefited from the criticism of, G. J. Nelson and D. E.

Rosen (AMNH), and W. L. Fink (Museum of Comparative Zoology, Harvard University).

Anisochromis straussi, new species

FIGURES 1, 2

DIAGNOSIS.—A species of Anisochromis with 25-27

segmented dorsal-fin rays (modally 26), 17-19 segmented anal-fin rays (modally 18), 33-35 total vertebrae (modally 34). Dorsal fin of terminal stage color

pattern (Figure 1) lacking a discrete dark spot or

blotch in interradial membranes at anterior end

of dorsal fin.

DESCRIPTION.—Osteology. Infraorbital bones 3-5

(includes lacrymal and relatively tiny dermosphenotic; lesser numbers apparently resulting from

fusions); where five infraorbitals are present, third

from anteriormost bears wide subocular shelf. One

extrascapular (lateral) on each side (supratemporal

canal passing only through skin medially; canals

from opposite sides just failing to join middorsally).

Orbitosphenoid absent. Basisphenoid present. Vomer toothed. Palatine toothless, well separated from

foreshortened mesopterygoid and ectopterygoid

(ectoterygoid reduced to small blade of bone without anteriorly extending process). Supramaxillaries

absent. Sesamoid articulars (coronomeckelians) absent. Outer row of large teeth in each jaw; patch

of much smaller teeth behind outer row anteriorly

in upper jaw; irregular row of much smaller teeth

behind outer row anteriorly in lower jaw.

' Infrapharyngobranchial 1 absent; infrapharyngobranchials 2,3, and toothplate of 4 present, toothed;

no interarcual cartilage between cartilageneously

tipped uncinate process of epibranchial 1 and infrapharyngobranchial 2; one or two well-developed

gill rakers on epibranchial 1, two or three ceratobranchial 1; basibranchials 1-3 present, 4 present

as cartilage; basibranchial 1 anteriorly ventral to

basihyal; urohyal articulates with ventral surface of

basibranchial 1. Basihyal toothless, with well-developed median keel anteriorly.
NUMBER 252

FIGURE 1.—Anisochromis straussi, terminal color pattern, USNM 216465, 22.6 mm SL: a, lateral

view; b, anterior portion of dorsal fin enlarged. Anisochromis kenyae, terminal color pattern,

USNM 216415, 25.1 mm SL: c, anterior portion of dorsal fin enlarged (Drawn by J. R. Schroeder.)

Dorsal and ventral hypohyals present; ceratohyal

(= anterior ceratohyal) without "beryciform" foramen, but with dorsal margin excavated; ceratohyal

and epihyal (= posterior ceratohyal) form suturing

joint on both medial and lateral surfaces; six branchiostegals: five attach on ceratohyal, one attaches

at joint between ceratohyal and epihyal; anterior

two branchiostegals attach to ventral notches i?i

ceratohyal.

Dorsal and ventral postcleithra present. Pectoralfin radials 4, ventralmost articulating only with

coracoid; fin rays 14 (rarely 13 or 15 unilaterally);

rays usually all branched except very short dorsalmost ray; two dorsalmost rays articulate with scapula; scapula foramen complete. Pelvic-fin rays I, 4;

spine feeble, inapparent; innermost ray much reduced in size, simple, often nubbin-like, other rays

branched. Dorsal and anal fins each comprise tiny

spine and branched rays (see Table 1 for dorsal and

anal-fin ray counts); last ray in each fin split to

base, both halves of ray articulating with same

pterygiophore (two halves counted as one ray in

enumerating dorsal- and anal-fin rays), posterior

half of split ray usually much reduced in size, unbranched; all but anteriormost two pterygiophores

of dorsal and anal fins consist of autogenous proximal, medial and distal radials; two anteriormost

pterygiophores in each fin consist of fused proximal

and medial radials, no (autogenous?) distal radials

present in anteriormost pterygiophore in each fin,

but distal radials autogenous in next to anteriormost pterygiophore; anteriormost dorsal-fin pterygiophore inserted in space between second and third

neural spines; except for anteriormost two pterygiophores in each fin, dorsal or anal-fin pterygiophorevertebra ratio is 1:1. Predorsal bones 2 (rarely 3).

Caudal fin with 14-16 branched rays all of which

articulate with hypurals; total rays, including procurrent elements, consistently 25; no procurrent

spur (Johnson, 1975); parhypural and hypurals 1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Fir.iRE 2—Anisochromis straussi, ocellated color pattern, USNM 215859, 23.4 mm SL. (Drawn by

J. R. Schroeder.)

and 2 fused into autogenous plate; hypurapophysis

well developed; hypurals 3 and 4 fused to each

other and to urostylar complex; hypural 5 autogenous; two epurals; no autogenous uroneurals; neural spine of preural vertebra 2 usually short, occasionally long.

Vertebrae 10 (rarely 11) + 23-25 = 33-35; epipleural ribs begin on first vertebra and continue

posteriorly to vertebra 21 or 22; pleural ribs begin

on third vertebra and continue posteriorly to vertebra 10, where they may be greatly reduced in size;

no autogenous hemal spines.

No scales on head or dorsal, anal, pectoral, and

pelvic fins. Body completely scaled except for portions of predorsal area and fleshy pectoral-fin base;

caudal fin scaled basally. Scales cycloid anteriorly on

body, becoming ctenoid posteriorly at about level

of pectoral-fin axil (note: ctenii not shown in Figures

1 and 2). Tubed lateral-line scales 30-38. Of 28

specimens on which bilateral counts were made, 7

were bilaterally symmetrical, 9 had higher left side

counts, 12 had higher right side counts; the difference between right and left counts ranged from

1 to 4 scales (x = 1.7). Tubed lateral-line scales terminate well anterior to posterior insertion of dorsalfin base; posteriormost tubed scale, unlike other

tubed scales, often notched on posterior margin.

Several scales of midlateral row on posterior half

of body of some specimens each bearing a pit centrally; anteriormost pitted scale often well in advance of level of posteriormost tubed scale.

Color of freshly preserved specimens (based on

general recollections and color slides of one terminal stage and two ocellated stage specimens). Terminal stage. Head generally bright reddish orange

with adjacent pale and black stripe-like markings

extending posterodorsally from orbit at about twoo'clock position; indistinct, slender, dusky stripe extending from ventral margin of orbit to posterior

tip of maxillary; diffuse, dusky markings elsewhere.

Head color rapidly grading into black anteriorly on

body. Body uniformly black. Black of body extending onto dorsal, anal, caudal, and pelvic fins, at

least basally. Dorsal and anal fins dusky distal to

black basal portions, distalmost margins immaculate. Caudal fin uniformly dusky distal to black

area. Pelvic fin almost entirely black with immaculate distal margin. Pectoral fin: fleshy base dusky,

rayed portion black basally, becoming pale dusky

distally.

Ocellated stage. Head generally pale greenish

ventrally, becoming olive brown dorsally; large,

dark-dusky to black spot on opercle narrowly, irregularly margined with brilliant white; adjacent

pale and black stripe-like markings extending posterodorsally from orbit at about two-o'clock position; with few scattered, irregular, pale and dusky

marks; indistinct, slender, dusky stripe, bordered

by pale spots or stripe, extending from ventral margin of orbit to posterior tip of maxillary. Body generally dusky orangish, grading into dusky greenish

posteriorly; faint indications of several dusky bands

present; prepectoral area olive brown with bright,

pale spots. Dorsal and anal fins dusky greenish ante-
NUMBER 252

TABLE 1.—Frequency distributions for certain meristic characters of Anisochromis straussi and A. kenyae

Species

straussi

kenyae

Species

straussi*

kenyae^

Segmented

dorsal-fin rays

25

9

19

26

70

2

27

2 25

25

28 29

1 1

X

.91

.09

30

2

4

17

5

19

Tubed

31

1

2

Segmented

anal-fin rays Total

18 19 X 32

74 2 17.96

1 17.05 1

lateral-line scales

32 33 34 35 36 37

6 9 13 17 7 7

2 1 1

33

11

19

38

1

vertebrae

34 35

69 1

1

X

34.38

30.84

33

33

X

.87

.00

* Includes bilateral counts on 28 specimens.

^Includes bilateral counts on 3 specimens. Three additional specimens, data

from which are not included in table, had unilateral counts of 29 or 30,

30 or 31, and 31 or 32.

riorly grading into dusky orangish posteriorly; distal margins immaculate. Caudal fin greenish in area

covered by scales, rays greenish or orangish, membranes variably dusky. Pelvic fin irregularly dusky

basally, immaculate distally. Pectoral fin pale greenish basally, pale dusky distally. Specimens obtained

during collections are remembered as having greenish heads and bodies, often with broad orangeyellow area on side, and white-margined, dark spot

on opercle.

Color pattern of preserved specimens. Terminal

stage. Head relatively immaculate dorsally and ventrally; dusky and black markings on side of head

as described for fresh specimens; adjacent black and

pale stripes extending from eye may vary in intensity and distinctiveness, and may be present only as

dark and pale spots; faint dusky stripe extending

anteriorly from orbit at about nine o'clock position; occasionally a dusky spot ventrally on opercle.

Black markings on body and fins essentially as in

fresh specimens. Linear series of pale spots present,

each spot at base of a dorsal-fin ray; occasionally a

few pale spots basally in anterior interradial membranes of dorsal fin and anterodorsally on body.

Ocellated stage. Dark and dusky markings as described for fresh specimens, white areas immaculate; faint, dusky stripe extending anteriorly from

orbit at about nine o'clock position. Some specimens, especially smaller ones, exhibit black spots

on caudal peduncle area, one or two of which are

usually at posterior end of dorsal-fin base; body

rarely with many scattered tiny black spots; up to

12 diffuse, dusky bands present on sides, some restricted to dorsal portion; abruptly pale, roughly

rectangular area often encompassing much of area

from venter to midside dorsally and from pectoralfin axil to level of anterior anal-fin rays posteriorly.

Many degrees of intermediacy in color pattern

between the ocellated and terminal stages are represented by available specimens; however, the specimens were sorted into two groups, each believed to

be homogeneous on the basis of color pattern, prior

to our finding that only one species was involved.

The size-frequency distributions for these sortings

is given in Table 2. It is apparent from this sorting

that specimens below 20.0 mm SL are all clearly

ocellated stages and those above 25.9 mm (almost

all above 24.9 mm) are terminal stages.

COMPARISONS.—Anisochromis straussi is very similar to A. kenyae, the only other species of Anisochromis, but differs from that species in coloration

(particularly of the terminal stage) and in having

higher mean numbers of dorsal and anal-fin rays,

total vertebrae, and tubed lateral-line scales.

J. L. B. Smith (1954b) described the "male"

(equals our terminal stage) of A. kenyae as having

the dorsal and anal fins red with a dusky margin,

a dark blotch anteriorly on the dorsal fin extending

posteriorly to the fifth ray, the blotch anteriorly

edged with yellow, and the fin ventral to the blotch
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Frequency distributions for the two color-pattern types of Anisochrotnis straussi

arranged by standard length classes

Color pattern

Terminal

Ocellated

16.5 17.5 18

Standard length middass (mm)

.5 19.5 20.5 21.5 22.5 23.5 24.5 25.5 26.5 27.5 28.5

yellow anteriorly grading into orange-red posteriorly. We noted no red or yellow color or dark

blotch in the dorsal or anal fins of freshly preserved

terminal stage specimens of A. straussi, in which the

dorsal and anal fins appeared black and dusky, just

as in preserved specimens. In terminal-stage specimens of A. straussi the dorsal fin (Figure \a,b) has

a wide, black basal stripe (punctuated with pale

spots), the pigment of which may be intensified

anteriorly, but does not form a noticeable blotch.

Immediately above the basal stripe is a wide dusky

stripe and the narrow, immaculate distal margin of

the fin. In preserved terminal-stage A. kenyae (Figure \c) the dorsal fin has a narrow, dusky basal

stripe margined dorsally by a broader, immaculate

stripe, which is margined dorsally by an even

broader dusky stripe; an intensified dark blotch occupies much of the distal area of the membranes

between the anterior two to four rays.

Frequency distributions and their means for certain meristic characters for the two species of

Anisochromis are compared in Table 1. Each pair

of means was compared by Mest and the p-value

for each pair was less than 0.001, indicating that the

differences are highly significant.

J. L. B. Smith (1954b) reported that A. kenyae

had 11 precaudal vertebrae. All of the 21 specimens

(including 20 paratypes) of A. kenyae we examined

had 10 precaudal vertebrae, as did 80 of 81 specimens of A. straussi checked for this character (the

exception had 11). Smith did not indicate how

many specimens of A. kenyae were checked for

precaudal counts nor how he defined precaudal

vertebrae. We define precaudal vertebrae as those

vertebrae that lack a hemal spine (the precaudal

vertebrae of the four cleared and stained specimens

of A. straussi we examined also lacked complete

hemal arches).

TYPE SPECIMENS.—All types were collected at

St. Brandon's Shoals within the area between 16°20'

to 16°43'S and 59°35' to 59°41'E during the period

30 March to 11 April 1976. All three of us participated in all the collections, during which we were

assisted by various other participants in the expedition to St. Brandon's: L. H. Strauss, P. West, C.

Marshall, W. O. Nordlinger, P. W. Manhard. More

detailed information is available from the original

field data sheets filed in the; Division of Fishes,

National Museum of Natural History, Smithsonian

Institution.

Holotype: USNM 216462, terminal-stage color

pattern, 25.5 mm SL, field number VGS 76-7.

Paratypes: USNM 215859, 26 specimens (including

4 cleared and stained), 18.7-26.1 mm SL, VGS 76-7;

USNM 216463, 19 specimens, 16.4-27.8 mm SL,

VGS 76-11; USNM 216465, 3 specimens, 22.6-23.9

mm SL, VGS 76-1; USNM 216464, 1 specimen,

26.8 mm SL, VGS 76-6; USNM 216466, 2 specimens, 26.5-28.3 mm SL, VGS 76-12; BMNH

1976.8.24.1-10, 10 specimens, 16.6-25.3 mm SL,

VGS 76-10; CAS 37640, 15 specimens, 16.1-24.9

mm SL, VGS 76-9; AMNH 35892, 6 specimens,

22.0-28.0 mm SL, VGS 76-17; RUSI 862, 2 specimens, 17.2-25.5 mm SL, VGS 76-8.

ETYMOLOGY.—Named in honor of Lewis H.

Strauss of Washington, D.C., who conceived, organized, produced, and participated in the expedition to St. Brandon's that netted a scientifically,

highly valuable collection of fishes and other marine organisms.

DISTRIBUTION AND HABITAT NOTES.—Anisochromis straussi is known only from the isolated and

relatively tiny area of St. Brandon's Shoals (= Cargados Carajos) in the southwestern Indian Ocean.

Specimens were obtained only in rotenone collections. All of the 16 rotenone collections made at

St. Brandon's were in rocky areas that included

dead and live coral (live coral was uncommon at

all but one of these areas). A. straussi was taken

in nine of these 16 collections at an overall depth

range of about 0.25-11 meters (recorded in the

field as estimates of 0.5-35 feet), and all nine col-
NUMBER 252

lections were proximate to areas exposed at low

tide. Of the seven rotenone collections where A.

straussi was not obtained, four were made at depths

greater than 15 meters and were well removed

from areas exposed at low tide. The other three

areas did not appear to differ much from stations

where A. straussi was obtained. We often noted

during collections that specimens of A. straussi

were lying on the bottom adjacent to isolated,

small (perhaps less than 0.5 meter in diameter), live

coral heads with surfaces composed of tiny fingerlike projections. Our presumption is that the

Anisochromis were living on the corals.

Anisochromis kenyae, apparently unlike A.

straussi, is widely distributed: occurring on the

east African coast, at least from 3°-14° south latitude (J. L. B. Smith, 1954b; M. M. Smith, pers.

comm.). A. kenyae was collected from pools in

reefs at about low-tide mark (J. L. B. Smith, 1954b).

Reproductive Biology

The reproductive tract of Anisochromis straussi

is of the usual perciform configuration, in which

the right and left gonads fuse posteriorly to form

a common oviduct that ends immediately behind

the anus. There is no ovipore; the eggs apparently

leave the body through a rupture in the connective

tissue at the end of the common oviduct.

In Anisochromis each gonad is a hollow sac with

a single dorsal ridge of germinal tissue. Transformation occurs when the germinal epithelium ceases

to produce oocytes, and seminiferous crypts proliferate and begin to produce sperm. At this time

sperm passages develop in the dorsal walls of the

gonad and join in a common sperm duct in the

dorsal and posterior wall of the common oviduct.

In specimen USNM 216464-1, a ripe female that

apparently spawned some time before it was collected (evidenced by oocytic debris in the germinal

ridge), the sperm duct is visible near the union of

the gonads but not farther posteriorly. This specimen does not have functional seminiferous crypts.

Specimen USNM 216463-3 has functional crypts,

early stage oocyte remnants, and a complete sperm

duct that runs in close proximity to the ureter,

but none of our serial sections are complete enough

for us to determine if the urinary and sperm ducts

exit separately or unite before leaving the body.

Specimen USNM 215859-9, a mature male with

few oocyte remnants, was sectioned frontally. The

posterior half of the testis has large crypts that are

filled with a weakly staining clear material. This

region appears similar to the so-called seminal

vesicles that appear in certain gobies, toadfishes,

and some serranoids. Their function is unknown.

The urinary bladder of both sexes is conspicuously enlarged with thick spongy walls. We are

not aware of similar elaboration of the urinary

bladder in any other fish, but this may not be

significant.

Table 3 presents the results of histological examination of the gonads of 12 specimens of Anisochromis straussi. In general, specimens with ocellated stage color pattern are smaller and are females, and specimens with terminal stage color

pattern are larger and are males, but there is considerable overlap. The overlap is not surprising

and is comparable to that found in other transforming hermaphrodites (C. L. Smith, 1967). The

presence of a nonfunctional lumen in the testis

(remnants of the ovarian lumen) and oocyte remains among the testicular acini are further evidence that this species is a protogynous hermaphrodite.

Relationships of the Anisochromidae

J. L. B. Smith (1954b) described the Anisochromidae in the same paper in which he described

the genus Anisochromis. Without explanation,

Smith stated that the Anisochromidae were most

closely related to the Pseudochromidae, in which

he (1954a) earlier recognized two subfamilies.

Pseudochrominae and Pseudoplesiopinae (Pseudochromidae was first proposed as a family-group

name by Muller and Troschel, 1849, and Pseudoplesiopidae by Bleeker, 1875, who included a group

Pseudoplesiopini among the three groups he recognized in his family Pseudochromidoides). The only

other important study treating the classification of

the Anisochromidae was that of Bohlke (1960),

who recognized three families Anisochromidae,

Pseudochromidae, Pseudoplesiopidae — henceforth

referred to jointly as the pseudochromoids) for the

fishes treated by J.L.B. Smith (1954a, b), but Bohlke

did not indicate whether he considered the families

to be closely related. Most recently, Lubbock (1975,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 3.—Results of histological examination of gonads of Anisochromis straussi (oocyte stage

as denned by C. L. Smith and Young, 1966)

Catalog

number

CAS 37640

CAS 37640

USNM 216463

CAS 37640

USNM 215859

CAS 37640

USNM 216463

USNM 216463

USNM 216463

USNM 216464

USNM 216463

AMNH 35892

Specimen

number

12

10

11

3

9

6

2

3

1

1

4

2

Standard

length (mm)

16.1

19.2

21.4

22.9

24.2

24.9

25.3

26.3

26.7

26.8

27.8

28.0

Color pattern

staqe

ocellated

ocellated

ocellated

terminal

terminal

ocellated

terminal

terminal

terminal

terminal

terminal

terminal

Gonad condition

immature female, stage 2 oocytes, no obvious male

tissue

immature female, stage 2 oocytes, no obvious male

tissue, no sperm duct

immature female, stage 2 oocytes, no obvious male

tissue

inactive male, some sperm, stage 2 oocytes

active male, many sperm, few stage 2 oocytes

active female, stage 4 oocytes, no obvious male

tissue

immature female, stage 2 oocytes, no obvious male

tissue

transforming, sperm duct complete, many active

sperm crypts, some sperm, stage 2 oocytes

abundant

inactive male, few sperm, stage 1 and 2 oocytes

active female, oocyte debris, late stage 3 oocytes,

partial sperm duct, no obvious male tissue

active male, abundant sperm, stage 2 oocytes

highly developed male, many spermatocytes, few

sperm, stage 2 oocytes

1976), in describing new species of Pseudochromis

and Chlidichthys (which Bohlke placed in the

Pseudochromidae and Pseudoplesiopidae, respectively), followed J. L. B. Smith (1954a) and did

not recognize the Pseudoplesiopidae as distinct

from the Pseudochromidae. Lubbock (pers. comm.)

suggested to Springer the possibility that the Anisochromidae ought to be synonymized with the

Pseudochromidae.

Bohlke's 1960 study was an attempt to define the

families of serranoid fishes with disjunct lateral

lines. The fishes of the seven families (pseudochromoids, Plesiopidae, Grammidae, Pseudogrammidae, Acanthoclinidae) Bohlke recognized in this

group have a long and involved classificatory history. Most have been included in the Serranidae

at one time or another, and all have specialized

character states of the lateral line, including: interrupted, incomplete posteriorly, multiple lateral

lines, and no lateral line at all (we consider the

primitive state for the lateral line in perciforms

to be a single, uninterrupted lateral line extending

the entire length of the body). Without explanation, however, Bohlke neglected to include some

families (e.g., Owstoniidae and Opistognathidae)

whose members have posteriorly incomplete lateral

lines and which have been considered (for instance,

Giinther, 1860; Boulenger, 1901, Jordan and Snyder, 1902; Fowler, 1934) to be closely related to,

or even included in, families he did treat in his

study. Nevertheless, Bohlke's study, as the most

recent and comprehensive treatment of the pseudochromoids, is a point of departure from which to

attempt to determine if these three families are

closely related and whether we should recognize

one or more families for them. To answer the first

of these two questions it was necessary to determine if the pseudochromoids form a monophyletic

group among the fishes Bohlke treated. We believe

the pseudochromoids are monophyletic and we

distinguish them from the other compared families

on the presence of two synapomorphies: the reduced number of dorsal-fin spines and the nature

of the connection between the uncinate process of

the first epibranchial and the second infrapharyngobranchial.
NUMBER 252

Based on the generalized perciform fishes Percichthys and Morone (Percichthyidae), we consider

the presence of 10 dorsal-fin spines to be the unspecialized condition for perciform fishes, among

which we include the serranoids (for a discussion

of some of the most generalized perciforms, see

Gosline, 1966).1 Bohlke's families appear to fall

into four groups based on number of dorsal-fin

spines: Group I, Anisochromidae (1 spine), Pseudoplesiopidae (1 or 2, primitively 2), Pseudochromidae (2 or 3, primitively 3); Group II, Pseudogrammidae 6 to 8, primitively 8 (note: the anteriormost

spine is vestigial, visible only in osteological preparations, and has been overlooked often by authors);

Group III, Grammidae (11 to 13, primitively 11),

Plesiopidae (11 to 14, primitively 11); Group IV,

Acanthoclinidae (18 to 21, primitively 18). The

Grammidae and Plesiopidae (Group III) appear to

be least specialized for number of dorsal-fin spines,

with specialization proceeding in two directions:

increase and decrease in number of spines. The

pseudochromoids have the fewest spines and are

well separated in this character from the group

(II) with the next most reduced number of spines.

In the pseudochromoids epibranchial 1 bears an

uncinate process that is cartilagenously tipped and

is connected ligamentously directly to infrapharyngobranchial 2. A similar connection between these

two bones occurs in the beryciform fishes and thus

might be considered to be an unspecialized state

when present in perciforms. In the Percichthyidae,

however, there is an interarcual rod of cartilage

between the uncinate process of epibranchial 1 and

infrapharyngobranchial 2. Rosen and Greenwood

(1976:25) assert that the presence of this interarcual cartilage is a synapomorphy of the perciforms, and many perciforms exhibit the cartilage,

including all the fishes Bohlke (1960) discussed

except the pseudochromoids. The lack of an interarcual cartilage in the pseudochromoids represents

either retention of a primitive, beryciform condition or a specialization, through loss, convergent on

the beryciform condition. Many specialized (usually

benthic) perciform fishes lack an interarcual cartilage (for instance: blennioids, stichaeoids, cottoids,

batrachoids, trichonotids, pholydichthyids, callinonymids, trichodontids, uranoscopids; Springer

and Freihofer, 1976). In view of the numerous

specializations shared by the pseudochromoids and

many perciforms with an interarcual cartilage (including, lack of an orbitosphenoid, fewer than six

segmented pelvic-fin rays, fewer than 19 principal

caudal-fin rays), we believe that the lack of an

interarcual cartilage in the pseudochromoids is a

specialization convergent with the primitive beryciform character state.

Interrelationships of the Pseudochromoids

Based on number of dorsal-fin spines, it might

appear that the Pseudogrammidae are the sister

group of the pseudochromoids, but we believe, as

did Bohlke, that the pseudogrammids are not

closely related to any of the other families he

treated.2 Aside from a disjunct lateral line, we find

no specialization common to the pseudogrammids

and any of the other families that is not also common to a large number of families Bohlke did not

treat (a disjunct lateral line occurs also in the

Nannopercidae, some Blenniidae, and some Brotulidae, for instance, and may not necessarily indicate

relationships). The Pseudogrammidae appear to be

generally less specialized than the other families

Bohlke treated, and do not exhibit some specializations that occur throughout the other families. For

instance (primitiveness here based on character

states in Percichthys), the pseudogrammids are

primitive in having seven branchiostegals, the epihyal and ceratohyal not sutured together (but sutured on medial surface in the percichthyid genus

Morone!), and in having hypurals 1-4 autogenous,

whereas the other families are more specialized in

having five or six branchiostegals, the epihyal and

1

 The Perciformes is a widely accepted group for which

monophyly has not been demonstrated. It cannot be demonstrated, therefore, that the pseudochromoids are a member

of the Perciformes. Our out-group comparisons are based on

the assumptions that the Perciformes is a monophyletic

group and the pseudochromoids are a member of it.

- Bohlke (1960) believed that the pseudogrammids were

closely related to Grammistops (Grammistidae). Gosline (1960,

1966) synonymized the Pseudogrammidae with the Grammistidae, and Schultz (1966) agreed. C. L. Smith and E. H. Atz

(1969) concluded on the basis of gonodal structure, and Randall, et al. (1971) on the basis of the toxin grammistin, that

the grammistids and pseudogrammids are not closely related.

Kendall (J976) considered the grammistids and pseudogrammids to form a portion of a subfamily Grammistinae, of the

Serramidae.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ceratohyal sutured together (both surfaces), hypurals 1 and 2 fused together, and in having

hypurals 3 and 4 fused together and also fused to

the urostylar complex. It, therefore, seems more

reasonable that one of the other families Bohlke

treated would be a better candidate as a sister

group for the pseudochromoids. This may be true,

but we were unable to find a shared specialization

that was unique to the pseudochromoids and one

or two of the other three families, with one possible exception, Lipogramma, which Bohlke (1960)

described and assigned, for lack of another possibility, to the Grammidae.

Lipogramma, which has no lateral line, exhibits

one specialization (and possibly a second), which is

found otherwise only in the pseudochromoids

among all the fishes Bohlke treated (including the

Pseudogrammidae): infrapharyngobranchial 1 is

absent (specialized), and there is a cordlike ligament attaching the lateral surface of the ceratohyal

to the dentary. We are uncertain of the ligament's

existence in Percichthys (our specimens were in

poor condition), but it is absent in Morone. The

loss of infrapharyngobranchial 1, or its lack of ossification, has occurred independently in a wide variety of fishes (for instance, blennioids, cottoids,

leptoscopids, stichaeoids) and may not necessarily

indicate relationships. We have not searched widely

for the ligament, which occurs in various character

states in at least the Clinidae sensu stricto (but not

other blennioids), Congrogadidae, and some Serranidae (but not others). Considerably more study

will be necessary to determine whether Lipogramma, or any of the other fishes Bohlke discussed, is the sister group of the pseudochromoids.

Fowler (1934) placed Parasphenanthias Gilchrist

(which Barnard, 1927:492, had synonymized with

Owstonia of the Owstoniidae) in the Pseudochromidae. Myers (1935) indicated that although Parasphenanthias and Owstonia were closely related,

they were not close to the Pseudochromidae, but

Myers was using Pseudogramma as a basis for his

comparison. Nevertheless, we agree with Myers that

the owstoniids are not closely related to the pseudochromoids. We also agree with Okada and Suzuki

(1956) that the Owstoniidae should be synonymized

with and under the Cepolidae, and consider them

as such in the following discussion.

The cepolids exhibit a number of specializations

that occur also in the pseudochromoids. Most of

these specializations are shared as well with the

other families of fishes that Bohlke treated, but

one specialization, the presence of only two or

three dorsal-fin spines, occurs only among the

pseudochromoids. We believe that this character

is convergent in the cepolids and pseudochromoids.

The cepolids are less specialized than the pseudochromoids (or any of the other families Bohlke

treated) in having autogenous hypurals 3 and 4

(these hypurals appear to be fused in some specimens or species of cepolids, but they are primitively autogenous within the family; Monod (1968,

fig. 683) misinterpreted the hypurals of Cepola: his

hypural 1 represents fused hypurals 1 and 2, his

hypural 2 is 3, and his hypural 3 is 4). The cepolids are also less specialized than the pseudochromoids in having an interarcual cartilage between

epibranchial 1 and infrapharyngobranchial 2 and an

infrapharyngobranchial 1, and are perhaps less specialized in lacking the ceratohyal-dentary ligament.

The cepolids are more specialized than the pseudochromoids (and other fishes Bohlke treated) in having: the first dorsal-fin pterygiophore inserted between the first and second neural spines (primitive

condition: between spines 2 and 3), the last dorsal

and anal-fin pterygiophores each supporting a single ray (primitive condition: supporting two elements; also known as the last-ray-split-to-the-base

condition), a single postcleithrum on each side

(primitive condition: two postcleithra, one dorsal

and one ventral, on each side), and no predorsal

bones (primitive condition: 3 bones; pseudochromoids have 2 or 3).

The conjunction of more primitive states for the

caudal-fin hypurals, interarcual cartilage, infrapharyngobranchial 1, and ceratohyal-dentary ligament, together with the more specialized states for

dorsal-fin pterygiophores, predorsal bones, and postcleithra, leads us to believe that the cepolids are

not the sister group of the pseudochromoids.

The Opistognathidae have been considered to be

close relatives of the Pseudochromidae (Giinther,

1860:254; Boulenger, 1901; Jordan and Snyder,

1902) and Acanthoclinidae (Gosline, 1968, fig. 12,

and pers. comm.), and although Bohlke (1960)

omitted consideration of the opistognathids, their

relationships must be examined. The opistognathids exhibit a number of specializations that are also
NUMBER 252 11

found in the pseudochromoids (lateral line incomplete and terminating below the dorsal fin, six

branchiostegals, epihyal and ceratohyal sutured together, hypurals 1 and 2 fused, hypurals 3 and 4

fused to each other and to urostylar complex), but

these are the same specializations that characterize

the other families Bohlke (1960) treated. We found

no specialization that the opistognathids shared

with the pseudochromoids to the exclusion of the

other families. We, therefore, have no basis for

relating the opistognathids more closely to the

pseudochromoids than to any of the other families.

There is, in addition, no basis for excluding the

Opistognathidae from Bohlke's serranoid group of

fishes with disjunct lateral lines. (The opistognathids also possess an interarcual cartilage, 9-12 dorsalfin spines, infrapharyngobranchial 1, no ceratohyaldentary ligament, last dorsal and anal-fin pterygiophores each supporting two external elements, and

first dorsal-fin pterygiophore inserted between the

second and third neural spines. The opistognathids

are more specialized than the other families Bohlke

treated in having: the lateral line imbedded in the

skin, rather than occurring on scales; no teeth on

infrapharyngobranchial 2, a convergent condition

occurring otherwise only in Acanthoplesiops of the

Acanthoclinidae among the fishes Bohlke treated;

and the condition of the segmented pelvic-fin rays

is unusual: the anterior two are stout and simple,

whereas the posterior three are weak and branched.)

In summary, considering the families Bohlke

(1960) discussed, together with the Opistognathidae

and Cepolidae, we believe that the pseudochromoid

fishes form a monophyletic group, whose sister

group we are unable to hypothesize reasonably.

Synonymization of the Anisochromidae and

Pseudoplesiopidae

Assuming that pseudochromoids form a monophyletic group, we wished to determine whether

each of the three families that the group comprises

is monophyletic. We surveyed the members of each

family for various characters in a search for autapomorphies, or synapomorphies shared by only

two of the three families. Of numerous characters

examined, the most useful are given in Table 4,

together with the character states for each. We are

unable to propose degree of specialization for characters 8 and 10 in the table, but based on more

generalized perciform fishes (particularly Percichthys), we propose the following directions for specialization of the other characters (least specialized

condition presented first):

1. From 5 to 4 to 3 segmented pelvic-fin rays.

2. From all branched, to some branched, to all

simple rays.

3. From 3, to 2, to 1 dorsal-fin spine, and from

strong to weak spines.

4. From all branched to most simple segmented

dorsal-fin rays.

5. From 3 to 2 to 1 anal-fin spine, and from

strong to weak spines.

6. From scales to no scales on dorsal fin.

7. From few medial radials fused with proximal

radials (or almost all medial radials autogenous),

to several fused, to all fused with their respective

proximal radials in each fin.

9. From ventralmost pectoral-fin radial joining

only coracoid, to joining both coracoid and scapula.

11. From two (anterior and posterior) disjunct

series of tubed lateral-line scales, to an ter odor sally

tubed series only, to only one tubed scale anterodorsally.

12. From head scaled to not scaled.

13. From palatine teeth present to teeth absent.

14. From more than 7 to less than 4 gill rakers

on lower limb of first arch.

15. From branchiostegal membranes separate to

membranes fused ventroposteriorly.

16. From parhypural autogenous to parhypural

fused to hypural 1.

17. From preural hemal spine 2 autogenous to

spine fused to centrum.

18. From ectopterygoid and mesopterygoid both

articulating closely with palatine, to both well

separated from palatine (resulting from shortening

of the anterior processes of the ectopterygoid and

mesopterygoid).

19. Ligament from ceratohyal: from connecting

to dentary at symphysis, to connecting on coronoid

(ascending) process of dentary (specialization inferred from condition in Lipogramma and the

Clinidae, and less certainly in Serranus, where the

ligament connects at the symphysis or point anterior to coronoid process).

The Anisochromidae exhibit at least five autopomorphies (12-15, 19) and can be considered to be
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 4.—Comparison of certain characters in the three families of pseudochromoid fishes

(family composition as defined by Bohlke, 1960; * denotes autapomorphic characters and, where

the same character in another family is accompanied by two asterisks, further denotes that the

phylogeny of the autapomorphy is presumed to have passed through the more primitive derived

state exhibited by the character with two asterisks; •• denotes synapomorphic characters and,

where the same character in another family is accompanied by a single asterisk, further denotes

that the synapomorphy is presumed to have occurred, as a percursor, in the phylogeny of the

family in which the character bears a single asterisk)

Characters

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

Pelvic-fin rays

Segmented pelvic-fin

rays

Dorsal-fin spines

Segmented dorsal-fin

rays

Anal-fin spines

Scales on dorsal fin

Medial radials in

dorsal and anal fins

Pectoral-fin rays

Ventralmost

pectoral-fin radial

Vertebrae

Tubed lateral-line

scales

Head scaled

Palatine teeth

Gill rakers on

lower limb of

first arch

Branchiostegal

membranes

Par.hypural

Hemal spine of

preurai centrum 2

Ectopterygoid and

mesopterygoid

Ligament from

ceratohyal attaches

to dentary

Pseudochromidae

1,5

all branched

II or III, at least some

strong

vary from all branched to

most simple

III, at least some strong

present or absent

several fused to proximal

radials in each fin**

16-20

joins coracoid

10-11 + 16-17 = 26-28

2 disjunct series:

1 anterodorsally,

1 midlateral posteriorly

yes

present

more than 7

margins on each side

separate ventroposteriorly

autogenous

autogenous

articulate closely with

palatine

at symphysis

Families

Pseudoplesiopidae

1,3 or 1,4**

all simple*

I or II, weak**

most simple*

I-III, weak**

absent**

all, or almost all, fused

to proximal radials*

17-19

joins coracoid and scapula*

10-13 + 17-20 = 27-33

1 scale anterodorsally*

yes

present

more than 7

margins on each side

separate ventroposteriorly

fused to hypural 1**

fused to centrum**

articulate closely with

palatine

at symphysis

Anisochromidae

1,4**

3 branched, 1 simple**

I*. weakest*

all branched

I*, weakest*

absent**

few fused to proximal

radials in each fin

13-15 (14 modally)

joins coracoid

10 (rarely 11) + 22-25 = 32-35

1 series anterodorsally**

no*

absent*

less than 4*

ventroposterior margins fused

across ventral surface of

head*

fused to hypural 1**

fused to centrum**

well separated from palatine*

on coronoid process*

monophyletic (not unexpected as only one genus

is involved). The Pseudoplesiopidae exhibit at least

four autapomorphies (2,4,9, 11) and may also be

considered to be monophyletic (the presence of

simple segmented dorsal-fin rays in pseudoplesiopids and some pseudochromids is here considered

to be a convergence as the unspecialized condition

for rays is also present in pseudochromids and

must be considered the primitive state for all

pseudochromids). The Pseudochromidae, however,
NUMBER 252 13

lack autapomorphous characters; thus, there is reason to doubt their monophyly. The pseudochromids exhibit one possible synapomorphy (7) with

the pseudoplesiopids but none with the anisochromids, and one might consider this a basis for recognizing the pseudochromids as the sister group of

the pseudoplesiopids. The anisochromids, however,

exhibit at least six synapomorphies (1,2,5,11,16,17)

with the pseudoplesiopids and the likelihood that

these two families form a sister group seems more

plausible than the pseudochromid-pseudoplesiopid

group possibility.

If the Pseudochromidae are a polyphyletic group

it is probable that some member genera are more

closely related to the Pseudoplesiopidae and others

more closely related to the Anisochromidae. In any

event, the Pseudochromidae presently appear to

form a primitive residue pseudochromoid group

and there is no basis for maintaining them as a

distinct family. J. L. B. Smith (1954a) combined

the Pseudochromidae and Pseudoplesiopidae (although maintaining them as subfamilies), and Lubbock (1975,1976), currently the most active student

of both groups, agreed (although he did not cite

subfamilies in his studies). J. L. B. Smith (1954b)

later recognized the Anisochromidae as distinct

from his Pseudochromidae. Based on synapomorphies, however, the Anisochromidae and Pseudoplesiopidae are more closely related to each other

than either is to the Pseudochromidae. Under the

circumstances there seems little reason to recognize

either two or three families among the pseudochromoids, and we here propose to combine them

all under the oldest of the three family names,

Pseudochromidae. Concurrently, we do not recognize subfamilies in the Pseudochromidae. In effect,

we have chosen the lowest (least equivocal) phylogenetic level for which there is evidence for monophyly among the pseudochromoids as the level

for family recognition.

Diagnosis of the Pseudochromidae

Small, often colorful, Indo-West Pacific reefdwelling perciform fishes, seldom attaining 125 mm

SL (to about 200 mm in Labracinus); many species

under 75 mm. Dorsal fin 1-111,21-37; spines often

inconspicuous; segmented rays varying from all

branched to most unbranched. Predorsal bones 2

or 3. Anteriormost dorsal-fin pterygiophore inserted

between second and third neural spines. Terminal

dorsal and anal-fin pterygiophore each supporting

two external elements (last ray split to base). Pelvicfin insertion below to well in advance of level of

anteriormost insertion of pectoral fin. Pelvic fin

consisting of a spine and: five branched rays, or

three branched and one simple ray, or three or four

simple rays. Vertebrae 10-13 + 16-25 = 26-35.

Hypurals 1 and 2 (and sometimes parhypural)

fused into autogenous plate. Hypurals 3 and 4

fused to each other and to urostylar complex. No

autogenous uroneurals. No interarcual cartilage between uncinate process of epibranchial 1 and infrapharyngobranchial 2. Infrapharyngobranchial 1

absent. Six branchiostegals. Epihyal and ceratohyal

suturally joined. Preopercle margin entire. Tubed

lateral-line scales in two disjunct series (one dorsoanteriorly, one midlateral posteriorly), one series

(dorsoanteriorly), or present as only one tubed

scaje anteriorly; centrally pitted scales often present

following dorsal tubed scale series or preceding

midlateral posterior series. Cordlike ligament present attaching lateral surface of ceratohyal either

to coronoid process or symphysial area of dentary.
Literature Cited

Barnard, K. H.

1927. Monograph of the Marine Fishes of South Africa, 2.

Annals of the South African Museum, 21:419-1065.

Bleeker, P.

1875. Sur la famille des Pseudochromidoides et revision de

ses especes insulidiennes. Verhandelingen der Koninklijke Akademie van Wetenschappen, 15:1-32, 3

plates.

Bohlke, J. E.

1960. Comments on Serranoid Fishes with Disjunct Lateral

Lines, with Description of a New One from the

Bahamas. Notulae Naturae, 330:1-11.

Boulenger, G. A.

1901. Notes on the Classification of Teleostean Fishes, I:

On the Trachinidae and Their Allies. Annals and

Magazine of Natural History, series 7, 8: 261-271.

Fowler, H. W.

1934. Fishes Obtained by Mr. H. W. Bell-Marley Chiefly

in Natal and Zululand in 1929 to 1932. Proceedings

of the Academy of Natural Sciences of Philadelphia,

86:405-514.

Gosline, W. A.

1960. A New Hawaiian Percoid Fish, Suttonia lineata,

with a Discussion of Its Relationships and a Definition of the Family Grammistidae. Pacific Science,

14(l):28-38.

1966. The Limits of the Fish Family Serranidae, with

Notes on Other Lower Percoids. Proceedings of the

California Academy of Sciences, series 4, 33(6):91-112.

1968. The Suborders of Perciform Fishes. Proceedings of

the United States National Museum, 124(3647): 1-78.

Giinther, A.

1860. Catalogue of the Acanthopterygian Fishes in the

Collection of the British Museum. Volume 2, xxi +

548 pages. London.

Johnson, G. D.

1975. The Procurrent Spur: An Undescribed Perciform

Caudal Character and Its Phylogenetic Implications.

Occasional Papers of the California Academy of Sciences, 121:1-23.

Jordan, D. S., and J. O. Snyder

1902. A Review of the Trachinoid Fishes and Their Supposed Allies Found in the Waters of Japan. Proceedings of the United States National Museum,

24(1263):461-497.

Kendall, A. W., Jr.

1976. Predorsal and Associated Bones in Serranid and

Grammistid Fishes. Bulletin of Marine Science,

26(4): 585-592.

Lubbock, R.

1975. Fishes of the Family Pseudochromidae (Perciformes)

in the Northwest Indian Ocean and Red Sea. Journal of Zoology, 176:115-157.

1976. Fishes of the Family Pseudochromidae (Perciformes)

in the Central Indian Ocean. Journal of Natural

History, 10:167-177.

Monod, T.

1968. Le complexe urophore des poissons teleosteens.

Memoires de L'Institut Fondamental D'Afrique

Noire, 81:1-705.

Miiller, J., and F. H. Troschel

1849. Horae Ichthyologicae. Volume 3, 28 pages, 5 plates,

1 foldout table. Berlin.

Myers, G. S.

1935. A New Genus of Opisthognathid Fishes. Smithsonian

Miscellaneous Collections, 91 (23): 1-5.

Okada, Y., and K. Suzuki

1956. On the Similarity of the Osteological Characters

Found between Owstoniidae and Cepolidae. Report

of Faculty of Fisheries, Prefectural University of

Mie, 2(2): 185-194.

Randall, J. £., K. Aida, T. Hibiya, N. Mitsuura, H. Kamiya,

and Y. Hashimoto

1971. Grammistin, the Skin Toxin of Soapfishes, and Its

Significance in the Classification of the Grammistidae. Publications of the Seto Marine Biological

Laboratory, 19(2-3): 157-190, figures 13-23.

Rosen, D. E., and P. H. Greenwood

1976. A Fourth Neotropical Species of Synbranchid Eel

and the Phytogeny and Systematics of Synbranchiform Fishes. Bulletin of the American Museum of

Natural History, 157(1): 1-69.

Schultz, L. P.

1966. Pseudorhegma diagramma, a New Genus and Species of Grammistid Fish, with a Key to Genera of

the Family and to the Species of the Subfamily

Pseudogramminae. Ichthyologica, The Aquarium

Journal, 37(4): 185-194.

1967. A Review of the Fish Genus Labracinus Schlegel,

Family Pseudochromidae, with Notes on and Illustrations of Some Related Serranoid Fishes. Ichthyologica, The Aquarium Journal, 39(1): 19-40.

Smith, C. L.

1967. Contribution to a Theory of Hermaphroditism.

Journal of Theoretical Biology, 17:76-90.

Smith, C. L., and E. H. Atz

1969. The Sexual Mechanism of the Reef Bass Pseudogramma bermudensis and Its Implications in the

Classification of the Pseudogrammidae (Pisces: Perci14
NUMBER 252 15

formes). Zeitschrift fur Morphologie und Ocologie Annals and Magazine of Natural History, series 12,

der Tiere, 65(4):315-326. 7:195-208.

Smith, C. L., and P. H. Young 19 - 4 b Th e Anisochromidae, a New Family of Fishes from

1966. Gonad Structure and Reproductive Cycle of the ,-. . . c . . , , . , • , A, . > »..

1

 ' East Africa. Annals and Magazine of Natural titsKelp Bass, Paralabrax clathratus (Girard), with . ,„ „ „„„ „„„ ,

„

 K

 .„,.. . r , c J tor*> serie s 12- 7:298-302, plate 6.

Comments on the Relationships of the Serranid

Genus Paralabrax. California Fish and Game, Springer, V. G., and W. C. Freihofer

54(4):283-292- 1976. Study of the Monotypic Fish Family PholidichthyiSmith, J. L. B. dae (Perciformes). Smithsonian Contributions to

1954a. Pseudolesiopsine Fishes from South and East Africa. Zoology, 216:1-43.

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Manuscripts intended for series publication receive substantive review within their

originating Smithsonian museums or offices and are submitted to the Smithsonian Institution

Press with approval of the appropriate museum authority on Form SI-36. Requests for

special treatment—use of color, foldouts, casebound covers, etc.—require, on the same

form, the added approval of designated committees or museum directors.

Review of manuscripts and art by the Press for requirements of series format and style,

completeness and clarity of copy, and arrangement of all material, as outlined below, will

govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art.

Copy must be typewritten, double-spaced, on one side of standard white bond paper,

with iyA" margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not

stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages.

Front matter (preceding the text) should include: title page with only title and author

and no other information, abstract page with author/title/series/etc., following the established format, table of contents with indents reflecting the heads and structure of the paper.

First page of text should carry the title and author at the top of the page and an unnumbered footnote at the bottom consisting of author's name and professional mailing address.

Center heads of whatever level should be typed with initial caps of major words, with

extra space above and below the head, but with no other preparation (such as all caps or

underline). Run-in paragraph heads should use period/dashes or colons as necessary.

Tabulations within text (lists of data, often in parallel columns) can be typed on the text

page where they occur, but they should not contain rules or formal, numbered table heads.

Formal tables (numbered, with table heads, boxheads, stubs, rules) should be submitted as camera copy, but the author must contact the series section of the Press for editorial attention and preparation assistance before final typing of this matter.

Taxonomic keys in natural history papers should use the alined-couplet form in the

zoology and paleobiology series and the multi-level indent form in the botany series. If

cross-referencing is required between key and text, do not include page references within the

key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon,

author, year:page), with a full reference at the end of the paper under "Literature Cited."

For the botany series, the long form (taxon, author, abbreviated journal or book title, volume,

page, year, with no reference in the "Literature Cited") is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed

at the bottom of the text page on which the reference occurs. Extensive notes must appear at

the end of the text in a notes section. If bibliographic footnotes are required, use the short

form (author/brief title/page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a

"Literature Cited" at the end of the text) must be used in place of bibliographic footnotes in

all scientific series and is strongly recommended in the history and technology series:

"(Jones, 1910:122)" or ".. . . Jones (1910:122)."

Bibliography, depending upon use, is termed "References," "Selected References," or

"Literature Cited." Spell out book, journal, and article titles, using initial caps in all major

words. For capitalization of titles in foreign languages, follow the national practice of each

language. Underline (for italics) book and journal titles. Use the colon-parentheses system

for volume/number/page citations: "10(2):5-9." For alinement and arrangement of

elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but

must be submitted at the end of the manuscript—with as many legends typed, doublespaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted separately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.)

can be intermixed throughout the printed text. They should be termed Figures and should

be numbered consecutively. If several "figures" are treated as components of a single larger

figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: "Figure 9b.." If illustrations are intended to be

printed separately on coated stock following the text, they should be termed Plates and any

components should be lettered as in figures: "Plate 9.b_." Keys to any symbols within an

illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as "mm, ft,

yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled-out fractions: "two-thirds." (3)

Spell out numbers "one" through "nine" in expository text, but use numerals in all other

cases if possible. (4) Use the metric system of measurement, where possible, instead of

the English system. (5) Use the decimal system, where possible, in place of fractions.

(6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular listings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front

matter and ALL legends, etc., at the back of the text—in the following order: (1) title page,

(2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes,

(7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.
«

•

,111/.