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
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 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
A B S T R A C 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 Contri-
butions 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 corre-
lated 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 Aniso-
chromidae, Pseudoplesiopidae, and Pseudochromidae form a monophyletic group.
Evidence is also presented that within this group the Pseudoplesiopidae and Aniso-
chromidae form a monophyletic group and each of these two taxa is definable
based on autapomorphies. The Pseudochromidae appear to be a possibly poly-
phyletic primitive residue group that is not definable on the basis of autapomor-
phies. 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. Pseudochro-
midae 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 Aniso-
chromidae, 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 list-
ings or discussions based on the original descrip-
tions (J. L. B. Smith, 1954b) of Anisochromis and
the Anisochromidae, there has been no new infor-
mation presented on these two taxa. During prepa-
ration of the description of our new species, we
became impressed by the similarity of Anisochromis
to fishes of the family Pseudochromidae and Pseudo-
plesiopidae. We were thus prompted to investigate
the systematic relationships of Anisochromis. We
also noted that eggs were often present in the go-
nads 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, Na-
tional 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 osteologi-
cal 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 Mu-
seum 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 Insti-
tute of Ichthyology, Grahamstown, South Africa;
USNM = acronym for former United States Na-
tional 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 fami-
lies, 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 bran-
chial and hyoid arches (poor condition of the speci-
mens precluded clearing and staining). It was not
always possible to determine genus or species for
the specimens of Pseudochromidae and Pseudople-
siopidae. 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); Belonep-
terygion fasciolatum, USNM 211306 (2). ANISOCHROMIDAE:
Anisochromis straussi, USNM 215859 (4). CEPOLIDAE: Cepola
pauclradiata, USNM 213502 (1). GRAMMIDAE: Gramma lo-
reto, USNM 199487 (3); Lipogramma anabantoides, USNM
216405 (2). OPISTOCNATHIDAE: Opistognathus whitehursti,
ANSP 120714 (1); Stalix moenensis, USNM 211963 (1). PER-
CICHTHVIDAE: Morone americana, USNM 109851 (3). PLESIO-
PIDAE: 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); Pseudo-
plesiops 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 fol-
lowing individuals for providing loans of specimens
or information on holotypes: M. Boeseman, Rijks-
museum van Natuurlijke Historic Leiden; A. Kott-
haus, Hamburg; H. Nijssen, Zoologisch Museum,
Amsterdam; M. M. Smith and R. Winterbottom,
RUSI; W. F. Smith-Vaniz, ANSP; P. J. P. White-
head and A. C. Wheeler, BMNH.
A. Spreitzer, NMNH, provided laboratory assist-
ance, and P. Melville, NMNH, prepared the histo-
logical 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 bene-
fited from the criticism of, G. J. Nelson and D. E.
Rosen (AMNH), and W. L. Fink (Museum of Com-
parative 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 seg-
mented anal-fin rays (modally 18), 33-35 total verte-
brae (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 dermosphe-
notic; 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. Vo-
mer toothed. Palatine toothless, well separated from
foreshortened mesopterygoid and ectopterygoid
(ectoterygoid reduced to small blade of bone with-
out anteriorly extending process). Supramaxillaries
absent. Sesamoid articulars (coronomeckelians) ab-
sent. 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; infrapharyngo-
branchials 2,3, and toothplate of 4 present, toothed;
no interarcual cartilage between cartilageneously
tipped uncinate process of epibranchial 1 and in-
frapharyngobranchial 2; one or two well-developed
gill rakers on epibranchial 1, two or three cerato-
branchial 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-devel-
oped 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" fora-
men, but with dorsal margin excavated; ceratohyal
and epihyal (= posterior ceratohyal) form suturing
joint on both medial and lateral surfaces; six bran-
chiostegals: 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. Pectoral-
fin radials 4, ventralmost articulating only with
coracoid; fin rays 14 (rarely 13 or 15 unilaterally);
rays usually all branched except very short dorsal-
most ray; two dorsalmost rays articulate with scap-
ula; scapula foramen complete. Pelvic-fin rays I, 4;
spine feeble, inapparent; innermost ray much re-
duced 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, un-
branched; all but anteriormost two pterygiophores
of dorsal and anal fins consist of autogenous proxi-
mal, 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 anterior-
most pterygiophore; anteriormost dorsal-fin pteryg-
iophore inserted in space between second and third
neural spines; except for anteriormost two pterygio-
phores in each fin, dorsal or anal-fin pterygiophore-
vertebra 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 pro-
current 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 autoge-
nous; two epurals; no autogenous uroneurals; neu-
ral spine of preural vertebra 2 usually short, occa-
sionally long.
Vertebrae 10 (rarely 11) + 23-25 = 33-35; epi-
pleural ribs begin on first vertebra and continue
posteriorly to vertebra 21 or 22; pleural ribs begin
on third vertebra and continue posteriorly to verte-
bra 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 por-
tions 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 differ-
ence between right and left counts ranged from
1 to 4 scales (x = 1.7). Tubed lateral-line scales ter-
minate well anterior to posterior insertion of dorsal-
fin 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 cen-
trally; anteriormost pitted scale often well in ad-
vance of level of posteriormost tubed scale.
Color of freshly preserved specimens (based on
general recollections and color slides of one termi-
nal stage and two ocellated stage specimens). Termi-
nal stage. Head generally bright reddish orange
with adjacent pale and black stripe-like markings
extending posterodorsally from orbit at about two-
o'clock position; indistinct, slender, dusky stripe ex-
tending 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 extend-
ing onto dorsal, anal, caudal, and pelvic fins, at
least basally. Dorsal and anal fins dusky distal to
black basal portions, distalmost margins immacu-
late. Caudal fin uniformly dusky distal to black
area. Pelvic fin almost entirely black with immacu-
late 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, ir-
regularly margined with brilliant white; adjacent
pale and black stripe-like markings extending post-
erodorsally from orbit at about two-o'clock posi-
tion; with few scattered, irregular, pale and dusky
marks; indistinct, slender, dusky stripe, bordered
by pale spots or stripe, extending from ventral mar-
gin of orbit to posterior tip of maxillary. Body gen-
erally 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 Anisochro-
mis 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 bi lateral counts on 28 specimens.
^Includes bi lateral 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; dis-
tal margins immaculate. Caudal fin greenish in area
covered by scales, rays greenish or orangish, mem-
branes variably dusky. Pelvic fin irregularly dusky
basally, immaculate distally. Pectoral fin pale green-
ish basally, pale dusky distally. Specimens obtained
during collections are remembered as having green-
ish heads and bodies, often with broad orange-
yellow area on side, and white-margined, dark spot
on opercle.
Color pattern of preserved specimens. Terminal
stage. Head relatively immaculate dorsally and ven-
trally; dusky and black markings on side of head
as described for fresh specimens; adjacent black and
pale stripes extending from eye may vary in inten-
sity and distinctiveness, and may be present only as
dark and pale spots; faint dusky stripe extending
anteriorly from orbit at about nine o'clock posi-
tion; 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 mem-
branes of dorsal fin and anterodorsally on body.
Ocellated stage. Dark and dusky markings as de-
scribed for fresh specimens, white areas immacu-
late; faint, dusky stripe extending anteriorly from
orbit at about nine o'clock position. Some speci-
mens, 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 re-
stricted to dorsal portion; abruptly pale, roughly
rectangular area often encompassing much of area
from venter to midside dorsally and from pectoral-
fin axil to level of anterior anal-fin rays posteriorly.
Many degrees of intermediacy in color pattern
between the ocellated and terminal stages are repre-
sented by available specimens; however, the speci-
mens 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 simi-
lar to A. kenyae, the only other species of Aniso-
chromis, 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 posteri-
orly. 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 speci-
mens 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 (Fig-
ure \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 oc-
cupies much of the distal area of the membranes
between the anterior two to four rays.
Frequency distributions and their means for cer-
tain 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 speci-
mens 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 partic-
ipated in all the collections, during which we were
assisted by various other participants in the expe-
dition 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 speci-
mens, 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 speci-
mens, 17.2-25.5 mm SL, VGS 76-8.
ETYMOLOGY.?Named in honor of Lewis H.
Strauss of Washington, D.C., who conceived, orga-
nized, produced, and participated in the expedi-
tion to St. Brandon's that netted a scientifically,
highly valuable collection of fishes and other ma-
rine organisms.
DISTRIBUTION AND HABITAT NOTES.?Anisochro-
mis straussi is known only from the isolated and
relatively tiny area of St. Brandon's Shoals (= Car-
gados Carajos) in the southwestern Indian Ocean.
Specimens were obtained only in rotenone collec-
tions. 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 finger-
like 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 lati-
tude (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. Transfor-
mation occurs when the germinal epithelium ceases
to produce oocytes, and seminiferous crypts prolif-
erate 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 col-
lected (evidenced by oocytic debris in the germinal
ridge), the sperm duct is visible near the union of
the gonads but not farther posteriorly. This speci-
men 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 conspic-
uously 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 exam-
ination of the gonads of 12 specimens of Aniso-
chromis straussi. In general, specimens with ocel-
lated stage color pattern are smaller and are fe-
males, and specimens with terminal stage color
pattern are larger and are males, but there is con-
siderable overlap. The overlap is not surprising
and is comparable to that found in other trans-
forming hermaphrodites (C. L. Smith, 1967). The
presence of a nonfunctional lumen in the testis
(remnants of the ovarian lumen) and oocyte re-
mains among the testicular acini are further evi-
dence that this species is a protogynous hermaphro-
dite.
Relationships of the Anisochromidae
J. L. B. Smith (1954b) described the Anisochro-
midae 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 (Pseudo-
chromidae was first proposed as a family-group
name by Muller and Troschel, 1849, and Pseudo-
plesiopidae by Bleeker, 1875, who included a group
Pseudoplesiopini among the three groups he recog-
nized 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, respec-
tively), 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 Aniso-
chromidae 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 (pseudo-
chromoids, Plesiopidae, Grammidae, Pseudogram-
midae, Acanthoclinidae) Bohlke recognized in this
group have a long and involved classificatory his-
tory. Most have been included in the Serranidae
at one time or another, and all have specialized
character states of the lateral line, including: inter-
rupted, 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 explana-
tion, 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 Sny-
der, 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 pseudo-
chromoids, 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 deter-
mine 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 re-
duced number of dorsal-fin spines and the nature
of the connection between the uncinate process of
the first epibranchial and the second infrapharyn-
gobranchial.
NUMBER 252
Based on the generalized perciform fishes Perci-
chthys and Morone (Percichthyidae), we consider
the presence of 10 dorsal-fin spines to be the un-
specialized 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), Pseudo-
plesiopidae (1 or 2, primitively 2), Pseudochromi-
dae (2 or 3, primitively 3); Group II, Pseudogram-
midae 6 to 8, primitively 8 (note: the anteriormost
spine is vestigial, visible only in osteological prepa-
rations, 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 infrapharyn-
gobranchial 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 inter-
arcual cartilage is a synapomorphy of the perci-
forms, and many perciforms exhibit the cartilage,
including all the fishes Bohlke (1960) discussed
except the pseudochromoids. The lack of an inter-
arcual cartilage in the pseudochromoids represents
either retention of a primitive, beryciform condi-
tion or a specialization, through loss, convergent on
the beryciform condition. Many specialized (usually
benthic) perciform fishes lack an interarcual carti-
lage (for instance: blennioids, stichaeoids, cottoids,
batrachoids, trichonotids, pholydichthyids, calli-
nonymids, trichodontids, uranoscopids; Springer
and Freihofer, 1976). In view of the numerous
specializations shared by the pseudochromoids and
many perciforms with an interarcual cartilage (in-
cluding, 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 beryci-
form 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 com-
mon to a large number of families Bohlke did not
treat (a disjunct lateral line occurs also in the
Nannopercidae, some Blenniidae, and some Brotu-
lidae, 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 specializa-
tions 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 epi-
hyal and ceratohyal not sutured together (but su-
tured 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 demon-
strated, 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 Grammis-
tidae, and Schultz (1966) agreed. C. L. Smith and E. H. Atz
(1969) concluded on the basis of gonodal structure, and Ran-
dall, 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 pseudogram-
mids to form a portion of a subfamily Grammistinae, of the
Serramidae.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ceratohyal sutured together (both surfaces), hy-
purals 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 pos-
sible exception, Lipogramma, which Bohlke (1960)
described and assigned, for lack of another possi-
bility, 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 liga-
ment 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 ossi-
fication, has occurred independently in a wide va-
riety 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 Serra-
nidae (but not others). Considerably more study
will be necessary to determine whether Lipo-
gramma, or any of the other fishes Bohlke dis-
cussed, 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 Pseudochro-
midae. Myers (1935) indicated that although Para-
sphenanthias 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 pseudo-
chromoids. 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 pseudo-
chromoids (or any of the other families Bohlke
treated) in having autogenous hypurals 3 and 4
(these hypurals appear to be fused in some speci-
mens or species of cepolids, but they are primi-
tively 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 cepo-
lids are also less specialized than the pseudochro-
moids in having an interarcual cartilage between
epibranchial 1 and infrapharyngobranchial 2 and an
infrapharyngobranchial 1, and are perhaps less spe-
cialized in lacking the ceratohyal-dentary ligament.
The cepolids are more specialized than the pseudo-
chromoids (and other fishes Bohlke treated) in hav-
ing: the first dorsal-fin pterygiophore inserted be-
tween the first and second neural spines (primitive
condition: between spines 2 and 3), the last dorsal
and anal-fin pterygiophores each supporting a sin-
gle ray (primitive condition: supporting two ele-
ments; 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; pseudochro-
moids have 2 or 3).
The conjunction of more primitive states for the
caudal-fin hypurals, interarcual cartilage, infra-
pharyngobranchial 1, and ceratohyal-dentary liga-
ment, together with the more specialized states for
dorsal-fin pterygiophores, predorsal bones, and post-
cleithra, 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 opistognath-
ids exhibit a number of specializations that are also
NUMBER 252 11
found in the pseudochromoids (lateral line incom-
plete and terminating below the dorsal fin, six
branchiostegals, epihyal and ceratohyal sutured to-
gether, 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 opistognath-
ids also possess an interarcual cartilage, 9-12 dorsal-
fin spines, infrapharyngobranchial 1, no ceratohyal-
dentary ligament, last dorsal and anal-fin pterygio-
phores 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 mono-
phyletic 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 au-
tapomorphies, 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 char-
acters 8 and 10 in the table, but based on more
generalized perciform fishes (particularly Percich-
thys), we propose the following directions for spe-
cialization 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 antero-
dorsally.
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 in-
ferred from condition in Lipogramma and the
Clinidae, and less certainly in Serranus, where the
ligament connects at the symphysis or point ante-
rior to coronoid process).
The Anisochromidae exhibit at least five autopo-
morphies (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 pseudoplesiop-
ids 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 rea-
son to doubt their monophyly. The pseudochro-
mids exhibit one possible synapomorphy (7) with
the pseudoplesiopids but none with the anisochro-
mids, and one might consider this a basis for recog-
nizing 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 (al-
though maintaining them as subfamilies), and Lub-
bock (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 synapomor-
phies, however, the Anisochromidae and Pseudo-
plesiopidae 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 pseudo-
chromoids, and we here propose to combine them
all under the oldest of the three family names,
Pseudochromidae. Concurrently, we do not recog-
nize subfamilies in the Pseudochromidae. In effect,
we have chosen the lowest (least equivocal) phylo-
genetic level for which there is evidence for mo-
nophyly among the pseudochromoids as the level
for family recognition.
Diagnosis of the Pseudochromidae
Small, often colorful, Indo-West Pacific reef-
dwelling 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). Pelvic-
fin 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 be-
tween uncinate process of epibranchial 1 and infra-
pharyngobranchial 2. Infrapharyngobranchial 1
absent. Six branchiostegals. Epihyal and ceratohyal
suturally joined. Preopercle margin entire. Tubed
lateral-line scales in two disjunct series (one dorso-
anteriorly, 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 pre-
sent attaching lateral surface of ceratohyal either
to coronoid process or symphysial area of dentary.
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14
NUMBER 252 15
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