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NOAA Technical Report NMFS Circular 428

Morphological Comparisons
of North American Sea Bass
Larvae (Pisces: Serranidae)

Arthur W. Kendall, Jr.

August 1979

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

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NOAA Technical Report NMFS Circular 428

j«Sv.

^fNT Of C

Morphological Comparisons
of North American Sea Bass
Larvae (Pisces: Serranidae)

Arthur W.Kendall, Jr.

August 1979

■a

■
a

U.S. DEPARTMENT OF COMMERCE

Juanita M. Kreps, Secretary

National Oceanic and Atmospheric Administration

Richard A. Frank, Administrator

National Marine Fisheries Service

Terry L. Leitzell, Assistant Administrator for Fisheries

The National Marine Fisheries Service (NMFS) does not approve, rec-
ommend or endorse any proprietary product or proprietary material
mentioned in this publication. No reference shall be made to NMFS, or
to this publication furnished by NMFS, in any advertising or sales pro-
motion which would indicate or imply that NMFS approves, recommends
or endorses any proprietary product or proprietary material mentioned
herein, or which has as its purpose an intent to cause directly or indirectly
the advertised product to be used or purchased because of this NMFS
publication.

CONTENTS

Page

Introduction 1

Methods and Materials 1

Results 2

Subfamily Serraninae 2

Centropristis 2

Paralabrax 4

Diplectrum 5

Serranus 6

Hypoplectrus 10

Schultzea 12

Serraniculus 12

Serraniculus pumilio 12

Discussion 15

Subfamily Anthinae 16

Plectranthias 16

Anthias 16

Pronotogrammus 20

Hemanthias 22

Hemanthias uivanus 22

Discussion 26

Subfamily Epinephelinae 26

Paranthias 30

Epinephelus 31

Mycteroperca 34

Mycteroperca microlepis 34

Discussion 35

Subfamily Grammistinae 38

Liopropoma 38

Rypticus 39

Pseudogramma 40

Pseudogramma gregoryi 41

Discussion 43

Discussion 45

Summary 45

Acknowledgments 47

Literature Cited 47

Figures

1. Larvae of Centropristis striata from the northwestern Atlantic Ocean 4

2. Spine-bearing bones of the opercular and posttemporal regions on a 10.6-mm Centropristis striata

larva 4

3. Larvae of Paralabrax sp. from the northeastern Pacific Ocean 5

4. Spine-bearing bones of the opercular and posttemporal regions on a 9.5-mm Paralabrax sp. larva ... 5

5. Larvae of Diplectrum sp. Type 1 from the northwestern Atlantic Ocean 6

6. Larvae of Diplectrum sp. Type 1 from the northeastern Pacific Ocean 7

7. Spine-bearing bones of the opercular and posttemporal regions on a 10.3-mm Diplectrum sp. Type 1

larva 8

8. Larvae of Diplectrum sp. Type 2 from the northeastern Pacific Ocean 8

9. Spine-bearing bones of the opercular and posttemporal regions on a 9.9-mm Diplectrum sp. Type 2

larva 8

10. Larvae of Serranus sp. from the northwestern Atlantic Ocean 9

11. Spine-bearing bones of the opercular and posttemporal regions on a 9.7-mm Serranus sp. larva 10

12. Laboratory-reared larvae of Hypoplectrus sp. from eggs collected near Miami, Fla 11

13. Spine-bearing bones of the opercular and posttemporal regions on a 12.5-mm Hypoplectrus sp. larva . 12

14. Young stages of Serraniculus pumilio from the northwestern Atlantic Ocean 13

iii

15. Larvae of Plectranthias garupellus from the northwestern Atlantic Ocean

16. Larvae oiAnthias sp. Type 1 from the northwestern Atlantic Ocean

17. Larvae oiAnthias sp. Type 2 from the northwestern Atlantic Ocean

18. Larva oiAnthias tenuis from the northwestern Atlantic Ocean

19. Larva oiAnthias sp. Type 3 from the northwestern Atlantic Ocean

20. Larvae of Anthias gordensis from the eastern Pacific Ocean

21. Spine-bearing bones of the opercular and posttemporal regions on a 10.2-mm Anthias sp. Type 1
larva

22. Larvae of Pronotogrammus aureorubens from the northwestern Atlantic Ocean

23. Larvae of Pronotogrammus eos from the eastern Pacific Ocean

24. Spine-bearing bones of the opercular and posttemporal regions on a 9.5-mm Pronotogrammus
aureorubens larva

25. Larvae of Hemanthias vivanus from the northwestern Atlantic Ocean

26. Larva of Hemanthias peruanus from the eastern Pacific Ocean

27. Spine-bearing bones of the opercular and posttemporal regions of a 4.3-mm Hemanthias vivanus
larva

28. Spine-bearing bones of the opercular and posttemporal regions on a 10.3-mm Hemanthias vivanus
larva

29. Development of the supraoccipital crest of Hemanthias vivanus

30. Details of the head spination on a Hemanthias vivanus larva

31. Larva of Paranthias furcifer from the eastern Pacific Ocean

32. Larva of a member of the Epinephelus striatus species group from the northwestern Atlantic
Ocean

33. Larva of an Epinephelus sp. (subgenus Dermatolepis or Alphestes) from the eastern Pacific Ocean . .

34. Spine-bearing bones of the opercular and posttemporal regions on a 10.2-mm Epinephelus niveatus
larva

35. Young stages of My cteroperca microlepis from the northwestern Atlantic Ocean

36. Spine-bearing bones of the opercular and posttemporal regions on a 9.8-mm Mycteroperca
microlepis larva

37. Larvae of Liopropoma sp. from the northwestern Atlantic Ocean

38. Spine-bearing bones of the opercular and posttemporal regions of a 12.0-mm Liopropoma
sp. larva

39. Larva of Rypticus sp. from the northwestern Atlantic Ocean

40. Larvae of Pseudogramma gregoryi from the northwestern Atlantic Ocean

41. Spine-bearing bones of the opercular and posttemporal regions on a 10.3-mm Pseudo-
gramma gregoryi larva

42. Larvae representative of four groups of American Serranidae

43. Interoperculars of larvae of representatives of four groups of American Serranidae

Tables

1. Some adult and larval characters of genera of North American serranids

2. Meristic character development of larvae of Serraniculus pumilio

3. Body proportions of larvae of Serraniculus pumilio

4. Meristic character development of larvae of Hemanthias vivanus

5. Body proportions of larvae of Hemanthias vivanus

6. Meristic character development of larvae of Mycteroperca microlepis

7. Body proportions of larvae of Mycteroperca microlepis

8. Meristic character development of larvae of Pseudogramma gregoryi

9. Body proportions of larvae of Pseudogramma gregoryi

17
18
19
20
20
21

22
23

24
25
26

26
28
29
31

32
33

34
36

38
40

41
41
42

44
46
47

14
15
27
28
37
38
43
44

Morphological Comparisons of North American
Sea Bass Larvae (Pisces: Serranidae)

ARTHUR W. KENDALL, JR.

ABSTRACT

Larvae of 17 of the 23 nominal genera of American serranid fishes are described. Representatives
of only two of these genera have been described previously from American waters. The genera fall into
four groups which closely follow subfamilial groupings based on adult characters. Larvae of the
Serraninae, representing seven genera, appear to be the most generalized and are most similar to
Morone-like percichthyid larvae. They have some larval specializations but exhibit rather direct de-
velopment of adult morphology. The serranine genera can be ranked in order of increasing larval spe-
cialization as follows: Serraniculus, Paralabrax, Centropristis, Diplectrum, and Serranus.
Hypoplectrus and one type of Diplectrum, although considered serranines on the basis of adult char-
acters, are quite different from the other serranine larvae observed. One line of divergence from the
serranines is the Anthiinae, with larvae of four genera being represented in the present collections.
These larvae have variously developed strong spines on the head and in the opercular region, also the
pelvic fin spine and some dorsal fin spines are strong and develop precociously. These spines are ser-
rated in more specialized genera. Among the anthiine genera described there is a progression of larval
specialization as follows: Plectranthias, Pronotogrammus, Anthias, and Hemanthias. A third
general type of serranid larvae is represented by members of the three American genera of the Epine-
phelinae and Gonioplectrus. These larvae all are similar in general appearance and specialized in
having elongate, strong serrate spines — primarily the second spine of the dorsal fin and the pelvic
spine. The fourth larval type is comprised of three genera whose affinities have been unclear.
Liopropoma is generally considered by others to be a serranid of the subfamily Liopropominae, Pseu-
dogramma has been placed by others in the family Pseudogrammidae, or considered to be a member of
a subfamily of the Grammistidae, the family in which Rypticus, the third genus, is placed. Because of
the similarity of their larvae and evidence from adult characters, I consider these genera to be
members of the serranid subfamily Grammistinae. Larvae of these genera share some larval charac-
ters with the serranines. Their most outstanding larval feature is the development of one or two great-
ly elongated flexible dorsal fin spines.

INTRODUCTION

Jordan and Eigenmann (1890) reviewed the American
and European serranids known at that time. The limits
of and relationships within the family have continued to
be sources of study and, as presently understood, in
American waters it consists of three of Jordan and Eigen-
mann's (1890) six subfamilies (Serraninae, Anthiinae,
and Epinephelinae) plus the Liopropominae (Gosline
1966). Their Grammistinae has been elevated to familial
status (Gosline 1966).

Serranid larvae are poorly known, although adults of
most species are common and important commercially.
Early descriptions of Mediterranean larvae indicated a
diverse morphology (Fage 1918). The larvae have vari-
ously developed head and fin spination that seem related
to the group within the family to which the species are
assigned. Larvae of only 2 of the 100 or so species of
American serranids have been described — Epinephelus
niveatus by Presley (1970) and Centropristis striata by
Kendall (1972). Descriptions of these and other serranid

'Northeast Fisheries Center Sandy Hook Laboratory, National Marine
Fisheries Service, NOAA, Highlands, N.J.; present address: Northwest
& Alaska Fisheries Center, National Marine Fisheries Service, NOAA,
2725 Montlake Boulevard East, Seattle, WA 98112.

larvae indicate that the family may be subdivided based
on specializations of the larvae. In this paper, sub-
familial and generic characteristics of American ser-
ranid larvae are defined from material from western
Atlantic and eastern Pacific waters and one representa-
tive of each of the four subfamilies is described in detail.

METHODS AND MATERIALS

Since only Hypoplectrus and Centropristis (Hoff
1970) among American serranids have been reared
beyond yolk exhaustion, the indirect method of assem-
bling series of similar appearing larvae from plankton
samples was used.

Representative larvae were drawn using a camera
lucida attachment on a dissecting microscope. Pigment
on the formaldehyde-preserved specimens was limited to
melanophores which is variously referred to as it ap-
peared on the larvae, e. g., as spots, patches, or blotches.
Body proportion measurements were made with an
ocular micrometer. The larval period was divided into
three stages: preflexion (before notochord flexion), flex-

2W. J. Richards, National Marine Fisheries Service, Southeast Fish-
eries Center, Miami, pers. commun. January 1974.

ion (during notochord bending), and postflexion (after
the notochord tip had completed flexion) following rec-
ommendations of Ahlstrom et al. (1976). Body length
(BL), the reference for proportion measurements, was
measured from the tip of the snout to the tip of the noto-
chord (notochord length (NL) in preflexion and flexion
larvae and in postflexion larvae it was measured from the
tip of the snout to the margin of the hypural bones (stan-
dard length (SL) as in Ahlstrom et al. (1976). Some
specimens were cleared and stained following Hollister
(1934) and Clothier (1950). The ventral portion of the
interopercular bone is not shown in the figures of the
opercular area because its outline was hidden by over-
lying tissue in the cleared and stained specimens.

Specimens included materials from collections at the
National Marine Fisheries Service (NMFS) Labora-
tories in La Jolla, Calif.; Miami, Fla.; Narragansett,
R.I.; and Sandy Hook, N.J.

Most of the material was collected as part of one of
three large-scale ichthyoplankton surveys — one in the
Atlantic Ocean and two in the Pacific Ocean. The survey
in the Atlantic was that of NMFS, Sandy Hook, aboard
the RV Dolphin in 1965-68 when ichthyoplankton was
sampled from Cape Cod, Mass., to Palm Beach, Fla.
Most of the Pacific material resulted from the CalCOFI
program, a long-continuing survey of waters off Cali-
fornia and Baja California. The other major source of
material from the Pacific Ocean was the EASTROPAC
program, which consisted of three multivessel cruises in
the eastern tropical Pacific Ocean. Appendix Table 1
lists collection data associated with specimens used for
the illustrations.

RESULTS

Subfamily Serraninae

The serranines have rather uniform characteristics
which help distinguish them from other serranids (Table
1). They are represented by seven genera in North Amer-
ican waters with 52 species and are moderate to small-
sized serranids apparently representing the base from
which other subfamilies arose (Smith and Atz 1969; Ken-
dall 1976). Most serranines are synchronous hermaphro-
dites; however, protogyny and secondary gonochorism
are present in some species (Smith and Young 1966) .
The lateral line is complete and not highly arched and
the scales are moderate to large and strongly ctenoid
(Jordan and Eigenmann 1890). The larvae of three Euro-
pean species of Serranus (Fage 1918; Bertolini 1933;
Aboussouan 1972) as well as those of Centropristis striata
(Kendall 1972) are the only serranines described.

'Serranids studied to date are hermaphroditic (Smith 1965; Kendall
1977), i.e., male and female gametes are produced in each individual.
Some genera are secondary gonochorists, i.e., the gonads show
histological traces of both sexes but only male or female gametes are pro-
duced in each individual. Hermaphroditism takes on two forms in serran-
ids: synchronous hermaphroditism, where both types of gametes are
produced simultaneously in one individual, and protogyny, where in-
dividuals mature first as females and later in life the gonad transforms
into testes and produces sperm.

Larvae of all but one (Schultzea) of the North Ameri-
can genera of serranines are herein described. Serranine
larvae, except those of Hypoplectrus (see below), can be
distinguished from larvae of other fishes on the basis of
several characters. Development in serranines is direct
and they do not possess many of the larval specializa-
tions of other serranids. They do not have elongate pre-
opercular spines, rather a series of blunt points. The
dorsal and pelvic fin spines are thin and only slightly
elongate in some genera. Larval pigment generally con-
sists of melanophores in characteristic positions mostly
along the ventral midline. Generally, there are spots at
the angle of the jaw, at the junction of the cleithra, be-
tween the bases of the pelvic fins, near the anus, at the
bases of the anal rays and some on the caudal peduncle.
Some of these spots are intensified in some species. Pig-
ment on other parts of the body is variable within the
subfamily but some is consistent within genera. The
body shape of the larva gradually changes to that of the
adult, without abrupt changes in proportions.

Centropristis. — Centropristis occurs only in the
northwest Atlantic where four species are recognized
(Kendall4). These are moderately large protogynous her-
maphrodites. Centropristis striata occurs farther north
than any other serranid on the east coast. A closely re-
lated species, C. melana, is widespread in the Gulf of
Mexico."' The two other species, C. philadelphica and C.
ocyurus, cooccur with C. striata in the South Atlantic
Bight. The species of Centropristis are distinguished by
meristic and other characters (Table 1).

The eggs of C. striata were the object of one of the early
detailed studies of fish embryology (Wilson 1891) and
Hoff (1970) illustrated eggs and prolarvae of C. melana.
Larvae of C. striata were not described until recently
(Kendall 1972), although Sette (see Merriman and Sclar
1952) and Pearson (1941) had apparently recognized
them in plankton samples.

I found only one type of larva assignable to
Centropristis in the present material. The description of
Kendall (1972) has been condensed and supplemented
with recent observations in the following account.

Larval morphology of Centropristis (Fig. 1) is typical
of other serranines. There is no armature development
associated with the fin spines, the preopercular bone has
only small, unserrate protruding spines (Fig. 2) and body
shape, once the caudal fin has formed, approximates
that of small adults. Fin spines are thin and not pro-
duced. Dorsal fin spines develop at about the same stage
as the soft rays. Pelvic fins develop after the other fins
have several rays ossified and grow isometrically
throughout development.

In early larvae (<5 mm) there is some dorsal body pig-
ment but afterwards nearly all larval pigment is asso-

JA. W. Kendall. 1977. Biological and fisheries data on black sea
bass. Centropristis striata (Linnaeus). Sandy Hook Lab. Tech. Ser.
Rep. No. 7, 29 p.

''Some (e.g., Miller 1959) consider C. melana a subspecies of C. striata.

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Figure 1.— Larvae of Centropristis striata from the northwestern Atlantic Ocean: a) 4.7 mm; b) 8.3 mm.

posttemporal

preopercular

supracleithrum

Figure 2. — Spine-bearing bones of the opercular and posttemporal
regions on a I0.(i-mm Centropristis striata larva.

ciated with the ventral midline. Melanophores (spots) in
characteristic positions are larger than others. Large
spots are seen on the posterior margin of the angular, at
the junction of the cleithra, between the bases of the
pelvic fins, near the anus, and near the insertion of the
anal fin. Several slightly smaller spots develop on the
caudal peduncle, and still smaller spots are seen at the
bases of the anal fin rays, between the larger caudal pe-
duncle spots, and at the bases of some caudal fin rays.
Several melanophores also occur on the posterior portion

of the optic lobes and on the dorsoposterior lining of the
coelomic cavity.

Paralabrax. — Paralabrax with eight species is the
only North American serranine limited to the eastern Pa-
cific (Table 1). It is ecologically similar to Centropristis
in the Atlantic in that it is larger than most other serran-
ines and occurs at higher latitudes than other serranines
on the Pacific coast. Smith and Young (1966) indicated
that Centropristis and Paralabrax are also closely re-
lated. The only species that has been studied is a sec-
ondary gonochorist, while most other serranines are syn-
chronous hermaphrodites (Smith and Young 1966).

Paralabrax larval development is typical of serran-
ines. Paralabrax larvae approach adult body pro-
portions soon after tail flexion is complete (Fig. 3). The
spinous and soft portions of the dorsal fin form at about
the same rate, as they do in Centropristis. The fins of
Paralabrax larvae are more heavily pigmented than
those of Centropristis but the ventral melanophores are
smaller and more uniform in size. Pelvic fins form with
pigment on the membrane as the dorsal fin starts to
form. Pigment develops ventrally at the base of the
pelvic fin, near the anus, along the base and at the inser-
tion of the anal fin, and on the caudal peduncle. Pigment
also forms on the angular, on the dorsal surface of the
brain, and at the base of the caudal fin. Some pigment
develops dorsally at the base of the spinous dorsal fin and
a large blotch of pigment develops on its membrane. The
pectoral fin rays are pigmented on their distal third and
the pelvic fin is heavily pigmented on and between the
rays. Opercular spination is as in other serranines (Fig.
4): the preopercular has about three spines dorsal and

Figure 3. — Larvae of Paralabrax sp. from the northeastern Pacific Ocean: a) 6.(1 mm; b) 9.0 mm.

two ventral to the one at the angle, on the preopercular
ridge there are a few blunt spines, and the subopercular
has a smooth posterior margin.

Diplectrum. — In contrast to reef-dwelling habits of
many other serranids, Diplectrum occurs mainly over
sand or mud bottoms on the continental shelves (Bortone
1977). These moderate-sized synchronous hermaphro-
dites have one or two clusters of strong and divergent
spines on the preopercular which is produced into a bony
flap. The Pacific species were reviewed by Rosenblatt
and Johnson (1974) and the entire genus by Bortone
(1977).

Diplectrum is the only serranine with larvae from both
Atlantic and Pacific waters. Since species of Diplectrum
have overlapping meristic features and geographic
ranges, the larvae cannot be assigned to species. Two dis-
tinct larval types with the unique meristic characters of
Diplectrum were found from both oceans. They will be
referred to as Type 1 and Type 2 and discussed sepa-
rately, since they share few larval characters in common.

Type 1 — Larvae of Diplectrum called Type 1 from the
Atlantic and Pacific Oceans are similar in development.
They exhibit most of the features of other serranine lar-
vae. The body shape approaches that of the adult and in
later larval stages, after the fin rays form, they are among
the most slender serranines (Figs. 5, 6). Fin develop-
ment is modified from the serranine pattern (see page 2)
in that the first dorsal fin spines and pelvic fin elements

posttemporal

supracleithrum

preopercular

bopercular

interopercular

Figure 4. — Spine-bearing bones of the opercular and posttemporal
regions on a 9.5-mm Paralobrax sp. larva.

form early. None of the early forming dorsal fin spines
are very elongate, as they are in some Serranus larvae.
Pigment on larvae from both oceans is present along the
ventral midline in positions characteristic of serranine
larvae (Table 1). Individual melanophores are more uni-
form in size than those of Centropristis. A series of about

Figure 5. — Larvae of Diplectrum sp. Type 1 from the northwestern Atlantic Ocean: a) 5.8 mm; b) 10.0 mm.

five spots occur ventrally on the caudal peduncle of
Atlantic larvae while there is only a spot at the insertion
of the anal fin and one two-thirds of the way back on the
caudal peduncle in Pacific larvae. At least one spot is
present at the base of the caudal fin on larvae from both
oceans. The membranes of the spinous dorsal and pelvic
fins of Atlantic larvae are variably pigmented with scat-
tered spots. Pacific larvae have intense pigment on the
membrane of the spinous dorsal fin centered around the
fourth spine. The pectoral and pelvic fins of Pacific lar-
vae have pigment on and between the fin rays. The pre-
opercular and subopercular have more armature than is
seen in other serranine genera (Fig. 7).

Type 2 — Larvae assigned here to Diplectrum Type 2
are distinct from Diplectrum Type 1. I have not found
such intrageneric differences in larvae of any other ser-
ranid genus. This would indicate that either these larvae
are not of Diplectrum but are of an undescribed genus,
possibly even in a closely related family with representa-
tives in both oceans, or that there are two quite distinct
types of Diplectrum, possibly worthy of subgeneric rank.

While Type 1 larvae are similar to other serranine lar-
vae, Type 2 larvae show specializations not seen in other
genera. The general body shape is similar to that of Type
1 but the snout is more elongate and is concave in dorsal
profile in smaller specimens (Fig. 8). The sequence and
extent of fin ray formation is different from that in other
serranines. The pectoral fin develops early, before most
other fins, as a fan-shaped structure and extends to the
anus. Only in grammistines is similar larval pectoral fin
development seen. The pelvic fins form before the rest of

the fins and are elongate, reaching beyond the anus. The
distal third of the rays of the paired fins are covered with
contracted melanophores. Elements of the dorsal, anal,
and caudal fins all develop at about the same time. Pig-
ment pattern differs from that seen in other serranines in
that the body anterior to the anus is unpigmented before
the fin rays are complete and later in development a
large diffuse internal pigmented area forms between the
lateral line and the anal fin, dorsal to the anterior half of
that fin. The bones of the opercular region are similar to
those of Diplectrum Type 1 except the subopercular
margin is smooth and does not have the small spines seen
in Diplectrum Type 1 (Fig. 9).

Serranus. — In the western Atlantic there are 13
species and in the eastern Pacific there are 5 nominal
species assigned to Serranus (Table 1). These are rather
small fishes ( <200 mm) sharing 17 characters in common
which, taken together, exclude all other genera (Robins
and Stark 1961). Their median fin ray counts fall out-
side the ranges of other American serranids except
Schultzea beta, which has only six branchiostegal rays
(Robins and Stark 1961) and Diplectrum, which general-
ly has more pectoral rays (Table 1).

Robins and Stark (1961) examined east coast Serranus
species in some detail. Since Serranus in the eastern Pa-
cific is poorly known with few specimens of some species
taken, some species may prove invalid.

While no American Serranus larvae have been de-
scribed, eggs and early larvae of Mediterranean species
of Serranus were among the first serranids described

Figure 6. — Larvae of Diplectrum sp. Type 1 from the northeastern Pacific Ocean: a) 4.3 mm; b) 5.0 mm; c) 6.1 mm; d) 13.0 mm.

posftemporal

opercular

preopercular

supracleithrum

Figure 7. — Spine-bearing bones of the opercular and posttemporal
regions on a 10.3-mm Diplectrum sp. Type 1 larva.

opercular

preopercular

supracleithrum

Figure 9.— Spine-bearing bones of the opercular and posttemporal
regions on a 9.9-mm Diplectrum sp. Type 2 larva.

Figure H. — Larvae of Diplectrum sp. Type 2 from the northeastern Pacific Ocean: a) 4.5 mm; b) 8.4 mm.

(Raffaele 1888). Larger larvae of Mediterranean species
have since been illustrated and described (Bertolini
1933) and there are brief descriptions of Serranus larvae
collected off Africa (Aboussouan 1972).

Serranus larvae were found in samples from the Atlan-
tic; however, none were found in Pacific collections.

Although there are differences among Serranus larvae in
extent of fin ray elongation, pigment, and body shape, all
found so far are separable from other larvae by the be-
ginning of the flexion stage. More detailed study may al-
low descriptions of the several types and their assign-
ment to species

Figure 10. — Larvae of Serranus sp. from the northwestern Atlantic Ocean: a) 3.7 mm; b) 5.0 mm; c) 5.5 mm; d) 9.4 mm.

Serranus development shows several modifications of
the basic serranine plan (page 2). The body is consider-
ably deeper from the head back to the anus from before
notochord flexion until after all the fin rays have formed
(Fig. 10). The pelvic fin forms before the other fins and
the elements are elongate in some larvae. The first three
to five spines of the dorsal fin form before the rest of the
fin and in some the third spine is elongate. Among Ser-
ranus larvae described so far these fin spines are longest
in 5. cabrilla (Fage 1918).

In Serranus larvae some of the characteristic ventral
pigment is intensified and some is eliminated. The more
contracted melanophores of other serranines are absent
in Serranus. Intensified melanophores are on the
angular, at the junction of the cleithra, and near the
anus. A spot at the insertion of the anal fin and one
midway back on the caudal peduncle are the largest
spots ventrally and are more pronounced in some larvae
than in others. Dorsally, on at least some larvae, there is
a spot near the base of the last dorsal spine. Another spot
ventral to the posterior third of the second dorsal fin is
variable in position from near the base of the fin to near
the lateral line. This spot opposes the one at the insertion
of the anal fin and in S. cabrilla these two spots are large
and distinctive (Fage 1918). Pigment spots are scattered
on the membranes of the spinous dorsal, anal, and pec-
toral fins in some larger larvae.

The opercular region is armed with blunt spines, which
are generally more pronounced than those of other ser-
ranines (Fig. 11). There seems to be some variation in
spine length among the species of Serranus since S.
cabrilla has longer spines (Fage 1918) than the larvae I
examined. The preopercular has two spines ventral to
and three spines dorsal to the one at the angle. There are
spines anterior to these on the ridge of the preopercular
bone in some specimens. The subopercular has two
spines dorsal to the one seen at its angle in other serran-

posttemporal

supracleithrum

preopercular

bopercular

interopercuiar

Figure II. — Spine-bearing bones of the opercular and posttemporal
regions on a 9.7-mm Serranus sp. larva.

ines. Among larval serranines, only Diplectrum has a
more ornately armed opercular region.

Hypoplectrus. — This genus of brightly colored small
reef fishes has long perplexed scientists. In the western
Atlantic there are several nominal species with different
color patterns but with identical meristic features and
body proportions. At present, these are generally consid-
ered valid species; however, other species of serranids
such as Mycteroperca rosacea and M. olfax have more
than one color phase (Rosenblatt and Zahuranec 1967).
Barlow (1975) observed spawning between two nominal
species and so considered them synonymous. Thresher
(1978) concluded that the color patterns observed in
Hypoplectrus demonstrated mimicry of nonpredatory
reef fishes. Apparently, coloration in Hypoplectrus is a
plastic character; and with the possibility of clones in a
synchronous hermaphrodite, different color patterns can
become established locally in a relatively short time. The
poorly known eastern Pacific Serranus lamprurus Jordan
and Gilbert was placed in Hypoplectrus by Meek and
Hildebrand (1925), but Robins and Stark (1961) stated it
is indeed a Serranus and is close to S. flaviventrus of the
Atlantic, although they gave no reason for their opinion.
Hypoplectrus lamprurus is deeper bodied than Serranus
and has a dorsal fin ray count of X, 14, whereas Serranus
species generally have X, 12 (Table 1). Although a de-
tailed study is needed, it seems that H. lamprurus should
still be considered a Hypoplectrus.

No larvae assignable to Hypoplectrus were found in
plankton collections. Their unique dorsal fin ray count
(X, 14) among east coast serranines assured that if larvae
had been present they would have been recognized. It is
possible that Hypoplectrus larvae are planktonic for only
a short time before they become demersal.

A series of Hypoplectrus sp. was reared from wild
planktonic eggs at the NMFS Laboratory in Miami
(Richards see footnote 2). I was able to examine the
specimens briefly and copy drawings of them. Also, three
cleared and stained specimens were examined in detail.

These specimens are different from other serranine
larvae. It is unlikely that rearing produced these differ-
ences and it is difficult to consider them as modifi-
cations of serranine larval characters. These Hypo-
plectrus larvae are not similar to any known serranid
larva and provide reason to doubt that Hypoplectrus be-
longs in the Serranidae.

The body shape indicates rather direct development
from larval to adult proportions (Fig. 12). The first few
spines of the dorsal fin develop early, during notochord
flexure, but the associated membrane is more fleshy than
in serranines. The pelvic fins form at about the same
stage as the first dorsal spines but the rays are shorter
than those of serranines with early developing pelvic fins.
The membranes of these early forming fins are
pigmented. The rest of the dorsal spines are pro-
gressively shorter than the third and the posterior few
form late after the rayed portion of the fin is developed.
Some of the rays of the second dorsal fin become longer

Figure 12. — Laboratory-reared larvae of Hypoplectrus sp. from eggs collected near Miami, Fla.: a) 3.4 mm; b) 4.7 mm; c) 5.7 mm; d)

8.5 mm.

than the longest spines. The rest of the fins form over a
short length interval of 4.7-5.7 mm.

The head and particularly tissues around the mouth
are fleshier than in serranines. The opercular region has
spines developed on several bones; however, they do not
appear on the larval drawings since they are covered by
flesh. The preopercular is edged with about 12 subequal
spines (Fig. 13). The subopercular has about three blunt
spines on its margin. Besides the three definitive spines
on the opercular, there are about four spines ventral to
them, a condition not seen in serranines.

Pigmentation consists of irregular shaped and sized
melanophores in characteristic positions. More pigment
is present on Hypoplectrus than on most other serran-
ines. A series of spots along the ventral midline consists
of a spot near the base of the pelvic fins, one below the
middle of the gut, one just anterior to the anus, one near
the insertion of the anal fin, and another on the caudal
peduncle. Spots ventral near the tip of the notochord
come to lie at the base of the caudal fin rays as they form.
A few spots near the base of the dorsal fin migrate to form
the dorsal fin membrane pigment. A few spots are scat-
tered on the dorsal surface of the brain posterior to the
eyes. All the fin membranes have pigment in later larval
stages. By 8.5 mm, small melanophores become scat-
tered over the dorsal portion of the anterior two-thirds of
the fish.

opercular

preoperc

supracleithrum

interopercular

Figure 13. — Spine-bearing bones of the opercular and posttemporal
regions on a 12.5-mm Hypoplectrus sp. larva.

Schultzea. — Schultzea is monotypic represented by S.
beta of the western Atlantic (Table 1). It lives more pe-
lagically than other serranines and is adapted to feed on
small, free-swimming prey. The mouth is modified to be
highly protrusible through extensive changes in the
shape of the premaxillary (Robins and Stark 1961). The
fin counts are similar to those of Serranus but there are
only six branchiostegal rays instead of seven (Robins and
Stark 1961).

Two juvenile specimens (24.0 and 25.4 mm) collected
by John McCosker (Steinhart Aquarium, San Francisco,
CA 94118) provided for study did not retain any larval
characters, so they cannot be compared with other
serranines and their relationships within the group can-
not be determined. However, from adult adaptations, it
appears that Schultzea is the most highly modified
American serranine.

Serraniculus. — Serraniculus contains one diminu-
tive species (<80 mm), S. pumilio, in the northwest
Atlantic (Table 1). Its life history has been studied and it
is a synchronous hermaphrodite (Hastings 1973).

A serranine larval type from the Atlantic had meristic
characteristics of S. pumilio, notably six branchiostegal
rays as opposed to seven for all other serranines except
Dules auriga and Schultzea beta, which occur south of
the area where these larvae were collected.

Serraniculus pumilio (Fig. 14). — Meristic element
development (Table 2) — A total of 41 specimens from 3.1
to 7.3 mm was studied to trace meristic character de-
velopment. Since the smallest larvae have a straight no-
tochord and vertebrae are the only ossified meristic ele-
ments, and the largest have adult complements of all
meristic characters except gill rakers and scales, the pat-
tern of meristic character development can be seen in
this series. Vertebrae ossify from anterior to posterior ex-
cept for the urostyle which ossifies before the two or three
vertebrae anterior to it. The smallest larva examined (3.1
mm) had 1 ossified vertebra. Numbers of vertebrae
gradually increase until by 4.6 mm some larvae have all
24 ossified. However, some larvae as large as 6.2 mm
have only 23 vertebrae. Throughout the series, there is
considerable variation in the number of ossified verte-
brae at a given length. Branchiostegal rays are ossified in
some larvae as small as 3.8 mm and the adult comple-
ment of six is seen in most specimens by 4.0 mm, when
the notochord has completed flexion. Other meristic ele-
ments ossify when the notochord completes flexion at 4.0
mm and larvae as small as 4.6 mm have adult fin ray
complements. Pelvic and spinous dorsal fin rays do not
develop precociously as they do in other serranid sub-
families and even in Diplectrum and Serranus. The dor-
sal fin spines and rays each develop at about the same
rate in S. pumilio.

Body proportions (Table 3) — Several body propor-
tions of 21 larvae from 2.9 to 8.3 mm were measured and
expressed as a percentage of body length (BL) to docu-
ment changes in body shape during larval development.
Total length increases rapidly from about 105% of BL at
2.9-3.9 mm, while the notochord is straight, to 115% of
BL at 4.0 mm after notochord flexion is complete; there-
after, it increases gradually to about 125% of BL as the
caudal fin forms. Preanal length increases from about
60% of BL in larvae <4.0 mm to about 65% of BL in 7.0-
to 8.0-mm larvae. Head length remains fairly constant at
about 39% of BL throughout larval development. Eye
length and snout length decrease slightly during de-
velopment from about 12% to slightly less than 10% of

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Figure 14. — Young stages of Serraniculus pumilio from the northwestern Atlantic Ocean: a) 3.8 mm; b) 5.8 mm; c) 55.3 mm.

BL. The greatest body depth decreases from 32% to
about 28% of BL. Depth at anus increases during early
development up to 4.0 mm and then holds constant at
about 25% of BL. Caudal peduncle depth increases from
<6% in BL in the 2.9-mm larvae to about 14% of BL in
4.4-mm larvae; thereafter, it changes little. A compari-
son of the proportions of the larvae after notochord flex-
ion with those of the adult shows that the adult body
shape of S. pumilio is established early in development,
in larvae as small as 4.2 mm.

Pigmentation — Throughout larval development, S.
pumilio is more heavily pigmented than any other

serranine. The characteristic serranine ventral spots are
present, but they are supplemented by many other spots.
Pigmentation is composed of small, individual melano-
phores making up a pattern, rather than the pattern be-
ing comprised of large melanophores in characteristic
positions as seen in other serranids.

The beginning of the pattern is already established in
2.6-mm larvae with three rows of melanophores on the
trunk: one along the dorsal midline, one along the lateral
line, and the third along the ventral midline. Dorsally,
there is a series of about four spots, which are larger than
those in the other two rows. The anteriormost of these

Table 2. — Meristic character development of larvae of Serraniculus pumilio. Specimens
between dashed lines are undergoing notochord flexion. For the caudal fin,"P" and "S"
indicate primary and secondary fin rays.

Body

Branchio-
stegal
rays

Verte-
brae

Pec-
toral
fin

Pel-
vic
fin

Caudal fin

Anal
fin

Dorsal fin
Spines Rays

length
(mm)

Dorsal Ventral
S P P S

Gill
rakers

3.1 NL

3.4

3.5

3.6

3.7

3.8

:■; S

3.8

4.0

4.3

4.4

1
3

3.8
4.1
4.2
4.3
3.9

17
17
17

1
12

1,2

3.8 SL

4.4

4.6

4.7

5.0

5.2

5.7

5.8

6.0

6.1

6.2

7.0

7.2

7.3

22
24
22
24
24
24
21
24
24
23
24
24
24
24
24

1,3

[,5

1,5

1.4

1,5

1,4

1,5

H,7

VIII

111,7

4
2

111,7

II. 1

11,1

VIII

111,7

11,1

VII

11,1

VII

III

111,7

IH,7

111,7

spots is just anterior to the origin of the dorsal fin and re-
mains pronounced throughout development. Addi-
tionally, a series of smaller spots forms along each side of
the dorsal fin. When all of the dorsal rays are formed, one
or two of the original large spots can be seen internal to
the base of the fin and the smaller spots have developed
into a series of nearly contiguous dashes on each side of
the fin base.

The lateral line series begins on 2.6-mm larvae as a
series of dashes from the head about two-thirds of the
length of the trunk. These dashes seem to develop in as-
sociation with the vertebrae and lie internally along the
lateral septa. The spots above the gut become internal
while those farther posterior remain on the surface. From
the anus back, as the original dashes become internal,
another series develops distal to them on the body sur-
face. Superficial spots develop from the anus to the ter-
mination of the dorsal fin between the dorsal and lateral

series of dashes. In larger larvae similar spots also extend
ventrally to a lesser extent from the lateral dashes. These
superficial spots are small but variable in size and posi-
tion and appear as a single pigmented blotch. Superfi-
cial pigment also develops as a line on the head passing
from the snout posteriorly at the level of the eye and
another line on the opercular region in line with the
lateral series of dashes on the trunk. A patch of small
spots also develops laterally in the area of the pectoral
fin. Ventrally, spots extend the length of the larva. A few
spots are present on the tip of the lower jaw, along the
isthmus, on the angular, and at the junction of the clei-
thra. A larger spot is present on the ventral midline be-
tween the junction of the cleithra and the base of the
pelvic fins. Along the gut the ventral spots diverge slight-
ly to form a series on each side of the midline. A few spots
are also present on the midline ventral to the posterior
third of the gut. The anteriormost of these is the largest

Table .'!. — Body proportions of larvae of Serraniculus pumilio. Specimens
between dashed lines are undergoing notochord flexion.

Percent of standard length

Body

Pre-

Caudal

Depth

length

Total

Body

Eye

Head

Snout anal

peduncle

(mm)

length

depth

length

length length

depth

anus

2.9 NL

103

13.0

10.0 58

5.7

3.7

104

11.0

12.0 59

8.0

3.9

107

12.0

10.0 63

10.0

4.0

107

11.0

11.0 61

9.8

4.0 SL

115

11.0

12.0 61

11.0

4.2

118

12.0

10.0 63

13.0

4.2

115

12.0

11.0 62

12.0

4.4

119

11.0

10.0 63

14.0

5.5

117

10.0

10.0 66

14.0

5.7

122

11.0

11.0 59

13.0

6.0

123

11.0

11.0 62

15.0

6.1

123

10.0

9.3 65

14.0

6.7

117

9.4

9.4 62

14.0

6.8

120

11.0

9.0 67

14.0

J 7

7.0

120

9.8

9.8 65

14.0

7.7

125

9.8

7.4 61

13.0

7.8

124

10.0

10.0 64

15.0

7.9

117

11.0

9.5 64

14.0

7.9

124

10.0

9.5 62

13.0

8.3

125

9.9

9.9 66

14.0

Adult1

27-31

8.5-10

34.5-36

8.5-10.5

12-13.5

'Gins

burg (1952

and is ventral to the deepest point of the gut. Posterior to
the gut, a series of spots extends along the ventral mid-
line to the caudal peduncle area in small larvae. As the
anal fin develops, these spots are positioned on either
side of its base. A series of spots also develops at the base
of the anal fin rays and extends posteriorly onto the
caudal peduncle. A spot develops ventrally near the tip
of the notochord. When the caudal fin develops, this spot
is situated ventrally at its base and several other spots
develop along the base of the fin. The posterior third of
the caudal peduncle is unpigmented throughout de-
velopment. Other small spots present during develop-
ment are on the membranes associated with the bran-
chiostegal rays, pectoral and pelvic fins, and internally
on the dorsolateral surface of the gut cavity.

Late in larval development the superficial spots inten-
sify and coalesce giving the larva, particularly the trunk,
a gray appearance and making S. pumilio the most
heavily pigmented serranid larva found to date.

Discussion. — Of the seven genera of North American
serranines, larvae of all but Schultzea have been ex-
amined. Larval material for five of these genera has been
gleaned from plankton collections, and their develop-
ment shows a similar pattern. Laboratory-reared mate-
rial of the other genus, Hypoplectrus, showed a pattern of
development dissimilar to that of other serranines. This
suggests that its adult characters, upon which is pres-
ently understood affinities are based, need to be re-
examined.

Two distinct types of larvae were found that had me-
ristic characters of Diplectrum. Both types of larvae oc-

curred in both oceans and more than two species of Di-
plectrum are known from each ocean. Diplectrum Type 2
larvae are not similar to those of other known serranines.
They do not have the characteristic serranine ventral
pigment, but have a large internal blotch dorsal to the
anal fin. The fin rays of the pectoral, pelvic, and caudal
fins are pigmented with small melanophores. The fin
rays are thin, but longer than in other serranines. On the
basis of larval characters, the affinities of Diplectrum
Type 2 cannot be determined, but it does not appear
closely related to any other serranine.

Among the other serranine genera, the extent of modi-
fications of the basic pattern of development may indi-
cate relationships. Among American serranids, the
serranine genera Serraniculus, Centropristis, and Para-
labrax show larval characters most similar to those of
Morone-type percichthyids (Mansueti 1958, 1964). In
these serranines the dorsal spines develop at the same
stage of development as the dorsal soft rays and are not
elongate. The pelvic fin rays are not precocious or elon-
gate during larval development. Spines of the opercular
region are less developed in these genera than in other
serranine genera. Pigmentation of Serraniculus larvae is
heavier than in other serranines and therefore, most
closely resembles that of Morone-like percichthyids.
Within the serranines it seems that there has been a
trend from a uniformly heavily pigmented larva to one in
which there are a few large spots in characteristic posi-
tions primarily along the ventral midline. Diplectrum
Type 1 and Serranus show modifications of the basic
serranine pattern in that the spinous dorsal fin develops
before the rest of the fin, and in some species of Serranus

some of the dorsal spines are elongate and tipped with
pigmented "flags." The pelvic fins are also early de-
veloping and become elongate, more so in Serranus than
in Diplectrum Type 1. Serranus is deeper bodied than
other serranines and Morone-type percichthyids. Also
some large pigment spots develop dorsally in Serranus.
Thus on the basis of larval characters, there seems to be a
progression of specialization in the order: Serraniculus,
Centropristis-Paralabrax, Diplectrum Type 1, Serranus.

Subfamily Anthiinae

This subfamily is presently under study for revision by
Anderson and Heemstra. This is a cohesive group of
small, brightly colored fishes that occur over hard bot-
toms near continental slopes. They are distinguished by
several morphological modifications from serranine char-
acteristics (Table 1). They have large scales (29-60
lateral line scales), an arched lateral line, deep bodies
with rather large heads, and at least 14 gill rakers, but
mostly more than 25 (Jordan and Eigenmann 1890,
Heemstra see footnote 6). There are about 15 species of
anthiines in American waters grouped in seven genera.
Four genera have representatives in both oceans but no
species are common to both oceans.

Ten types of larvae assignable to four genera {Anthias,
Pronotogrammus, Plectranthias, and Hemanthias) are
present in the material I examined. No American
anthiine larvae have been described previously although
larvae of two species of Anthias from the Mediterranean
Sea have been described (Bertolini 1933; Sparta 1932)
and several anthiines from the southeast Pacific have
been illustrated (Fourmanoir 1976).

Anthiine larvae have deep bodies and stout, produced
preopercular spines. They also have strong spines in their
dorsal, anal, and pelvic fins; but the spines are not as
produced as they are in epinephelines. There seems to be
a trend in anthiines toward development of armature in
the form of serrations on the spinous fin rays and on
spines of various head bones such as the opercular series,
frontals, parietals, and supraoccipitals. In some species
spiny scales develop at a small size, while the fish are
still planktonic. Pigment consists of a few blotches in
characteristic places on the trunk and head.

Plectranthias. — This genus is represented in the
western Atlantic by one species (Robins and Stark 1961).
The genus contains another species found in the Celebes
and Arafura Seas. Plectranthias garupellus, the Atlantic
species, is distinguished from other anthiines in the west-
ern Atlantic by the high number of dorsal soft rays (16)
and the low number of pectoral rays (13) (Table 1). The
spinous dorsal fin is quite angular with the third spine
being the longest. There are two strong antrose spines on
the lower limb of the preopercular. It is known from
Florida and the Bahamas, but is apparently not readily
available to normal collecting procedures.

6P. C. Heemstra, CSIRO, Cronulla, N.S.W., Australia, pers. commun.
April 1973.

Only a few P. garupellus larvae were found, so a de-
tailed description is not possible. However, the larval
characters indicate relationships of the species within
the subfamily and will be discussed briefly. Plectran-
thias garupellus shows little development of character-
istic anthiine larval features (Fig. 15). None of the fin
spines are serrate. The third dorsal spine in small larvae
is elongate but not stout. The head has spines in areas
characteristic of other anthiine larvae but they are weak-
ly developed and not serrate. A simple spine protrudes
from the frontal ridge above the eye. Too few specimens
were available for staining and illustrating the opercular
series but the preopercular has two ridges of blunt spines
with an elongate, thin spine at the angle of the posterior
ridge. The interopercular has an elongate thin spine lying
proximal to the long preopercular spine and the other
opercular bones have some spines protruding. The post-
temporal and supracleithral spines protrude bluntly.
There is no protruding supraoccipital crest.

The pigment pattern of larval P. garupellus is dis-
tinctive. There are three characteristic blotches of pig-
ment on the trunk. Two blotches are dorsal, one below
the sixth and seventh dorsal fin spines, and the other
below the four posteriormost dorsal soft rays. The third
blotch is on the ventral midline just posterior to the in-
sertion of the anal fin. Other spots occur on some speci-
mens, such as one ventrally on the caudal peduncle.

Anthias. — Anthias is circumtropical with several
species in the western Atlantic and one in the eastern Pa-
cific Ocean (Table 3). They are rather small, brightly
colored, deep-bodied fishes. Anthias anthias larvae have
been described (Fage 1918; Roule and Angel 1930; Ber-
tolini 1933) and they closely resemble Anthias larvae I
have examined. I found four types of anthiine larvae in
western Atlantic material and one type in Pacific
material, which were quite similar and larger specimens
had meristic features of Anthias species (Figs. 16-20).
Only one western Atlantic type could be identified to
species {Anthias tenuis) on the basis of meristic charac-
ters of the larvae. Anthias nicholsi seems to be the most
common adult of this genus in the western Atlantic and
Anthias Type 1 is the most common larval type col-
lected; therefore, Anthias Type 1 may be Anthias
nicholsi. Since there is only one Anthias species in the
eastern Pacific (Anthias gordensis), the larvae are pre-
sumed to be that species (Fig. 20).

Anthias larvae are deep bodied and have large heads,
as is characteristic of all anthiines. The second dorsal
spine is elongate but thin in at least some species. The
first three dorsal spines form while the rest of the dorsal
fin is still an undifferentiated finfold. When all the dorsal
rays have formed, the third spine is the longest and the
spines approach the proportions of adults. The dorsal fin
spines are rather stout during later larval development,
but are never serrate. The elongate spine has pigment on
its associated membrane, giving it a flaglike appear-
ance. Pigment is retained on the membrane between the
second and third dorsal spines into the late larval period.
There is a deep notch in the dorsal fin, the last four

Figure 15. — Larvae of Plectranthias garupellus from the northwestern Atlantic Ocean: a) 5.5 mm; b) K.O mm.

spines being quite short. The pelvic fin forms early in
Anthias larvae and the first soft ray is produced. The
pelvic fin and first three spines of the dorsal fin develop
before other fin rays. The membrane of the pelvic fin is
pigmented, like the dorsal fin. The pelvic fin spine is
stout but not elongate or serrate.

The dorsal and anal fins form concurrently except for
the precocious anterior dorsal spines. The caudal fin
completes formation after the dorsal and anal fins have
formed. The pectoral fin is last to form rays, the dorsal

rays of the pectoral fin forming first, as in all serranids
studied to date.

Anthias larvae have characteristic head spines. In
some species there is a simple supraoccipital spine.
There is a serrate ridge above the eye. The degree of ser-
ration varies among the species, some having 3 to 6 serrae
while others have up to 12. The most conspicuous spines
are in the opercular region (Fig. 21). There are two
spinescent ridges on the preopercular. The anterior ridge
has 3 to 5 broad points. The posterior ridge has 4 to 8

Figure 16. — Larvae of Anthias sp. Type I from the northwestern Atlantic Ocean: a) 3.K mm; b) 5.3 mm.

strong points, some of them serrate. This ridge is marked
by a large serrate spine at the angle of the preopercular.
Lying directly proximal to this preopercular spine is a
similar spine on the interopercular, serrate and just as
large. This spine is obscured by the one on the pre-
opercular and is easily overlooked. The posttemporal
region has a number of protruding spines, some of which
are serrate.

Pigment on Anthias larvae varies among the species;
however, there seems to be a generic pattern. Usually a
small melanophore is at the posterior end of the anal fin,
one is on the ventral edge of the caudal peduncle, one or a
few spots are at the base of the central rays of the caudal

fin, usually a few spots are on the skull above the optic
lobes, and in some species there is a spot at the symphy-
sis of the lower jaw. Dorsal trunk pigment varies among
the species. In the Atlantic there are four types of
Anthias larvae based on dorsal trunk pigment and other
characters. Anthias Type 1 has a spot on the trunk mus-
culature ventral to the middle of the dorsal fin (Fig. 16).
In Anthias Type 2 there is a line of pigment ventral to the
posterior part of the dorsal fin (Fig. 17). Anthias tenuis
has some pigment between the dorsal body margin and
the lateral line ventral to the last few rays of the dorsal
fin (Fig. 18). Anthias Type 3 has no dorsal trunk pigment
(Fig. 19).

Figure 17. — Larvae of Ant hias sp. Type 2 from the northwestern Atlantic Ocean: a) 4.7 mm; b) 5.7 mm; c) .H.4 mm.

Figure 18.— Larva of Anthias tenuis from the northwestern Atlantic Ocean, 6.7 mr

Figure 19. — Larva of Anthias sp. Type 3 from the northwestern Atlantic Ocean, 5.1 mm.

These four types of Anthias larvae are distinguished by
other characters also. Of the types, Type 2 and Anthias
tenuis have a protruding supraoccipital crest. Also Type
2 is scaled while it is still pelagic and has larval pre-
opercular spines, pigment, and body shape. Although
there are not enough specimens for a thorough analysis,
there appear to be differences in the number of spines
above the eye with Type 1 and Anthias tenuis having the
fewest (about 4), Type 3 having more, and Type 2 having
the most.

Anthias gordensis has no dorsal trunk pigment, no pro-
truding supraoccipital crest, and a moderate number of
spines in the ridge above the eye. Thus, Anthias
gordensis larvae correspond most closely to Anthias Type

3 larvae among the Atlantic types. As more becomes
known about the relationships within this genus, clues
about the identity of Anthias Type 3 may be sought in
the Atlantic species most closely resembling the Pacific
Anthias gordensis.

Pronotogrammus . — There are three American species
of Pronotogrammus: P. aureorubens in the Atlantic and
P. eos and P. multifasciatus in the Pacific. These fish are
more slender and have smaller heads than other Ameri-
can anthiines.

In plankton samples, I found one larval type in each
ocean which can be ascribed to Pronotogrammus on the
basis of meristic features (Figs. 22, 23). The Atlantic type

Figure 20. — Larvae of Anthias gordensis from the eastern Pacific Ocean: a) 5.2 mm; b) fi.O mm.

has meristic features of P. aureorubens and the Pacific
type those of P. eos. The two larvae share common char-
acters that differentiate them from larvae of other
genera. None of the fin rays are serrate or much elongate
but the second and third dorsal spines form before the
other elements and the pelvic fin is formed early. The
opercular region is heavily armed with several serrate
spines (Fig. 24). The preopercular has a long serrate
spine at its angle and the interopercular has a similar

spine proximal to the one on the preopercular. The post-
temporal and supracleithrum have protruding serrate
spines. The supraoccipital crest protrudes as a simple
ridge until it is "grown over" in late-stage larvae. There
are several spines above the eye on a ridge of the frontal
bone. Midlateral trunk pigment is characteristic of both
types of Pronotogrammus larvae. In P. aureorubens this
pigment consists of 5 to 12 dashes along the midlateral
septum, roughly the length of the anal fin base. In P. eos

posttemporal

opercular

preopercular

'i \^ V supracleithrum

nteropercular

Figure 21. — Spine-bearing bones of the opercular and posttemporal
regions on a 10.2-mm Anthias sp. Type I larva.

the pigment consists of a group of melanophores that
form a blotch of pigment laterally on the trunk over the
last few rays of the anal fin. Other pigment consists of
several melanophores on the surface of the optic lobes of
the brain. A few spots of pigment are at the base of the
caudal fin on both species, and on P. eos some pigment is
on the pelvic fin membrane. Lower jaw pigment is pres-
ent on P. eos and two ventral midline spots are on P.
aureorubens — one at the insertion of the anal fin and one
midway on the caudal peduncle.

Hemanthias. — Hemanthias is represented by three
American species: H. vivanus and H. leptus in the Atlan-
tic and H. peruanus in the Pacific.

I found one type of Hemanthias larvae in each ocean,
the Atlantic species with 47 lateral line scales (Fig. 25) is
presumably H. vivanus, the most northerly species.
From larval catches, H. vivanus is the most abundant
anthiine off the east coast of the United States. The lar-
vae from both coasts share several characters which sep-
arate them from larvae of other genera. They are the
most ornately spined anthiine larvae. They have well-
developed spines in the opercular region, marked by an
elongate, serrate spine at the angle of the preopercular
and one underlying it on the interopercular. The inter-
opercular also has a series of heavy spines ventral to the
larger one. Both species have a protruding supra-
occipital crest which is serrate, resembling a "cocks-
comb." There are numerous ridges of complex spines on
the frontals and parietals. The frontals also have a ridge
of spines on their ventral margins. The posttemporal and
supracleithral bones have protruding, serrate spines. The
second through fourth dorsal spines develop early and
the third spine is quite elongate early in development.
The first dorsal spine develops after these three. The
pelvic fin also develops early. In both species the pelvic
fin spine is serrate and stout. In H. vivanus the second
and third dorsal spines as well as the first anal spine are

also serrate. Pigment on H. vivanus consists of opposing
spots on the trunk medial to the posterior extent of the
second dorsal and anal fins. There is also a spot ventrally
on the caudal peduncle and one at the caudal fin base.
Spots also occur on the dorsal surface of the optic lobes.
Hemanthias peruanus has pigment on the membranes of
the spinous dorsal and pelvic fins (Fig. 26). Both species
become scaled with ornate ctenoid scales while they still
possess larval characters.

Hemanthias vivanus. — Meristic element develop-
ment (Table 4) — In the smallest recognizable larvae on
hand (3.1-4.0 mm), some meristic elements are already
forming: the third dorsal fin spine and the pelvic spine
are present. The third dorsal spine remains prominent
throughout development with pronounced serrations. By
4.1-4.5 mm the dorsal fin has added one or two spines
(the second and fourth) and some specimens are forming
gill rakers. Between 4.6 and 5.0 mm the vertebral column
has two to six vertebrae ossified, the first dorsal fin has
three to five spines, the anal fin had one spine on about
half of the specimens, and the branchiostegal rays range
from two to six. By 5.1-5.5 mm, six branchiostegal rays,
one spine, and one ray in the pelvic fin, and four to nine
first dorsal spines are present. Between 5.6 and 6.0 mm
the vertebral column is adding vertebrae and 14 to 20 are
seen, the dorsal fin has 8 or 9 spines, there are 7 to 9 gill
rakers, and branchiostegal ray complement is complete
at 7. Between 6.1 and 6.5 mm there are 18 to 23 verte-
brae, 9 or 10 dorsal spines, 1 to 4 pelvic rays, and 8 to 12
gill rakers. The second dorsal, anal, pectoral, and caudal
fins are developing rapidly during this period showing
wide ranges of numbers of elements formed. Between 6.6
and 7.5 mm, the same pattern of wide ranges of formed
elements is seen, indicating rapid formation. Some spec-
imens have adult complements of vertebrae, second dor-
sal, anal, caudal, and pelvic fin rays. By 7.6-8.0 mm, the
adult complements of all characters examined except gill
rakers are present. Notochord flexion takes place at 5.5
mm and specimens >6.0 mm are developing scales. That
is, scales form when the only meristic character with its
adult complement is the branchiostegal rays.

Serrae are well developed on the third dorsal spine and
the pelvic spine. The second and fourth dorsal and the
first and second anal spines also develop serrae.

Body proportions (Table 5) — Larvae from 3.1 to 9.2
mm were measured to trace the proportional develop-
ment of several body dimensions. The only proportion
that shows noticeable difference with size is total length,
which increases from 110' V of BL to 120% of BL as the
larvae grow. This difference reflects the formation and
growth of the caudal fin. Greatest body depth remains
fairly constant ranging from 29 to 44' c of BL; the depth
at the anus is 30* c of BL; and caudal peduncle depth is
12' b of BL. Head length is about equal to greatest depth
ranging from 34 to 46' b of BL, snout length is 12c? of BL
as is eye diameter. The preanal length ranges from 53 to
65'V of BL. Thus, H. vivanus larvae are large headed and
deep bodied with a trunk that tapers quickly from the
anus to the caudal peduncle.

Figure 22. — Larvae of Pronotogrammus aureorubens from the northwestern Atlantic Ocean: a) 4.6 mm; b) 6.0 mm; c)

9.(1 mm.

Head spine development (Figs. 27, 28) — The serrate
"cockscomb" spine protruding from the supraoccipital is
pronounced in the smallest larvae (3.1 mm) observed.
Serrate ridges above the eye and transverse ridges of
spines on the surface of the cranium as well as beginnings
of spines in the posttemporal and opercular region are
also present. During larval growth, these spines increase
in complexity and size, those in the opercular area show-
ing the greatest change (Figs. 29, 30). The supra-
occipital crest remains the largest cranial spine and is
strongly serrate. The ridges on the cranium form a com-
plex pattern covering the area between the eyes and the
supraoccipital region (Fig. 28). The serrate supraorbital

ridges develop about eight strong, blunt points. In the
opercular region, the preopercular has a long serrate
spine at its angle and several spines along the rest of the
posterior margin of the bone (Fig. 30). The inter-
opercular has a long serrate spine which lies directly
proximal to the long spine on the preopercular and
several smaller spines ventral to the long one. The sub-
opercular has a few small spines on its posterior margin
and some are serrate. The opercular bone develops the
three spines characteristic of serranids on its posterior
margin during the larval period. The posttemporal has
several serrate spines and the supracleithrum has a pos-
teriorly directed, elongate serrate spine. Apparently

Posttemporal

supracleithrum

preopercular

Figure 23. — Larvae of Pronotogrammus eos from the eastern
Pacific Ocean: a) 3.6 mm; b) 7.8 mm.

these spines are resorbed in the late larval or early
juvenile period but specimens to describe these events
are lacking.

Pigmentation — Pigmentation is present on the small-
est larvae examined (3.1 mm) and remains essentially
unchanged during larval development. Two spots are
present on the ventral midline of the caudal peduncle,
one just posterior to the insertion of the anal fin and the
other, slightly smaller, at the anterior end of the caudal
fin base. Another spot develops at the base of the ventral
lobe of the caudal fin and extends along the bases of
several rays. Dorsally a spot is present on some larvae at
the insertion of the second dorsal fin. This spot seems to
be quite superficial, seen on only one side of the larva in
some cases, and apparently abraded off of other larvae. A
few melanophores are present at the symphysis of the
lower jaw, on the dorsal surface of the optic lobes of the
brain, on the fin membrane between the third and fourth
dorsal spines, and on the membranes between the rays of

Figure 24. — Spine-bearing bones of the opercular and posttem-
poral regions on a 9.5-mm Prontogrammus aureorubens larva.

Figure 25.— Larvae of Hemanthias vivanus from the northwestern Atlantic Ocean: a) 4.2 mm; b) 5.3 mm; c) 6.8 mm.

Figure 2(>.— Larva of Hemanthias peruanus from the eastern Pacific Ocean, 9.3 mm.

post temporal

posttemporal

opercular

preopercular

supracleithrum
subopercular

ropercuiar

Figure 27.— Spine-bearing bones of the opercular and posttemporal
regions of a 4.3-mm Hemanthias vivanus larva.

the pelvic fins. Internal pigment on the dorsolateral sur-
face of the gut shows through the body wall.

Discussion. — Larvae of four genera of American
anthiines have been found. These larvae are separated
from other serranids by their deep bodies, large heads,
produced preopercular and interopercular spines, a
tendency for development of armature on the head, and
serrations of fin spines. Within the anthiines, the genera
can be ranked according to the development of these
characters. Plectranthias has little development of these
characters. The preopercular spine is elongate and ser-
rate, but the interopercular spine is not serrate. There is
no protruding supraoccipital crest, neither the frontal

supracleithrum

preopercular

interopercular

Figure 28. — Spine-bearing bones of the opercular and posttemporal
regions on a 10.3-mm Hemanthias vivanus larva.

ridge over the eye nor the fin spines is serrate. Anthias
develops more extreme larval characters. The pre-
opercular and interopercular spines are stronger and
both are serrate. The ridge above the eye has several ser-
rae. In some species there is a nonserrated protruding

Table 4. — Meristic character development of larvae of Hemanthias vivanus. Specimens
between dashed lines are undergoing notochord flexion. For the caudal fin, "P" and "S"
indicate primary and secondary fin rays.

Body

Branchio-
stegal
rays

Verte-
brae

Pec-
toral
fin

Pel-
vic
fin

Caudal fin

Anal
fin

Dorsal fin
Spines Rays

length
(mm)

Dorsal Ventral
S P P S

Gill

rakers

3.2 NL

3.3

3.5

3.6

3.7

3.8

3.9

4.0

4.1

4.2

4.3

4.4

4.5

4.7

4.8

4.9

5.0

5.2

5.3

5.4

3
3
3
2
3
3
4
4

3
5
3
6
6
5
5
5
I J

III

111
111
III
III
III
III
III
III
111
IV
1\
IV
IV

III

V
V
V

VI
IV
V

2
2

damage

5.5

5.6 SL

5.6

5.7

5.8

5.9

6 I

6.3

6.4

6.5

6.8

7.0

20
14
18
16
16
15
14
18
18
23
18
20
22
23
20
26
18

II. 1

11,1

11.9

II. 1

11,1

[1,9

11,1
11,1

11,9
11,1

IX
VIII

IX
VIII
X
X
X
X

IX
X
X
X
X
X

8
9
10

Table 4.— Continued.

Bodv

Branchio-
stegal

Verte-

Pec-
toral

Pel-
vic

Cau

dal fin

Anal

Dorsal fin

length

Dorsal

Ventral

Gill

(mm)

rays

brae

fin

Spines

Rays

rakers

'7.1

1,5

11,9

'7.3

1,5

11,9

'7.3

1,2

n,i

'7.7

1,5

11,9

'7.7

1,5

11,9

'Scales present.

Table

-Body proportions of larvae of Hemanthias vivanus. Specimens
between dashed lines are undergoing notochord flexion.

Percent of standard length

Body

Pre-

Caudal

Depth

length

Total

Body

Eye

Head

Snout

anal

peduncle

(mm)

length

depth

length

depth

anus

3.1 NL

106

4.0

103

:\2

4.4

114

4.6

108

4.6

112

4.6

109

4.6

109

4.7 SL

116

5.0

116

5.0

113

5.2

116

5.2

118

1(1

5.4

123

5.5

117

5.7

121

6.0

118

6.0

122

6.3

122

6.5

124

6.8

117

(11

7.0

124

7.8

124

8.0

113

8.1

122

9.2

121

supraoccipital crest. The fin spines are strong and the
third dorsal spine and the pelvic spine are elongate.
Some species develop ctenoid scales while retaining lar-
val characters. Pronotogrammus develops more extreme
larval characters including a strong protruding supra-
occipital crest and heavy opercular armature. These lar-
vae, however, are not scaled. Among American anthiines,
Hemanthias exhibits the most extreme development of
larval characters. The head is armored with a serrate
supraoccipital crest, numerous serrate ridges on the fron-
tals and parietals, and heavy serrate spines on bones in
the opercular series. Ctenoid scales develop during the
larval period. In Hemanthias vivanus larval anthiine
characters reach their apex with the development of ser-
rations on several dorsal and anal fin spines.

Figure 29. — Development of the supraoccipital crest of
Hemanthias vivanus: a) 4.3 mm; b) 5.8 mm; c) 6.6 mm.

Figure 30.— Details of the head spination on a Hemanthias vivanus larva, 7.2 mm.

Subfamily Epinephelinae

The Epinephelinae is represented by three genera in
American waters from both oceans. These are Paranthias
with 1 species, Mycteroperca with 13 species, and Epi-
nephelus with 21 species grouped in 5 subgenera (Table
1). Epinephelus is worldwide in warm water while the
other two genera are indigenous to the Americas. The
monotypic genus Gonioplectrus has close affinities to the
epinephelines based on larval and adult characters (Ken-
dall and Fahay in press). Members of this subfamily are
quite disparate in size as adults, some reaching only 30
cm, while others reach 180 cm and a weight of 320 kg.
Until recently the subgroupings of Epinephelus were
given generic status but Smith (1971) regarded them as
subgenera.

Larvae of several species of Epinephelus have been il-
lustrated and described. The species are outstanding in
having the second spine of the dorsal fin and the pelvic
spine strong, serrated, and produced to nearly the length
of the larva. These larvae are frequently associated with
the entire family and have been cited as representatives
of the family (e.g., Rosenblatt and Zahuranec 1967;
Smith 1971). Among the earliest published illustrations
of Epinephelus larvae are those of Fage (1918) who called
them Serranus scriba. Bertolini (1933) reviewed Fage's
work and indicated that the fish was Epinephelus guaza.
Sparta (1935) figured the eggs of Epinephelus guaza and
the larvae of Epinephelus alexandrianus. Fage's (1918)
figures and description were reviewed by Vodjanitzki and
Kazanova (1954). Ukawa et al. (1966) and Mito et al.
(1967) described larval development of reared Epine-
phelus akaara from Japan. Guitart Manday and Juarez
Fernandez (1966) described eggs and early stage larvae of
Epinephelus striatus obtained from aquarium held
adults.

Aboussouan (1972) illustrated and described three
types of larvae from the west coast of Africa that he
ascribed to the genus Epinephelus. However, only one of
these, E. aeneus, has the larval characteristics of an epi-
nepheline. The other two do not have elongated fin
spines; in fact, the spinous dorsal and pelvic fins develop
after the soft dorsal and anal. The preopercular margin is
comprised of several spines with the one at the angle only
slightly longer than the rest and not serrate. Pigment is
made up of small melanophores as opposed to the large
caudal peduncle blotch of epinephelinesm. These larvae
are almost certainly not epinephelines, rather one is pos-
sibly a sciaenid and the other a haemulid, judging from
the number of fin elements and general larval
morphology. Presley (1970) described Epinephelus
niveatus larvae from the Florida Straits. Smith (1971) il-
lustrated a larval Epinephelus from off the Carolinas.
Fowler (1944) described a new genus and species (Serri-
hastaperca exul) from a larval fish from the eastern Pa-
cific. As Heemstra (1974) has shown, this is probably a
larval Epinephelus, possibly Epinephelus panamensis.
Fourmanoir (1976) illustrated and briefly described three
epinepheline larvae from the southeastern Pacific. Ken-
dall and Fahay (in press) described a larval Gonio-

plectrus hispanus and concluded that although it had
some characters of the anthiines, it was more closely
related to the epinephelines.

Although the figures are not as detailed as might be
desired, and the descriptions are generally brief, most
Epinephelus larvae described thus far appear quite
similar. They all possess stout, elongate, serrate pelvic
fin spines and second dorsal spines. The preopercular is
armed with an elongate spine at its angle and several
smaller spines. The first spine of the dorsal fin is short
and serrate, the second long and serrate, and the third
somewhat elongate and serrate. The succeeding spines
become progressively shorter and less robust until it is
difficult to distinguish the last dorsal spine from the
first dorsal soft ray. Pigmentation generally consists of a
few spots on the surface of the skull, a spot at the junc-
tion of the cleithra, internal pigment lateral to the gut,
and a large spot on the caudal peduncle. The elongate fin
spines also have some pigmentation on their membranes.

The larval material I have examined contains many
species of epinepheline larvae all agreeing closely with
the above descriptions. There are no forms which possess
only some of this suite of characters, rather they are all
uniform and distinguishable from other types of serranid
larvae. Besides specimens having meristic characters of
Epinephelus sp., there are larvae representing the two
other American genera, Mycteroperca and Paranthias.
Larvae of these genera can be separated at present on the
basis of meristic characters but not larval morphology.

Paranthias. — The genus Paranthias contains one
species (P. furcifer) which is found on both sides of the
Americas. It is separated from other epinephelines by
morphology and habits. It converges with anthiines in
several of its adaptations, although on skeletal and other
characters its affinities clearly lie with the epinephelines
(Table 1). As opposed to the demersal, piscivorous habits
of other members of the subfamily, P. furcifer occurs in
aggregations somewhat above the bottom where it feeds
on plankton (Randall 1968). Morphological character-
istics related to these habits include a deeply forked
caudal fin, dorsal and ventral profiles nearly equally
curved, a smaller head with a small upturned mouth,
and more gill rakers (35-40) than other epinephelines
(Smith 1971).

As larvae, the three genera of American epinephelines
can only be separated using meristic characters. These
are sufficiently formed only late in development. Sepa-
ration of Paranthias from the other two genera depends
on the number of dorsal fin rays. The posteriormost
spines in the dorsal fin of epinephelines are short, late
forming, and first form as rays. The total count of dorsal
spines and soft rays in Paranthias (27-28) is commonly
seen in members of the other two epinepheline genera
also. At present, only large, well-developed larvae of
Paranthias furcifer have been identified.

As mentioned, Paranthias furcifer larvae share larval
characters with all other epinephelines found so far (Fig.
31). The pelvic spine and second spine of the dorsal fin
are long and serrate. They bear pigmented "flags" near

Figure 31. — Larva of Paranthias furcifer from the eastern Pacific Ocean, X.(i mm.

their tips, which are often abraded off in netted speci-
mens. The first and third dorsal spines and second anal
spine are also serrate. The third dorsal fin spine is mod-
erately elongate. The spine at the angle of the pre-
opercular is long and serrate and there are usually two
spines on the preopercular bone dorsal and ventral to the
spine at the angle. There is a serrate ridge of spines dor-
sal to the eye and the supracleithrum has a protruding
blunt serrate spine. Pigment spots are present over the
optic lobes of the brain and laterally over the gut. There
is a large, intense blotch midlaterally on the caudal pe-
duncle.

A 34-mm specimen collected at the surface as part of a
large school showed loss of larval characters. The body
was fully scaled and the supraocular ridge and the pro-
truding supracleithral spines were unpronounced. The
second dorsal spine was only slightly larger than the
third and all were well formed and nearly equal in length.
The second spine retained minute serrations. The pelvic
spine also retained serrations and was about as long as
the second dorsal spine, not reaching the posterior end of

the pectoral fin. The first ray of the pelvic fin was longer
than the spine. The spine at the angle of the pre-
opercular was still slightly elongate and serrate; but
rather than two spines on each side of the spine at the
angle, there were several (6 ventral and 20 dorsal). The
specimen I examined was cleared and stained so no as-
sessment of pigment pattern could be made.

Epinephelus. — This is the only American epi-
nephrine genus with representatives in other parts of the
world. Epinephelus is the most speciose genus of ser-
ranids, containing 21 American species grouped in 5 sub-
genera. Fifteen species occur in the Atlantic and 11 in the
Pacific, with 5 species common to both oceans. They are
separated from other American epinephelines by a com-
bination of characters (Table 1). Those species with IX
dorsal spines have fewer than 18 rays, whereas
Paranthias furcifer has a dorsal fin count of IX, 18-19.
Epinephelus species differ from Mycteroperca by having
8 or 9 anal rays, rather than 10 to 12. Other diagnostic

characters include characteristic skull crests and the
presence of a postocular process (Smith 1971).

As discussed earlier, larvae of several species of Epi-
nephelus from around the world have been described
(e.g., Bertolini 1933; Mito et al. 1967; Fourmanoir 1976).
Presley (1970) described larvae he assigned to Epi-
nephelus niveatus from Florida. His description is rather
brief, and considering the similar appearance of all epi-
nephrine larvae that have been described and all those I
have found, it is probably inadequate to separate that
species from others in the area.

Larval epinephelines are not separable into genera un-
til medial fin rays have formed. Only on the basis of these
counts could Epinephelus larvae be distinguished from
those of Mycteroperca (by a lower anal fin ray count) and
from Paranthias (by a different dorsal fin count). Two
larval specimens of Epinephelus are illustrated (Figs. 32,
33). They are both about the same length and appear
similar. One specimen (Fig. 32) from the Atlantic has XI,
16 dorsal fin elements and III, 9 anal fin elements making
it most likely a member of the Epinephelus striatus
species group and probably either E. morio or E. gut-
tatus. The other specimen (Fig. 33) is from the eastern

Pacific with XI, 19 dorsal fin elements and III, 9 anal fin
elements making it most likely either E. dermatolepis
dermatolepis or E. alphestes multiguttatus . Although
these specimens are in different subgenera, no charac-
ters other than meristics could be found to distinguish
them from other genera within the subfamily.

The following account describes features of all larval
Epinephelus specimens seen so far, since no characters
have yet been found to identify the species. Most of the
characters mentioned with the account of the subfamily
apply to the specimens identified as Epinephelus sp. The
second spine of the dorsal fin is elongate and serrate as is
the pelvic spine. The first and third dorsal fin spines are
also serrate and the fourth may be as well. The anal fin
forms with only the first two spines as such until late in
development. The third anal fin spine forms as a ray and
later becomes a spine. The first two anal spines may be
serrate. The preopercular margin is armed with an elon-
gate, serrate spine at its margin and one or two blunt
spines dorsal and ventral to the one at the angle. Later in
development more spines form on this margin and it
becomes serrate. There is a serrate ridge above the eye
and the posttemporal and supracleithral bones have pro-

Figure 32. — Larva of a member of the Epinephelus striatus species group from the northwestern Atlantic Ocean,

7.6 mm.

Figure 33. — Larva of an Epinephelus sp. (subgenus Dermatolepis or Alphestes) from the eastern Pacific Ocean,

8.4 mm.

trading, serrate spines. The pigment pattern consists of a on the lateral surface of the gut, and a large blotch on the

few melanophores on the surface of the optic lobes, some caudal peduncle. The caudal peduncle blotch is var-

iously placed, from along the ventral midline to laterally
on the lateral line. It tends to become internal on larger
specimens. The elongate dorsal and pelvic spines are var-
iously pigmented but the pigment is absent on many net-
ted specimens. The pigment is seen on the membranes
associated with the extended portion of the spines.

Besides the transformation of the third anal spine from
a ray to a spine during development, as many as two pos-
terior spines of the dorsal fin undergo similar transfor-
mation. The second dorsal fin spine is elongate and the
third through seventh are progressively smaller. The 7th
through the 11th dorsal spines are quite small during lar-
val development. It seems that the first nine spines
develop as such, but the last two develop as soft rays and
later transform into spines. However, in specimens which
have only 9 or 10 spines as adults, no such transforma-
tion may occur.

The bones in the opercular region of a 10-mm Epi-
nephelus sp. (possibly E. niveatus) are characteristic of
the genus (Fig. 34). The preopercular bone has the most
produced spines of any of the bones in the region. As
mentioned above, it has a long, serrate spine at its angle
and two spines dorsal and ventral to the one at the angle.
These spines may also be serrate. The opercular has a
broad fan shape with three blunt spines. The area ventral
to the lower spine is broad. The subopercular has a spine
on its posteroventral margin. The interopercular has a
spine on its posterodorsal edge. The posttemporal and
supracleithrum each have a protruding serrate spine.

posttemporal

opercular

supracleithrum

preopercular

Figure 34. — Spine-bearing bones of the opercular and posttemporal
regions on a 10.2-mm Fpinephelus niveatus larva.

Mycteroperca. — This is the second most speciose epi-
nepheline genus with eight species in the Atlantic and
five in the Pacific Ocean (Table 1). They are distin-
guished from the other two epinepheline genera by hav-
ing 10 to 13 anal rays and straight, parallel lateral skull
crests that join the rim of the orbit anterior to the mid-
orhital area (Smith 1971).

Larvae with developed III, 10-13 anal fin elements and
epinepheline larval characteristics were found in samples
from both oceans. They had other diagnostic characters
of Mycteroperca. These larvae had the full suite of char-
acters of epinepheline larvae and, on the basis of larval
characters, could not be separated from other epi-
nepheline larvae. They had elongate, serrate second dor-
sal and pelvic spines and a long serrate spine at the angle
of the preopercular. There was a serrate, bony ridge
above the eye and protruding, serrate spines in the post-
temporal region. They had a few pigment spots on the
dorsal surface of the optic lobes of the brain, some on the
lateral surface of the gut, and a large blotch on the
caudal peduncle. Some pigment was present on the
membranes of the elongate fin spines.

Mycteroperca microlepis (Figs. 35, 36). — The mer-
istic characteristics of Mycteroperca microlepis fall
within the ranges for several species of Mycteroperca in
the western Atlantic (Table 1). However, among these
species, M. microlepis is the most northerly occurring.
Late larvae and early juveniles with characteristics of M.
microlepis have been caught in the Middle Atlantic
Bight.

In plankton samples from as far north as Cape Hat-
teras, N.C., large epinepheline larvae (>7 mm) with
meristic characters of M. microlepis are frequently
found. In these samples smaller larvae with similar larval
characters but incomplete meristic characters are also
found. The following description is based on these epi-
nepheline larvae taken off North Carolina. Larger larvae
and juveniles (12-35 mm) from nearshore and estuarine
collections along the Atlantic coast from Florida to New
Jersey were examined to note the development of
juvenile pigment and the regression of the elongate
spines of early larvae. No other species of Mycteroperca
could be identified in the samples with certainty.

Meristic element development (Table 6) — Develop-
ment of most meristic elements occurs over a length
range of 4.0-9.8 mm. In the smallest larvae available (4
mm), the dorsal fin already has the first three spines ossi-
fied, the pelvic fin has one spine and one ray, and there
are three branchiostegal rays and two gill rakers. Be-
tween 5 and 6 mm, the notochord is flexing and most
meristic features are forming ossified elements. The ossi-
fied vertebrae increase from 4 to 18, forming from ante-
rior to posterior except for the urostyle. Several verte-
brae are partially developed in most specimens. The
branchiostegal rays reach their adult complement of 7
and there are 6 to 10 gill rakers. The pelvic fin has three
rays ossified and the pectoral fin starts to form rays. The
dorsal fin increases from three to five spines and the anal
starts to form. The caudal fin has its adult complement
of nine dorsal and eight ventral primary rays as the no-
tochord becomes fully flexed at 6 mm. When notochord
flexion is complete, by 7 mm, most meristic features are
well formed, approaching adult counts. Vertebrae in-
crease to 22 with the urostyle forming prior to the 2 or 3
vertebrae anterior to it. The pectoral fin increases to 12
rays and the pelvic fin attains its adult complement of 1

spine and 5 rays. The anal fin increases rapidly having 2
spines and 11 rays. The first soft ray of the anal fin
becomes the third spine later in development, as is char-
acteristic of most adult serranids with three anal spines.
The dorsal fin is difficult to count separately because the
posterior spines are quite small at formation and, like the
third anal spine, originate as soft rays. It appears that
there are 10 spines and 15 rays in the dorsal fin. In addi-
tion to the 9 or 10 gill rakers on the lower limb, 1 forms on
the upper end of the first gill arch. Procurrent caudal
rays start to ossify. Between 7 and 10 mm, the meristic
elements, except gill rakers and scales, reach their adult
complements. Not until about 20 mm do the 3rd anal
spine and 11th dorsal spine change from soft rays to
spines.

Body proportions (Table 7) — Proportions of various
measurements of larvae and juveniles of Mycteroperca
microlepis to body length (BL) were examined. Most
proportions show little ontogenetic change; however,
total length increases sharply with the formation of the
caudal fin and the depth at the anus and caudal pedun-
cle depth also increase between 4 and 6 mm. The greatest
body depth remains between 29 and 35% of BL through-
out the size range examined. The head length stays
about 31 to 43% of BL. The preanal length is 51 to 66% of
BL. Eye and snout length are about 9 to 14% of BL. The
elongated larval spines were measured to trace their
relative growth. The second dorsal spine was broken on
several smaller specimens but it appeared to increase
from 40% of BL at 4.6 mm to 60% from 5 to 10 mm and
then decrease to 11% by 35 mm. The pelvic spine showed
a similar pattern of increase and decrease, reaching a
maximum relative size of 68% of BL at 8.3 mm and de-
creasing to 12% of BL by 35 mm. The preopercular spine
increases from about 0.5% of BL at 5 mm to 1.1% at 8
mm and decreases to 0.2% by 35 mm.

Pigmentation — The pigment pattern seen on the
smallest larvae examined remains basically unchanged
throughout larval development. On the heads of 4-mm
larvae there is a spot on the dorsoposterior surface of
each optic lobe, and there are several spots in the same
area on 6-mm specimens. A spot develops internally on
the nape near the dorsal fin origin which later occurs in
the area of the neural spines of the first two vertebrae.
The dorsolateral surface of the gut cavity is covered with
rather dense pigment throughout larval development.
Ventrally, on most specimens there is a spot at the
cleithral junction. A large spot occurs on the ventral
region of the caudal peduncle in small larvae. In 7.5-mm
specimens this spot has moved dorsally to lie over the
lateral line in the midlateral caudal peduncle area. This
spot extends internally to the area of vertebrae 20 to 21.
The size and vertical position of this spot varies some-
what among the specimens observed from along the
ventral midline to the lateral line. A few small spots that
develop on the ventral tip of the notochord lie near the
bases of the caudal rays as the caudal fin is formed.

Opercular series spines (Fig. 36) — Spine-bearing bones
of the opercular region of a 9.8-mm specimen of Mycter-

operca microlepis appear similar to those of other epi-
nephelines. The preopercular bone has an elongate,
serrate spine at its angle which reaches a maximum of
1.3% of BL in 8-mm specimens. Smaller spines form
along the vertical and ventral edges of the preopercular.
During larval development, there are two spines in each
area with all but the uppermost spine being serrated.
Later in development, as the elongate spines regress,
additional small spines appear along the edge of the pre-
opercular to form the serrate margin of the adult. The
interopercular bears two blunt spines along its posterior
margin. The subopercular has one blunt spine at its
ventralposterior angle. The opercular is fan-shaped with
three blunt spines on its posterior margin. The supra -
cleithrum has a posteriorly directed, serrate spine and
the lateral posttemporal ridge is serrate.

Discussion. — The 35 presently recognized species of
epinephelines in American waters constitute a cohesive
group (Smith 1971). Recently, five genera have been
given subgeneric rank, leaving the species grouped in
only three genera. Epinephelus, the most speciose Amer-
ican genus with 21 species, is a speciose genus in other
parts of the world also, e.g., there are 34 species in Japan
(Katayama 1960). Among American genera, Paranthias
has evolved obvious ecological and morphological dis-
tinctions from the general "grouper" characteristics. A
clearer understanding of the relationships within this
subfamily must await its comprehensive revision on a
worldwide basis.

Epinepheline larvae representing all three American
genera were found. Their identification was based solely
on meristic characters. No larvae of epinephelines could
be separated on the basis of any larval characters. They
shared all recognized larval characters. Thus it appears
that evolutionary differences within this group are not
reflected in larval morphology.

Epinepheline larvae are quite distinct from larvae of
other serranids. The other subfamily with serrated
spines, Anthiinae, does not have the extremely elongate,
serrate fin spines seen in epinephelines. In anthiines the
interopercular has a produced spine which lies under the
spine at the angle of the preopercular, both of which are
usually serrate. Epinephelines have only a blunt spine on
the interopercular. Anthiines also are heavier bodied
than epinephelines. Gonioplectrus shares trenchant lar-
val characters with epinephelines but has less variation
in dorsal spine length and a stouter body, both reminis-
cent of anthiine larvae. These resemblances to anthiine
larvae may represent convergence. However,
Gonioplectrus may represent an evolutionary plateau
between anthiines and epinephelines; this remains for
further study to elucidate. Serranines have simple ar-
mature on the preopercular and no fin spine serrations.
Grammistines have the second spine of the dorsal fin
produced into a long, filamentlike structure, in contrast
to its stout-serrated form in epinephelines. Apparently,
in American waters at least, intermediates between
epinephelines and other groups of serranids have been
lost.

Figure 35.

-Young stages of Mycteroperca microlepis from the northwestern Atlantic Ocean:

mm; d) 22. (i mm.

a) 4.0 mm; b) 7.4 mm; c) 14.2

Table 6. — Meristic character development of larvae of Mycteroperca microlepis. Specimens
between dashed lines are undergoing notochord flexion. For the caudal fin, "P" and "S"
indicate primary and secondary fin rays.

Body

Branchio-
stegal

Verte-

Pec-
toral

Pel-
vic

Caudal fin

Anal

Dorsal fin

length

Dorsal

Ventral

Gill

(mm)

rays

brae

fin

Spines

Rays

rakers

4.0 NL

1,1

4.6

1,2

III

5.1

1,2

III

5.6

t,3

III

5.3

1,3

III

5.3

1,3

5.3

1,3

5.5

damage

1,6

6.0 SL

1,4

11,9

1+9

6.1

1,4

1,6

7.0

1,5

11,11

1 + 10

7.4

1,5

11,11

2+ 9

7.9

1,5

rj.,13

3 + 10

8.1

1,5

n,i2

3+10

9.2

1,5

11,11

3+12

9.8

1,5

11,12

4 + 11

■M

Table 7. — Body proportions of larvae of Myctoperca mierolepis. Specimens between dashed lines are undergoing

notochord flexion.

Percent of standard length

Body

length
(mm)

Total

length

Body
depth

Eye

length

Head
length

Caudal
Snout Preanal peduncle
length length depth

Depth

at
anus

Second
spine of
dorsal fin

Pelvic

fin
spine

Preoper-
cular
spine

4.6 NL

104

30.4

10.9

32.6

9.6 52.2 7.6

15.2

39.1

0.43

5.3
5.3
5.3
5.5
5.6
5.9

106
109
L13

110
107
112

32.1
34.0
34.0
36.4
32.1
32.2

11.3

11.9
12.1
12.7
11.1
10.8

32.1
35.8
35.8
40.0
35.7
42.4

9.4
10.6
12.1
13.6
10.9

50.9
56.6

54.7
56.6
55.4
64.4

8.5

8.3
9.4

10.2
7.9

10.5

18.9
20.8
26.4
25.5
19.6
22.0

69.2

49.1

54.7
58.5

0.58
0.75
0.94
0.91
0.55
1.02

6.0 SL

120

36.7

13.0

38.3

11.5

60.0

11.5

23.3

61.7

0.93

6.1

120

34.4

12.3

41.0

13.0

67.2

10.2

26.2

—

59.0

0.92

7.0

120

34.3

12.1

37.1

11.6

61.4

11.6

24.3

54.3

61.4

0.99

7.4

122

33.8

11.9

39.2

11.5

60.8

11.1

21.6

—

62.2

1.15

7.9

120

32.9

11.9

38.0

10.8

60.8

11.9

24.1

—

60.8

1.04

s 1

121

34.6

9.9

37.0

11.6

59.3

13.6

24.7

59.3

58.0

0.99

8.3

119

33.7

12.0

38.6

13.3

57.8

12.0

25.3

65.1

67.5

1.33

9.2

123

33.7

12.0

38.0

12.0

57.6

12.0

23.9

59.8

64.1

1.02

9.4

121

35.1

12.8

38.3

10.6

59.6

12.8

25.5

58.5

63.8

1.06

9.8

123

33.7

11.2

39.8

14.3

66.2

11.2

26.5

60.2

65.3

1.02

12.0

123

35.0

11.7

38.3

10.8

57.5

13.3

28.3

49.2

55.0

1.00

13.6

125

33.1

9.6

37.5

11.8

58.1

11.8

27.9

39.7

42.6

0.92

14.0

131

33.6

10.7

37.9

11.4

62.9

10.7

25.7

39.3

47.9

—

16.8

123

32.7

10.1

39.3

9.5

61.9

11.9

25.6

40.5

39.3

0.83

17.6

122

31.8

11.4

35.3

8.5

58.0

10.2

24.4

40.3

46.0

0.91

19.3

115

31.1

10.4

36.3

9.3

58.5

10.9

24.9

21.3

20.2

0.62

21.1

122

31.8

10.9

31.3

13.7

60.7

10.9

27.0

20.4

29.4

0.52

21.5

120

30.2

10.7

37.2

10.7

52.1

11.2

25.6

20.5

17.2

0.65

24.4

125

32.8

11.1

42.6

11.1

63.5

11.1

32.8

13.5

14.8

0.33

35.4

120

28.5

9.0

36.2

9.0-

61.6

10.7

27.1

11.0

12.1

0.14

opercular

preopercular

supracleithrum

subopercular

interopercular

Figure 36. — Spine-bearing bones of the opercular and posttemporal
regions on a 9.8-mm Mycteroperca mierolepis larva.

Subfamily Grammistinae

Larvae of three genera, Liopropoma (larvae of Pikea
may be included here), Rypticus, and Pseudogramma,

have been recognized and will here be considered
members of a single group, the subfamily Grammistinae.
These larvae are similar in appearance and readily
separable from larvae of other fishes. Considerable work
has been done on the taxonomic affinities of these fishes
but their relationships remain perplexing. Liopropoma
has been considered a member of the serranid subfamily
Liopropominae (Katayama 1960). Rypticus has been
considered a member of the family Grammistidae, a
closely related offshoot of the serranids (Gosline 1966).
Pseudogramma has been considered a pseudogrammid, a
group which has been considered a subfamily of the
grammistids or a separate family (see Randall et al.
1971). Reasons for considering these fishes members of a
single group are detailed in Kendall (1976, 1977).

Liopropoma. — Pikea has been included with
Liopropompa in the subfamily Liopropominae. They
have identical predorsal bone patterns and similar
meristic formulae (Table 1). Chorististium has been syn-
onymized with Liopropoma. Two genera, Joboehlkia
and Flagelloserranus, described from preadult material,
apparently belong with this group. Pikea has three north-
western Atlantic species and Liopropoma includes five
species in the northwest Atlantic, one in the eastern
Pacific, and several from the central and western Pacific.
Within this group, larvae of Liopropoma and/or Pikea

have been found in material from the northwest Atlantic.
These larvae appear similar to those described by Kott-
haus (1970) as the genus Flagelloserranus. He described
two species of Flagelloserranus, one from the Indo-
Pacific Ocean and one from the western Atlantic Ocean.
Fourmanoir (1971, 1976) indicated that similar speci-
mens from near New Caledonia were young of
Liopropoma. All the characters Kotthaus (1970) used to
define this genus and separate it from Liopropoma ap-
pear to be transient larval or juvenile characters. The
similarity of meristic and other characters, the unique
predorsal bone pattern, and the lack of adults of Flagel-
loserranus indicate that these are larval Liopropoma or
Pikea. Thus, Kotthaus (1970) provides descriptive
material for larval development of Liopropoma-Pikea so
it will be considered only briefly here.

I considered larval types resembling Flagelloserannus,
one with pigment on the ventral caudal peduncle surface
and one without, members of the genera Liopropoma
and/or Pikea. However, since the meristic characters and
areas of occurrence of the several species of these two
genera overlap, I did not attempt to allocate the larvae to
species.

Their general body shape is similar to that of the ser-
ranines, although the gut is shorter and there is a space
between the anus and the origin of the anal fin (Fig. 37).
The caudal peduncle is both longer and deeper than it is
in serranines.

The most outstanding developmental feature is the
presence, even in small larvae, of two elongate, thin dor-
sal spines. These develop before other fin rays, reach a
length of up to three times the fish length and become
the second and third dorsal spines. These spines are deli-
cate and are broken in many specimens. Kotthaus (1970)
described the presence of thick tissue surrounding these
spines; the tissue around the second spine having two
vanelike swellings on its distal third, the tissue around
the third spine being tubular for its entire length. The
distal portion of both spines is pigmented with several
large melanophores. The remaining fin rays develop their
adult proportions without any pronounced elongations.
The ventral fins develop more slowly than those of most
other serranids. The pectoral fins do not develop
precociously, as they do in Rypticus and Pseudogramma.

Except for the pigment on the elongate dorsal fin
spines, most larvae examined were unpigmented. Prior
to notochord flexion some larvae bear a melanophore on
the ventral midline anterior to the insertion of the anal
fin. Whether this spot disappears in larger larvae or
larger specimens with this pigment were not collected
cannot be determined presently. Some spots develop on
the hindbrain surface in larger larvae, probably
representing the onset of juvenile pigment.

The bones of the preopercular region are armed with
characteristic spines (Fig. 38). The preopercular has four
blunt spines above and three below the one at the angle.
Flagelloserranus meteori from the Indo-Pacific has a
similar preopercular with two spines above and three
below the one at the angle (Kotthaus 1970). Flagelloser-
ranus danae from the western Atlantic has a similar pre-

opercular except it has a hooked dorsalmost spine and
the vertical limb is longer than in F. meteori (Kotthaus
1970). The opercular of the larvae examined in this study
as well as those of Kotthaus (1970) is fan-shaped with
three spines on its margin. Both the subopercular and
interopercular have a blunt spine on the posterior edge in
the larvae I examined; however, Kotthaus (1970)
reported a smooth margin in both species he recognized.
A low posttemporal ridge that penetrates the epidermis
of the larvae I examined was not mentioned by Kotthaus
(1970).

Rypticus. — Occurring in the Atlantic and eastern
Pacific Oceans, this genus has meristic characters that
are unusual among serranids (Table 1) and small, em-
bedded cycloid scales. The unique gonadal morphology
(Smith and Atz 1969) and predorsal bone patterns (Ken-
dall 1976) also indicate that Rypticus is highly special-
ized and no other known fish can be derived from it.
These are secretive, nocturnal reef fishes of small to
moderate size (50-260 mm). Altogether, there are about
11 species, with 8 of them occurring in the western Atlan-
tic (Randall et al. 1971). Atlantic members of the genus
have been reviewed by Courtenay (1967).

Aboussouan (1972) illustrated and briefly described a
single 9.0-mm larva as Rypticus saponaceus and a 4.5-
mm specimen as "Rypticus (?)." Courtenay (1967) il-
lustrated a 15.5-mm juvenile of Rypticus saponaceus.

Only one larval specimen with meristic characters of
Rypticus was found in collections from the Pacific, while
the genus was represented by many specimens in collec-
tions from the Atlantic. Small larvae were so similar in
body shape and lack of pigment to Liopropoma and
Pseudogramma that it was difficult to separate them un-
til some fin elements were well formed. The only pigment
seen on the larvae was on the fleshy material enveloping
the single elongated first dorsal spine (Fig. 39). This
pigmented "flag" is more than half the body length; how-
ever, it is fragile and often broken in plankton-collected
material. Aboussouan (1972) seems to show rather heavy
pigment on the large fanlike pectoral fin of the 9.0-mm
larva he illustrated; however, I saw none on the material
I examined. Besides the elongate dorsal spine and the
large pectoral fin, the rays of the second dorsal and anal
fins are also uniformly long and the caudal fin appears
rounded. The pelvic fins are quite short in contrast to
their large size in most other serranids. In most fish lar-
vae with an elongate dorsal spine, the pelvic spines are
comparably elongate (Ahlstrom'); Rypticus, however,
has an elongate dorsal spine but a relatively undevel-
oped pelvic fin. Aboussouan (1972) reported an element
which resembled a spine in the anal fin; I found none in
the specimens I examined. Most serranids have three
anal spines, the third of which originates during larval
development as a soft ray. Adult Rypticus have no anal

7E. H. Ahlstrom, National Marine Fisheries Service, Southwest Fish-
eries Center, La Jolla, CA 92038, class notes, August 1971.

Figure 37. — Larvae of Liopropoma sp. from the northwestern Atlantic Ocean: a) fi.3 mm; b) 7.0 mm.

spines. If larvae of some Rypticus have an anal spine, it is
apparently resorbed as the fish develops.

The body shape is similar to that of the serranines; but
the nape is deeper and the bases of the second dorsal and
anal fins are fleshier.

The bones of the preopercular region are similar in lar-
val spination to those of Liopropoma, although speci-
mens of the same size as those observed of Liopropoma

were not available for study, so comparisons may be in-
valid.

Pseudogramma. — Rhegma has been synonymized
with Pseudogramma (Gosline 1960) and at times con-
sidered a serranid, a grammistid, or a pseudogrammid
(Kendall 1977). Larval and other characters indicate
that this genus is closely related to Rypticus and

posttemporal

opercular

supracleithrum

preopercular

Figure 38.— Spine-bearing bones of the opercular and posttemporal
regions of a 12.0-mm Liopropoma sp. larva.

Liopropoma and they all were derived from epi-
nephrine serranids. Pseudogramma contains one species
from each of the following areas: the central Pacific, the
eastern Pacific, and the western Atlantic. They have in-
complete lateral lines and moderate-sized scales
(Schultz 1966).

Larvae assignable to this genus by their meristic
features (Table 1) have been found in samples from the
western Atlantic; and since this genus is represented in

this area by a single species, it is assumed to be Pseudo-
gramma gregoryi. Although there is only a small series in
the present collections, it will be described in detail,
since both of the other genera in this group contain sev-
eral species, none of which could be identified using lar-
val characters. Also Liopropoma has been dealt with
previously (Kotthaus 1970).

Pseudogramma gregoryi (Fig. 40). — The proper
generic name and number of species assigned to the
Pseudogrammidae in the northwestern Atlantic have
been sources of confusion. Gosline (1960) and Schultz
(1966) listed three species, one in Pseudogramma and
two in Rhegma. At present, Pseudogramma gregoryi is
the only recognized pseudogrammid species in this area,
all others being synonymized with it. However, the
reported range of meristic characters among species syn-
onymized with P. gregoryi indicates that there may be
more than one valid species.

Meristic element development (Table 8) — A series of
18 larvae from 3.3 to 10.3 mm was cleared and stained.
At 3.3 mm, the notochord is straight but 12 vertebrae (10
precaudal and 2 caudal), 4 branchiostegal rays, 15 pec-
toral rays, 1 dorsal spine, and 2 gill rakers are already
ossified. The notochord undergoes flexion from 3.6 to 4.2
mm and during this time the vertebrae increase to 19, the
branchiostegal rays to 6, the pectoral rays to 16, the dor-
sal spines to 2, and the gill rakers to 4. The caudal fin
begins formation and has six dorsal lobe and six ventral
lobe primary rays. The dorsal fin spines which become
elongate during larval development are the second and
third. Most of the meristic complements are complete in

Figure 39.— Larva of Rypticus sp. from the northwestern Atlantic Ocean, (i.H mm.

Figure 40.— Larvae of Pseudogramma gregoryi from the northwestern Atlantic Ocean: a) 4.7 mm; b) 6.1 mm; c) 6.1 mm; d) 10.2

mm.

Table 8. — Meristic character development of larvae of Pseudogramma gregoryi. Specimens
between dashed lines are undergoing notochord flexion. For the caudal fin,"P" and "S"
indicate primary and secondary fin rays.

Body Branchio-
length stegal
(mm) rays

Verte-
brae

Pec-
toral
fin

Pel-
vic
fin

Caudal fin
Dorsal Ventral
S P P S

Anal
fin

Dorsal fin
Spines Rays

Gill
rakers

3.3 NL 4

10+2

3.6

4.1
4.1
4.2

10+8 15

10+8 16

10+9 16

4 6
6 6
6 6

4.2 SL
4.3
4.4
4.5
4.6
4.7
5.3
5.6
5.9
6.7
7.8
9.6
10.3

10 + 14
10 + 11
10 + 16
10 + 12
10+14
10 + 16
10 + 16
10+16
10 + 16
10 + 16
10+16
10+16
10+16

16
15
15
15
15
15
15
15
15
15
15
15
15

1,3
3

1,4
1,4
1,5
1,5
[,5
1,5

1,13
9

1,16
10

1,16
111,15
111,16
111,15
111,15
111,15
III, 15
111,15
111,15

VIII

VIII
VIII
VIII
VIII
VIII
VIII
VIII
VIII
VIII

17
18

19
19

18
18
19
19

1 + 7
1 + 7
1 + 7
1 + 5

8
3 + 7

3 + 7
3+8

4 + 8
3+9
3+10

5-mm specimens (branchiostegal rays, vertebrae, pec-
toral rays, anal fin rays, dorsal fin spines and rays, and
primary caudal fin rays). The pelvic fin is notably late in
formation, relative to the other fins, and does not attain
its adult complement of rays until 6.7 mm, after all other
elements have reached their adult numbers. Thus, the
sequence of meristic character development in Pseudo-
gramma gregoryi is quite different from that of other
serranids in that the pectoral fin forms before other fins
and the pelvic fin forms last. The third anal spine
develops as a spine rather than as a ray that transforms
into a spine as it does in other serranids.

Body proportions (Table 9) — A series of larvae from 3.9
to 11.4 mm was measured to trace changes in body pro-
portions during larval development. Due to the limited
number of specimens available, some partly damaged
larvae had to be used for these measurements; therefore,
some measurements may not represent actual body
shape. As the notochord flexes, total length increases
from about 110% to about 125% of BL, caudal peduncle
depth increases from 6-10% to about 14% of BL, and the
preanal length increases from about 45% to about 55% of
BL. The rest of the proportions remain rather constant
over the size range at hand. The body depth is about 25%
of BL (23.2-31.7%) and the depth at the anus is only
slightly less (22.5-29.5%). The measurements associated
with the head are: eye, 8.3 to 11.6% of BL; snout, 4.7 to
10.0% of BL; and head length, 26.6 to 37.8% of BL. Pseu-
dogramma gregoryi then, is shallower bodied and has a
shorter head and preanal length than other serranids.

Pigmentation — Larvae of Pseudogramma gregoryi are
notable, as are those of Liopropoma and Rypticus, for
their lack of pigment. The only body pigment of P.
gregoryi during larval development occurs on the dorso-
lateral surface of the gut cavity. Some pigment develops
on the membrane associated with the elongated second
spine of the dorsal fin and also on the tips of the pectoral

fin rays. Since many specimens were bleached, had
broken dorsal spines or were in otherwise poor condition,
it was not possible to determine exactly when this pig-
ment first appeared. It was only present in larger larvae
among those examined.

Spines in the opercular region (Fig. 41) — Several bones
in the opercular region develop unserrated spines during
the larval period. Their character was observed in a 10.3-
mm larva. The spines are more pointed in P. gregoryi
than in Liopropoma. The preopercular has five subequal
spines on its posterior margin. In contrast to other ser-
ranids where the spine at the preopercular angle is
longest, the spine dorsal to the one at the angle is longest
in P. gregoryi. The interopercular has a sharp spine on its
posterior edge, as does the subopercular. The suboper-
cular also has a strengthening ridge that lies under the
opercular and extends posterodorsally near the opercu-
lar margin. The opercular has three posteriorly directed
spines that are pointed and closer together than those of
other genera examined. The posttemporal and supra-
cleithrum lack noticeable spines protruding from the lar-
val surface.

Discussion. — Larvae of the three genera described
here (Pseudogramma gregoryi, Liopropoma, and Ryp-
ticus) share characters which unite them with each other
and separate them from other groups of serranid larvae.

1. They are practically devoid of pigment at all sizes.
Some Liopropoma larvae have one to three melano-
phores on the caudal peduncle and the fleshy parts of the
elongated dorsal spines are pigmented. Nevertheless,
these genera have markedly less pigment than other
serranid genera I examined.

2. They all possess one or two elongate, thin, dorsal
spines with fleshy sheaths. These are broken in many

Table 9. — Body proportions of larvae of Pseudogramma gregoryi. Specimens
between dashed lines are undergoing notochord flexion.

Percent of standard length

Body

Pre-

Caudal

Depth

length

Total

Body

Eye

Head

Snout

anal

peduncle

(mm)

length

depth

length

depth

anus

3.9NL

111

27.4

9.7

30.6

6.5

37.1

6.5

24.2

4.1

106

29.2

9.2

29.2

7.7

46.2

9.2

26.2

4.3

110

23.2

10.1

30.4

8.7

44.9

10.1

23.2

4.3

110

25.0

10.2

30.9

8.8

44.1

8.8

25.0

4.3

112

26.1

11.6

30.4

8.7

46.4

10.1

26.1

4.6

112

25.7

9.5

31.1

—

50.0

9.5

23.0

4.5 SL

120

26.8

11.3

32.4

7.0

52.2

14.1

26.8

4.5

118

23.9

9.9

29.6

9.9

46.5

11.3

22.5

4.8

113

27.6

10.5

27.5

6.6

42.1

13.2

26.3

4.8

116

27.6

9.2

27.6

—

46.0

13.2

26.3

4.8

118

26.3

9.2

30.3

9.2

50.0

14.5

23.7

5.0

118

26.3

10.0

31.3

10.0

47.5

12.5

23.8

5.2

117

31.7

9.8

37.8

9.8

59.8

11.0

24.4

5.3

112

27.4

10.7

31.0

7.1

50.0

16.7

26.2

5.7

122

26.4

9.9

29.7

7.7

50.5

15.4

25.3

5.9

119

26.6

8.5

26.6

5.3

46.8

13.8

23.4

6.0

121

26.0

10.4

34.4

8.3

45.8

14.6

25.0

6.1

120

26.8

9.3

29.9

8.2

48.5

15.5

25.8

6.0

119

25.0

8.3

28.1

6.3

44.8

12.5

24.0

7.2

121

26.7

9.5

29.5

6.7

50.5

16.2

27.6

8.0

118

25.2

9.4

29.9

4.7

50.4

14.2

27.6

9.7

125

26.0

8.4

31.8

9.1

53.2

13.0

27.3

9.8

125

6.3

9.0

28.8

7.1

53.8

13.5

28.2

10.4

122

28.3

9.0

30.7

6.6

56.0

14.5

29.2

11.4

124

25.4

8.8

32.0

7.7

53.6

14.4

26.5

posttemporal

supracleithrum

opercula

preopercula

subopercular

interopercular

Figure 41. — Spine-bearing bones of the opercular and posttemporal
regions on a lll..'1-nim Pseudogramma gregoryi larva.

field-collected specimens; but the bases of these elon-
gate spines are larger than those of other spines and this
is useful in recognizing these larvae.

3. The pectoral fins develop rays earlier and are larger
than in other serranids. In other serranid genera the pec-

toral fins form last; however, in these genera (except Lio-
propoma) they form while only a few dorsal spines and
caudal fin rays are developing.

4. The body shape is little elevated and the caudal pe-
duncle is deep, producing a tubular-shaped body similar
to that seen in some labrid and scarid larvae. Serranid
larvae in the subfamilies Anthiinae and Epinephelinae
are deeper bodied than grammistines. Only larvae of
serranines have the depressed condition of grammistines.

These three genera have been considered members of
separate families or subfamilies; but on the basis of lar-
val morphology and other characters (Kendall 1976),
they appear more closely related to each other. Several
other genera have been aligned with those described here
but their larvae are unknown, so an accurate assessment
of the relationships within this group based on larval de-
velopment is not possible at present. However, Hubbs
and Chu (1934) illustrated late larvae of Diploprion bi-
fasciatum, a genus considered a grammistid by Randall
et al. (1971). Their specimens had greatly elongated flex-
ible second and third dorsal spines. Jeboehlkia gladifer,
described from a small specimen by Robins (1967), has
an elongated but stiff first dorsal fin spine and is prob-
ably a small specimen of a species closely related to those
described here. Fourmanoir (1976) illustrated and briefly
described larvae of Grammistes sexilineatus and
Aporops bilinearis, both with an elongate dorsal spine
and preopercular spines similar to the grammistines de-
scribed here. The unique first pterygiophore of gram-
mistine and related fishes (Kendall 1976) may help sup-
port the elongate larval dorsal spines, as the enlarged

predorsal bone supports the vexillum of larval carapids
(Strasburg 1965). Possibly all fishes belonging in the
Grammistinae have one or two elongated dorsal spines as
larvae or have lost them secondarily. As larvae of more
members of this group are described, relationships be-
tween them may become clarified.

DISCUSSION

This study of larvae of most American serranid genera
permits an examination of the phylogeny of the group on
the basis of larval characters. Among the percichthyids,
only the larvae of Morone have been adequately de-
scribed (Mansueti 1958, 1964). Since they develop in es-
tuarine areas whereas serranids develop at sea, their
morphology may show adaptations not present in ser-
ranids. Nevertheless, the larvae of Morone appear
generalized and it appears that unspecialized serranines
could have been derived from them. Some of the larval
characters of Morone considered trenchant in this
respect are their late-developing spinous dorsal and
pelvic fins, pigment consisting of evenly sized melano-
phores scattered over the body, and the lack of heavy ar-
mature in the opercular area.

Serraninae larvae demonstrate various degrees of
specialization from the Morone pattern. There seems to
be a trend among the genera toward a deeper body,
earlier development of the spinous dorsal and pelvic fins,
larger spines on bones in the opercular region, and pig-
ment concentrated in blotches mainly along the ventral
midline. Hypoplectrus and a type here designated
Diplectrum Type 2 larvae do not show these features, so
their taxonomic position based on other characters
should be reevaluated.

Among the other serranids there seems two major lines
of divergence from the serranines. These are the
anthiines and the epinepheline-grammistines (Kendall
1976).

Four genera of anthiine larvae (Plectranthias, Anthias,
Pronotogrammus, and Hemanthias) were recognized and
among them there appears to be a trend toward in-
creased development of strong spines in the opercular
region and on the head. Also fin spines in the dorsal and
pelvic fins become strong. The strong ctenoid scales
develop precociously. Serrations develop on the spines in
the opercular region and on the thick fin spines. Among
the anthiine genera and among the species within the
genera these features are variously developed. These
features probably afford protection against predation
and thus make possible a longer planktonic larval period
to increase the dispersion of these fish.

The American epinephelines contain three genera.
One of these genera, Epinephelus, has several subgenera.
Although there are considerable differences in size and
body shape of adults of members of this subfamily, indi-
cating different ecological requirements, the larvae are
similar. Several species of epinepheline larvae have been
described from other parts of the world and these are
similar to American larvae. In fact, no characters were
found to separate larval epinephelines to genera. Epi-

nepheline larvae possess a suite of specialized characters
that apparently enable them to have a long pelagic ex-
istence. They have extremely elongate and serrate sec-
ond dorsal and pelvic spines. The preopercular also has a
long serrate spine at its angle and there is a serrate supra-
orbital crest. Pigment is generally confined to the ventral
area of the caudal peduncle, the hindbrain, the lateral
surface of the gut cavity, and the membranes of the elon-
gate spines. Gonioplectrus larvae resemble epinepheline
larvae in most respects, differences being in "degree"
rather than "kind" (Kendall and Fahay in press).
Whether Gonioplectrus should be considered an epi-
nepheline awaits further analysis of adult morphology.
Since there are no other larvae with only some of the epi-
nepheline suite of characters or degrees of development
of the characters, it is not possible to relate these fish to
any others on the basis of their larvae. However, other
evidence indicates that the epinephelines arose from
serranine-type fishes (Smith 1971).

Larvae representing three genera in the grammistine
line were found. No consensus on the relationship of
these three genera is available; but their predorsal bone
patterns (Kendall 1976) as well as larval similarity indi-
cate that they are part of a single lineage. The most out-
standing feature of these larvae is the development of one
or two elongate, flexible, dorsal spines. Each spine has a
variously pigmented membranous sheath around it.
Otherwise, the larvae are practically devoid of pigment.
In several features of development, these fish are similar
to serranines. Their body shape is little elevated and
there are no strong spines associated with the head or
fins. The sequence of fin development is similar to that in
serranines, except the pectoral fin develops early in Pseu-
dogramma. In summary, it appears that these fishes
could have developed from fishes with serranine-like lar-
vae and that epinepheline larval specializations occurred
in that lineage after the grammistine line diverged from
it. Based on predorsal bone patterns, Pogonoperca is
intermediate between serranines and epinepheline-
grammistines (Kendall 1976) so its larvae may show par-
tial development of epinepheline-grammistine
characters.

SUMMARY

There are four distinct groups among the American
serranids based on the morphology of their larvae. These
groups represent the subfamilies Serraninae, Anthiinae,
and Epinephelinae and the genera Liopropoma, Ryp-
ticus, and Pseudogramma, here grouped as the subfami-
ly Grammistinae. Within the serranines and anthiines,
larvae of several genera were found and a progression of
morphological characters which seem to make the larvae
better fit for a longer planktonic existence was seen.

The four groups of American serranid larvae can be
partially characterized as follows (Figs. 42, 43):

Serraninae — Body proportions show rather direct de-
velopment. There are no elongate spines in the opercu-
lar region, rather a series of blunt points. The fin spines
are thin and only slightly elongated in some. Most larval

Figure 42. — Larvae representative of four groups of American Serranidae: a) Serraninae (Paralabrax sp., 6.0 mm); b) Anthiinae (Anthias
tenuis, 6.7 mm); c) Epinephelinae (Mycteroperca mierolepis, 7.4 mm); d) Grammistinae (Rypticus sp., 6.6 mm).

Figure 43. — Interoperculars of larvae of representatives of four
groups of American Serranidae: a) Serraninae (Centropristis
striata, 10.6 mm); b) Anthiinae (Hemanthias vivanus, 10.3 mm);
c) Epinephelinae (Epinephelus niveatus, 10.2 mm); d) Grammistinae
(Pseudogramma gregoryi, 10.3 mm).

pigment consists of melanophores in characteristic posi-
tions along the ventral midline.

Anthiinae — These are deep-bodied larvae with pro-
duced spines on several bones in the opercular region,
some of which are serrated. The pelvic and some dorsal
fin spines are strong, and serrate in some, but not very
elongate. Pigment consists mainly of large blotches and
dashes in characteristic positions on the trunk.

Epinephelinae — The body is roughly "kite" shaped,
deepest at insertion of second spine of the dorsal fin and
pelvic fin spine. Elongate serrate spines are present on
the preopercular, but there are no serrations on other
bones in the opercular series. The second dorsal fin spine
and pelvic fin spine are greatly elongate and heavily ser-
rate. Pigment mainly consists of a large blotch on the
caudal peduncle, heavy pigment over the body cavity,
and some pigment on the membranes associated with the
produced fin spines.

Grammistinae — The body is roughly tubular with a
deep caudal peduncle. Bones in the opercular series are
armed with variously elongated, simple spines. One or
two dorsal fin spines become quite elongate, but are thin
and flexible with pigmented membranous sheaths
around them. Bodies of the larvae are practically devoid
of pigment throughout development.

ACKNOWLEDGMENTS

This work was a thesis project at Scripps Institution of
Oceanography. Elbert H. Ahlstrom and Richard H.
Rosenblatt provided much invaluable advice and
guidance during this study.

I thank the following people for their help in obtaining
specimens in collections under their control and for

engaging in useful discussions about this work: Phillip C.
Heemstra, CSIRO, Cronulla, N.S.W., Australia; C.
Lavett Smith, American Museum of Natural History;
Leslie Knapp, Smithsonian Oceanographic Sorting
Center; and William J. Richards, National Marine
Fisheries Service, Miami, Fla., who also reviewed the
manuscript. L. A. Walford, New Jersey Marine Science
Consortium, and H. Geoffrey Moser, National Marine
Fisheries Service, La Jolla, gave much encouragement
and helpful advice.

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1963. Patterns of the ramus lateralis accessorius and their
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Appendix Table 1. — Data associated with larval fish used for illustrations. Institution abbreviations: NE: NMFS, Northeast
Fisheries Center; SW: NMFS, Southwest Fisheries Center; SE: NMFS, Southeast Fisheries Center; UNC: University of North
Carolina.

Figure

Species

Length
(mm)

Insti-
tution

Cruise

Station

Lat.
(N)

Long. Date

(W) Day/Mo/Yr

la
b

Centropristis striata
Centropristis striata

Centropristis striata

4.7

D-67-4

FF-5(D

31°38'

79°56'

5 67

8.3

D-71-5

N-2p

34°56'

75°55'

4 71

0.6

D-67-4

CC-5U)

32°20'

78°25'

5 67

3a
b

Faralabrax sp.
Paralabrax sp.

6.0

9.0

6509B
S-5509

118.39 28°19' 115°24' 17 10 65

(CalCOFI sample — no precise data)

4 Paralabrax sp. 9.5 SW C-51

5a Diplectrum sp. Type 1 (Atl.)

b Diplectrum sp. Type 1 (Atl.)

6a Diplectrum sp. Type 1 (Pac.)

b Deplectrum sp. Type 1 (Pac.)

c Diplectrum sp. Type 1 (Pac.)

d Diplectrum sp. Type 1 (Pac.)

7 Diplectrum sp. Type 1 (Atl.) 10.3 NE D-67-4

8a Diplectrum sp. Type 2

b Diplectrum sp. Type 2

9 Diplectrum sp. Type 2 9.9 SW ETP

5.8

N E

D-67-4

10.0

D-67-16

4.3

5706S

5.0

5706S

6.1

5706S

13.0

5706S

130.30

26°21'

113°49'

GG-4U)

31<W

80°51'

HH-5(2)

30°22'

80°48'

107G02

29°42'

114°07'

107G02

29°42'

114°07'

107G02

29°42'

114°07'

143G100

29°19'

109°14'

CC-5(1)

33°20'

78°25' 14 5 67

4.5

P-5412

157.10

22°35'

109°19'

7 54

8.4

5612-H

160G30

22°39'

107°49'

7 56

47.244

6°59'

1°54' 27 8 67

10a
b

Serranus sp.
Serranus sp.
Serranus sp.
Serranus sp.

Serranus sp.

3.7

D-71-5

M-6

35°07'

74°55'

.VI)

35°03'

75°10'

5.5

D-66-3

M-5(2)

35°11'

75°07'

9.4

D-68-1

NN-2(2)

27°11'

80°03'

9.7

D-66-12

L-4(l)

35°46'

75°05'

12a
b

Hypoplectrus sp.
Hypoplectrus sp.
Hypoplectrus sp.
Hypoplectrus sp.

3.4

4.7

5.7

8.5

Reared at NMFS, Miami, Fla. from eggs collected nearby
Reared at NMFS, Miami, Fla. from eggs collected nearby
Reared at NMFS, Miami, Fla. from eggs collected nearby
Reared at NMFS, Miami, Fla. from eggs collected nearby

13 Hypoplectrus sp.

14a Serraniculus pumilio

b Serraniculus pumilio

c Serraniculus pumilio

15a Plectranthias garupellus

b Plectranthias garupellus

16a Anthias sp. Type 1

b Anthias sp. Type 1

17a Anthias sp. Type 2

b Anthias sp. Type 2

c Anthias sp. Type 2

18 Anthias tenuis

19 Anthias sp. Type 3

20a Anthias gordensis

b Anthias gordensis

21 Anthias sp. Type 1

12.5

Reared at NMFS, Miami, Fla. from eggs collected nearby

3.8

D-67-16

DD-4U)

32°55'

78°51'

5.8

D-67-16

DD-5(2)

32°47'

78°44'

55.3

D-71-3

SA-2

29°51'

81°10'

5.5

D-67-16

NN-4(1)

27°12'

79°51'

7.0

D-68-1

PP-2U)

26°47'

79°56'

3.8

D-66-5

N-4(2)

34°42'

75°48'

5.3

D-71-5

BB-7

32°53'

77°26'

4.7

D-68-1

DD-6(1)

32°36'

78°34'

5.7

D-71-5

M-4

35°13'

75°12'

8.4

D-68-1

EE-6

32°08'

79°10'

6.7

BLM

IV-2

26°10'

96°39'

5.1

D-71-5

BB-8

32°39'

77°17'

5.2

5706S

109G55

29°52'

113°04'

6.0

5706S

115G40

28°52'

112°44'

10.2

D-68-1

EE-6

32°08'

79°10'

Appendix Table 1.— Continued.

Figure

Species

Length
(mm)

Insti-
tution

Cruise

Station

Lat.

(N)

Long.

(W)

Date

Day/Mo/Yr

22a Pronotogrammus aureorubens

b Pronotogrammus aureorubens

c Pronotogrammus aureorubens

23a Pronotogrammus eos

b Pronotogrammus eos

24 Pronotogrammus aureorubens

4.6

D-68-1

PP-3(1)

26°47'

79°50'

6.0

D-68-1

CC-7(2)

32°54'

78°07'

9.0

D-68-1

AA-7

33°37'

76°47'

3.6

5706S

133G40

26°16'

110°50'

7.8

052

00°04'

84°58'

9.5

D-68-1

HH-7

30°19'

313'

1 2

25a
b

Hemanthias viuanus
Hemanthias uiuanus

Hemanthias uiuanus

Hemanthias peruanus

Hemanthias uiuanus

29a
b
c

Hemanthias vwanus
Hemanthias uiuanus
Hemanthias uiuanus

Hemanthias uiuanus

Paranthias furcifer

Epinephelus striatus

Epinephelus sp.

Epinephelus niueatus

35a
b
c
d

Mycteroperca microlepis
Mycteroperca microlepis
Mycteroperca microlepis
Mycteroperca microlepis

Mycteroperca microlepis

37a
b

Liopropoma sp.
Liopropoma sp.

Liopropoma sp.

Rypticus sp.

40a
b
c
d

Pseudogramma gregoryi
Pseudogramma gregoryi
Pseudogramma gregoryi
Pseudogramma gregoryi

41 Pseudogramma gregoryi

42a Paralabrax sp.

b Anthias tenuis

c Mycteroperca microlepis

d Rypticus sp.

43a Centropristis striata

b Hemanthias uiuanus

c Epinephelus niueatus

d Pseudogramma gregoryi

4.2

D-71-5

LK-5

35°59'

74°33'

5.3

D-71-5

BB-8

32°39'

77°17'

6.8

D-71-5

BB-8

32°39'

77°17'

9.3

6611J

147.30

23°36'

111°42'

9.3

D-71-5

N-7

34°13'

75°35'

10.3

D-68-1

HH-7

30°19'

80°13'

4.3

D-71-5

N-7

34°13'

75°35'

5.8

D-71-5

N-6

34°23'

75°39'

6.6

D-71-5

N-7

34°13'

75°35'

7.2

BLM-2

1-3 Day

27°34'

96°07'

8.6

5612

157G10

22°30'

10915'

7.6

D-67-4

GG-7(2)

30°54'

80°00'

8.4

162

11°10'

92°00'

10.2

D-67-16

GG-7

30°54'

80°00'

4.0

35°03'

75°10'

7.4

35°03'

75°08'

14.2

UNC

T-9

New Rivei

,N.C.

22.6

UNC

(UNC 7934)

Bogue Sd

, N.C.

9.8

D-71-5

AA-7

33°37'

76°47'

6.3

D-67-4

NN-4(2)

27°12'

79°51'

7.0

35°03'

75°08'

12.0

OR II 702

0-2-13

26°47'

84°34'

6.6

D-67-8

AA-4U)

34°11'

77°11'

4.7

D-67-4

MM-4(1)

27°46'

79°56'

6.1

OR II 7739

20°00'

80°13'

6.1

D-66-3

N-5U)

34°33'

75°44'

10.2

D-67-4

JJ-6U)

29°49'

80°14'

10.3

OR II 7239

20°00'

80°13'

6.0

s\\

6509B

118.39

28°19'

115°24'

6.7

BLM

IV-2

26°10'

96°39'

7.4

35°03'

75°08'

6.6

D-67-8

AA-4(1)

34°11'

77°11'

10.6

D-67-4

CC-5U)

33°20'

78°25'

10.3

D-68-1

HH-7

30°19'

80° 13'

10.2

D-67-16

GG-7

30°54'

80°00'

10.3

OR U 7239

20°00'

80°13'

NOAA TECHNICAL REPORTS
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Full text of "Morphological comparisons of North American sea bass larvae (Pisces: Serranidae) / Arthur W. Kendall, Jr"

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