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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 
 
NOAA TECHNICAL REPORTS 
 National Marine Fisheries Service, Circulars 
 
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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 
 
 24 
 25 
 26 
 
 26 
 
 26 
 28 
 29 
 31 
 
 32 
 33 
 
 34 
 36 
 
 38 
 40 
 
 41 
 41 
 42 
 
 44 
 46 
 47 
 
 3 
 
 14 
 15 
 27 
 28 
 37 
 38 
 43 
 44 
 
 IV 
 
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- 
 
 2 W. 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 
 (Kendall 4 ). 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- 
 
 J A. 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 
 
 10 
 
a 
 
 b 
 
 d 
 
 
 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. 
 
 11 
 
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 
 
 12 
 
a 
 
 b 
 
 ?-- T*s?3r 
 
 rk&^$>. 
 
 'MM 
 
 v'^-vSW ^ '-% ra$* 
 
 
 
 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 
 
 L3 
 
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 
 
 3.5 
 
 3.6 
 
 3.6 
 
 3.6 
 
 3.7 
 
 3.8 
 
 3.8 
 
 3.8 
 
 3.8 
 
 :■; S 
 
 3.8 
 
 3.8 
 
 4.0 
 
 4.0 
 
 4.0 
 
 4.3 
 
 4.4 
 
 1 
 3 
 
 4 
 
 8 
 
 5 
 
 11 
 
 11 
 
 10 
 
 15 
 
 in 
 
 8 
 
 3 
 
 3 
 
 16 
 
 14 
 
 12 
 
 10 
 
 16 
 
 13 
 
 14 
 
 3.8 
 4.1 
 4.2 
 4.3 
 3.9 
 
 16 
 
 17 
 17 
 17 
 
 is 
 
 1 
 12 
 
 1,2 
 
 VI 
 
 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 
 
 7.0 
 
 7.0 
 
 7.2 
 
 7.3 
 
 19 
 
 22 
 24 
 22 
 24 
 24 
 24 
 21 
 24 
 24 
 23 
 24 
 24 
 24 
 24 
 24 
 
 13 
 
 15 
 
 14 
 
 9 
 
 14 
 
 15 
 
 15 
 
 6 
 
 9 
 
 15 
 
 8 
 
 8 
 
 13 
 
 14 
 
 15 
 
 14 
 
 1,3 
 
 1,3 
 
 [,5 
 
 I 
 
 1,5 
 
 1,5 
 
 1,5 
 
 1.4 
 
 1,5 
 
 1,5 
 
 1,4 
 
 1,4 
 
 1,5 
 
 1,5 
 
 1,5 
 
 1,5 
 
 
 H,7 
 
 VIII 
 
 9 
 
 1 
 
 1 
 
 111,7 
 
 IX 
 
 9 
 
 2 
 
 2 
 
 111,7 
 
 X 
 
 11 
 
 4 
 2 
 
 3 
 
 111,7 
 
 X 
 
 in 
 
 3 
 
 2 
 
 111,7 
 
 X 
 
 11 
 
 5 
 
 4 
 
 111,7 
 
 X 
 
 11 
 
 5 
 
 
 II. 1 
 
 IV 
 
 
 4 
 
 
 11,1 
 
 VIII 
 
 
 5 
 
 5 
 
 111,7 
 
 X 
 
 10 
 
 5 
 
 
 11,1 
 
 VII 
 
 
 5 
 
 
 11,1 
 
 VII 
 
 
 4 
 
 
 III 
 
 IV 
 
 
 4 
 
 
 111,7 
 
 X 
 
 11 
 
 5 
 
 6 
 
 IH,7 
 
 X 
 
 11 
 
 6 
 
 6 
 
 111,7 
 
 X 
 
 11 
 
 6 
 
 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 
 
 14 
 
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 
 
 at 
 
 (mm) 
 
 length 
 
 depth 
 
 length 
 
 length 
 
 length length 
 
 depth 
 
 anus 
 
 2.9 NL 
 
 103 
 
 32 
 
 13.0 
 
 39 
 
 10.0 58 
 
 5.7 
 
 19 
 
 3.7 
 
 104 
 
 31 
 
 11.0 
 
 38 
 
 12.0 59 
 
 8.0 
 
 25 
 
 3.9 
 
 107 
 
 33 
 
 12.0 
 
 38 
 
 10.0 63 
 
 10.0 
 
 27 
 
 4.0 
 
 107 
 
 34 
 
 11.0 
 
 39 
 
 11.0 61 
 
 9.8 
 
 27 
 
 4.0 SL 
 
 115 
 
 33 
 
 11.0 
 
 36 
 
 12.0 61 
 
 11.0 
 
 28 
 
 4.2 
 
 118 
 
 34 
 
 12.0 
 
 39 
 
 10.0 63 
 
 13.0 
 
 28 
 
 4.2 
 
 115 
 
 36 
 
 12.0 
 
 42 
 
 11.0 62 
 
 12.0 
 
 30 
 
 4.4 
 
 119 
 
 33 
 
 11.0 
 
 39 
 
 10.0 63 
 
 14.0 
 
 29 
 
 5.5 
 
 117 
 
 31 
 
 10.0 
 
 38 
 
 10.0 66 
 
 14.0 
 
 27 
 
 5.7 
 
 122 
 
 31 
 
 11.0 
 
 38 
 
 11.0 59 
 
 13.0 
 
 25 
 
 6.0 
 
 123 
 
 31 
 
 11.0 
 
 38 
 
 11.0 62 
 
 15.0 
 
 26 
 
 6.1 
 
 123 
 
 29 
 
 10.0 
 
 36 
 
 9.3 65 
 
 14.0 
 
 29 
 
 6.7 
 
 117 
 
 29 
 
 9.4 
 
 37 
 
 9.4 62 
 
 14.0 
 
 26 
 
 6.8 
 
 120 
 
 31 
 
 11.0 
 
 38 
 
 9.0 67 
 
 14.0 
 
 J 7 
 
 7.0 
 
 120 
 
 29 
 
 9.8 
 
 36 
 
 9.8 65 
 
 14.0 
 
 25 
 
 7.7 
 
 125 
 
 23 
 
 9.8 
 
 32 
 
 7.4 61 
 
 13.0 
 
 28 
 
 7.8 
 
 124 
 
 29 
 
 10.0 
 
 39 
 
 10.0 64 
 
 15.0 
 
 26 
 
 7.9 
 
 117 
 
 29 
 
 11.0 
 
 38 
 
 9.5 64 
 
 14.0 
 
 25 
 
 7.9 
 
 124 
 
 28 
 
 10.0 
 
 39 
 
 9.5 62 
 
 13.0 
 
 25 
 
 8.3 
 
 125 
 
 27 
 
 9.9 
 
 39 
 
 9.9 66 
 
 14.0 
 
 26 
 
 Adult 1 
 
 
 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 
 
 L5 
 
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. 
 
 6 P. 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 
 
 L6 
 
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 
 
 17 
 
a 
 
 b 
 
 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). 
 
 18 
 
b 
 
 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. 
 
 19 
 
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 
 
 20 
 
a 
 
 b 
 
 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 
 
 21 
 
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 12 c ? 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. 
 
 22 
 
a 
 
 b 
 
 c 
 
 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 
 
 23 
 
b 
 
 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. 
 
 24 
 
a 
 
 b 
 
 Figure 25.— Larvae of Hemanthias vivanus from the northwestern Atlantic Ocean: a) 4.2 mm; b) 5.3 mm; c) 6.8 mm. 
 
 25 
 
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 
 
 26 
 
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 
 
 3.6 
 
 3.7 
 
 3.7 
 
 3.7 
 
 3.7 
 
 3.8 
 
 3.8 
 
 3.8 
 
 3.8 
 
 3.9 
 
 3.9 
 
 3.9 
 
 4.0 
 
 4.0 
 
 4.1 
 
 4.1 
 
 4.1 
 
 4.1 
 
 4.2 
 
 4.2 
 
 4.2 
 
 4.2 
 
 4.3 
 
 4.4 
 
 4.4 
 
 4.4 
 
 4.5 
 
 4.7 
 
 4.7 
 
 4.8 
 
 4.9 
 
 4.9 
 
 4.9 
 
 4.9 
 
 4.9 
 
 4.9 
 
 5.0 
 
 5.0 
 
 5.2 
 
 5.3 
 
 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 
 
 3 
 
 damage 
 
 5.5 
 
 15 
 
 11 
 
 IX 
 
 5.6 SL 
 
 5.6 
 
 5.7 
 
 5.8 
 
 5.8 
 
 5.9 
 
 6 I 
 
 6.3 
 
 6.3 
 
 6.3 
 
 6.4 
 
 6.5 
 
 6.5 
 
 6.8 
 
 6.8 
 
 6.8 
 
 7.0 
 
 20 
 14 
 18 
 16 
 16 
 15 
 14 
 18 
 18 
 23 
 18 
 20 
 22 
 23 
 20 
 26 
 18 
 
 II 
 
 14 
 
 II 
 
 II 
 
 II. 1 
 
 11,1 
 
 II 
 
 II 
 
 II 
 
 11,1 
 
 II 
 
 11.9 
 
 II. 1 
 
 11,1 
 
 [1,9 
 
 11,1 
 11,1 
 
 11,9 
 11,1 
 
 IX 
 VIII 
 
 IX 
 
 IX 
 VIII 
 
 IX 
 VIII 
 X 
 X 
 X 
 X 
 
 IX 
 X 
 X 
 X 
 X 
 X 
 
 14 
 
 12 
 
 8 
 9 
 10 
 
 11 
 
 11 
 
 13 
 
 9 
 
 27 
 
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 
 
 fin 
 
 S 
 
 P 
 
 P 
 
 S 
 
 fin 
 
 Spines 
 
 Rays 
 
 rakers 
 
 '7.1 
 
 7 
 
 26 
 
 14 
 
 1,5 
 
 
 8 
 
 7 
 
 
 11,9 
 
 X 
 
 14 
 
 13 
 
 '7.3 
 
 7 
 
 26 
 
 16 
 
 1,5 
 
 
 9 
 
 8 
 
 
 11,9 
 
 X 
 
 14 
 
 14 
 
 '7.3 
 
 7 
 
 23 
 
 2 
 
 1,2 
 
 
 6 
 
 4 
 
 
 n,i 
 
 X 
 
 
 12 
 
 '7.7 
 
 7 
 
 26 
 
 17 
 
 1,5 
 
 
 9 
 
 8 
 
 
 11,9 
 
 X 
 
 14 
 
 15 
 
 '7.7 
 
 7 
 
 26 
 
 17 
 
 1,5 
 
 
 9 
 
 8 
 
 
 11,9 
 
 X 
 
 14 
 
 15 
 
 '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 
 
 at 
 
 (mm) 
 
 length 
 
 depth 
 
 length 
 
 length 
 
 length 
 
 length 
 
 depth 
 
 anus 
 
 3.1 NL 
 
 106 
 
 36 
 
 14 
 
 34 
 
 12 
 
 60 
 
 8 
 
 28 
 
 4.0 
 
 103 
 
 29 
 
 10 
 
 35 
 
 11 
 
 57 
 
 5 
 
 :\2 
 
 4.4 
 
 114 
 
 39 
 
 11 
 
 37 
 
 14 
 
 53 
 
 11 
 
 31 
 
 4.6 
 
 108 
 
 36 
 
 11 
 
 41 
 
 15 
 
 58 
 
 10 
 
 27 
 
 4.6 
 
 112 
 
 38 
 
 12 
 
 42 
 
 15 
 
 55 
 
 10 
 
 30 
 
 4.6 
 
 109 
 
 36 
 
 12 
 
 39 
 
 14 
 
 57 
 
 11 
 
 27 
 
 4.6 
 
 109 
 
 38 
 
 12 
 
 42 
 
 14 
 
 59 
 
 11 
 
 27 
 
 4.7 SL 
 
 116 
 
 41 
 
 15 
 
 43 
 
 16 
 
 61 
 
 12 
 
 35 
 
 5.0 
 
 116 
 
 38 
 
 13 
 
 39 
 
 13 
 
 58 
 
 11 
 
 30 
 
 5.0 
 
 113 
 
 41 
 
 15 
 
 48 
 
 15 
 
 60 
 
 14 
 
 38 
 
 5.2 
 
 116 
 
 37 
 
 12 
 
 40 
 
 12 
 
 55 
 
 12 
 
 31 
 
 5.2 
 
 118 
 
 39 
 
 13 
 
 41 
 
 1(1 
 
 60 
 
 13 
 
 31 
 
 5.4 
 
 123 
 
 40 
 
 15 
 
 43 
 
 15 
 
 63 
 
 14 
 
 30 
 
 5.5 
 
 117 
 
 44 
 
 16 
 
 43 
 
 15 
 
 62 
 
 14 
 
 36 
 
 5.7 
 
 121 
 
 38 
 
 13 
 
 40 
 
 15 
 
 58 
 
 L3 
 
 31 
 
 6.0 
 
 118 
 
 43 
 
 16 
 
 46 
 
 13 
 
 65 
 
 16 
 
 35 
 
 6.0 
 
 122 
 
 42 
 
 14 
 
 44 
 
 15 
 
 60 
 
 14 
 
 36 
 
 6.3 
 
 122 
 
 39 
 
 13 
 
 44 
 
 13 
 
 60 
 
 14 
 
 31 
 
 6.5 
 
 124 
 
 40 
 
 14 
 
 42 
 
 14 
 
 61 
 
 14 
 
 32 
 
 6.8 
 
 117 
 
 33 
 
 13 
 
 38 
 
 14 
 
 (11 
 
 13 
 
 29 
 
 7.0 
 
 124 
 
 42 
 
 14 
 
 43 
 
 11 
 
 61 
 
 14 
 
 32 
 
 7.8 
 
 124 
 
 37 
 
 14 
 
 42 
 
 14 
 
 59 
 
 13 
 
 29 
 
 8.0 
 
 113 
 
 40 
 
 11 
 
 38 
 
 11 
 
 54 
 
 11 
 
 30 
 
 8.1 
 
 122 
 
 40 
 
 15 
 
 41 
 
 14 
 
 60 
 
 11 
 
 32 
 
 9.2 
 
 121 
 
 35 
 
 12 
 
 37 
 
 14 
 
 55 
 
 12 
 
 28 
 
 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. 
 
 28 
 
Figure 30.— Details of the head spination on a Hemanthias vivanus larva, 7.2 mm. 
 
 29 
 
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 
 
 30 
 
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 
 
 31 
 
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. 
 
 32 
 
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- 
 
 33 
 
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 
 
 34 
 
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. 
 
 35 
 
a 
 
 b 
 
 36 
 
d 
 
 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 
 
 fin 
 
 S 
 
 P 
 
 P 
 
 S 
 
 fin 
 
 Spines 
 
 Rays 
 
 rakers 
 
 4.0 NL 
 
 3 
 
 
 
 1,1 
 
 
 
 
 
 
 UI 
 
 
 2 
 
 4.6 
 
 4 
 
 
 
 1,2 
 
 
 
 
 
 
 III 
 
 
 3 
 
 5.1 
 
 5 
 
 4 
 
 
 1,2 
 
 
 1 
 
 2 
 
 
 
 III 
 
 
 6 
 
 5.6 
 
 6 
 
 6 
 
 
 t,3 
 
 
 5 
 
 5 
 
 
 
 III 
 
 
 7 
 
 5.3 
 
 5 
 
 4 
 
 
 1,3 
 
 
 4 
 
 4 
 
 
 
 III 
 
 
 6 
 
 5.3 
 
 6 
 
 6 
 
 2 
 
 1,3 
 
 
 6 
 
 6 
 
 
 
 IV 
 
 
 8 
 
 5.3 
 
 6 
 
 8 
 
 2 
 
 1,3 
 
 
 7 
 
 7 
 
 
 
 IV 
 
 
 H) 
 
 5.5 
 
 7 
 
 18 
 
 6 
 
 damage 
 
 9 
 
 8 
 
 
 1,6 
 
 V 
 
 
 9 
 
 6.0 SL 
 
 7 
 
 20 
 
 9 
 
 1,4 
 
 
 9 
 
 8 
 
 1 
 
 11,9 
 
 X 
 
 11 
 
 1+9 
 
 6.1 
 
 7 
 
 19 
 
 6 
 
 1,4 
 
 
 9 
 
 8 
 
 
 1,6 
 
 Yl 
 
 
 9 
 
 7.0 
 
 7 
 
 22 
 
 12 
 
 1,5 
 
 1 
 
 9 
 
 8 
 
 2 
 
 11,11 
 
 X 
 
 15 
 
 1 + 10 
 
 7.4 
 
 7 
 
 24 
 
 12 
 
 1,5 
 
 1 
 
 9 
 
 8 
 
 2 
 
 11,11 
 
 X 
 
 15 
 
 2+ 9 
 
 7.9 
 
 7 
 
 24 
 
 14 
 
 1,5 
 
 3 
 
 9 
 
 8 
 
 3 
 
 rj.,13 
 
 X 
 
 19 
 
 3 + 10 
 
 8.1 
 
 7 
 
 24 
 
 15 
 
 1,5 
 
 3 
 
 9 
 
 8 
 
 2 
 
 n,i2 
 
 X 
 
 18 
 
 3+10 
 
 9.2 
 
 7 
 
 24 
 
 17 
 
 1,5 
 
 4 
 
 9 
 
 8 
 
 5 
 
 11,11 
 
 X 
 
 18 
 
 3+12 
 
 9.8 
 
 7 
 
 24 
 
 16 
 
 1,5 
 
 5 
 
 9 
 
 8 
 
 5 
 
 11,12 
 
 X 
 
 18 
 
 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 
 
 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 
 
 38 
 
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 
 
 7 E. H. Ahlstrom, National Marine Fisheries Service, Southwest Fish- 
 eries Center, La Jolla, CA 92038, class notes, August 1971. 
 
 39 
 
b 
 
 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 
 
 40 
 
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. 
 
 41 
 
a 
 
 b 
 
 d 
 
 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. 
 
 42 
 
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 
 
 15 
 
 
 
 
 I 
 
 2 
 
 3.6 
 
 4.1 
 4.1 
 4.2 
 
 10+8 15 
 
 10+8 16 
 
 10+9 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 
 
 IV 
 
 II 
 
 VIII 
 
 II 
 
 VIII 
 VIII 
 VIII 
 VIII 
 VIII 
 VIII 
 VIII 
 VIII 
 VIII 
 
 17 
 18 
 
 18 
 
 18 
 
 19 
 19 
 
 18 
 
 18 
 18 
 19 
 19 
 
 1 + 7 
 1 + 7 
 1 + 7 
 1 + 5 
 
 8 
 
 7 
 
 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 
 
 43 
 
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 
 
 at 
 
 (mm) 
 
 length 
 
 depth 
 
 length 
 
 length 
 
 length 
 
 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 
 
 41 
 
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 
 
 45 
 
c 
 
 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). 
 
 46 
 
a 
 
 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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 species of the subfamily Pseudogramminae. Ichthyol. Aquarium 
 J. 37:185-194. 
 SMITH, C. L. 
 
 1965. The patterns of sexuality and the classification of serranid 
 fishes. Am. Mus. Novit. 2207, 20 p. 
 
 1971. A revision of the American groupers: Epinephelus and allied 
 genera. Bull. Am. Mus. Nat. Hist. 146:69-241. 
 SMITH. C. L., and E. H. ATZ. 
 
 1969. The sexual mechanism of the reef bass Pseudogramma ber- 
 mudensis and its implications in the classification of the Pseudo- 
 grammidae (Pisces: Perciformes). Z. Morphol. Tiere 65:315-326. 
 
 SMITH, C. L., and P. H. YOUNG. 
 
 1966. Gonad structure and the reproductive cycle of the kelp bass. 
 Paralabrax clathratus (Girard), with comments on the relation- 
 ships of the serranid genus Paralabrax. Calif. Fish Game 52:283- 
 292. 
 
 SPARTA, A. 
 
 1932. Contributo alia conoscenza dello sviluppao post-embrionale 
 
 in Athias sacer Bloch. Rev. Comit. Talassogr. 195, 11 p. 
 1935. Contributo alia conoscenze dello sviluppo nei Percidi. Uova 
 ovariche mature di Epinephelus guaza L. e stadi post-embrionali e 
 larvali di Epinephelus alexandrinus Cuv. e Val. Rev. Comit. 
 Talassogr. 224, 12 p. 
 STRASBURG, D. W. 
 
 1965. Description of the larva and familial relationships of the fish 
 Snyderidia canina. Copeia 1965:20-24. 
 
 THRESHER, R. E. 
 
 1978. Polymorphism, mimicry, and the evolution of the hamlets 
 (Hypoplectrus. Serranidae). Bull. Mar. Sci. 28:345-353. 
 UKAWA, M., M. HIGUCHI, and S. MITO. 
 
 1966. Spawning habits and early life history of a serranid fish, Epi- 
 nephelus akaara (Temminck et Schlegel). Jpn. J. Ichthyol. 
 13:156-161. 
 
 VODJANITZKI, W. A., and I. Z. KAZANOVA. 
 
 1954. Opredelitel pelagicheskikh ikrinok i likinok rub Chernogo 
 Morya [Key for identification of the pelagic eggs and fish larvae 
 in the Black SeaJ. Tr. Vses. Nauchno-Issled. Inst. Morsk. Rybn. 
 Khoz. Okeanogr. 28:240-323. 
 WILSON, H. V. 
 
 1891. The embryology of the sea bass (Serranus atrarius). Bull. 
 U.S. Fish. Comm. 9:209-277. 
 
 48 
 
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 
 
 NE 
 
 D-67-4 
 
 FF-5(D 
 
 31°38' 
 
 79°56' 
 
 12 
 
 5 67 
 
 8.3 
 
 NE 
 
 D-71-5 
 
 N-2p 
 
 34°56' 
 
 75°55' 
 
 29 
 
 4 71 
 
 0.6 
 
 NE 
 
 D-67-4 
 
 CC-5U) 
 
 32°20' 
 
 78°25' 
 
 14 
 
 5 67 
 
 3a 
 b 
 
 Faralabrax sp. 
 Paralabrax sp. 
 
 6.0 
 
 9.0 
 
 SW 
 
 SW 
 
 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 
 
 NE 
 
 D-67-16 
 
 4.3 
 
 SW 
 
 5706S 
 
 5.0 
 
 SW 
 
 5706S 
 
 6.1 
 
 SW 
 
 5706S 
 
 13.0 
 
 SW 
 
 5706S 
 
 130.30 
 
 26°21' 
 
 113°49' 
 
 10 
 
 8 
 
 53 
 
 GG-4U) 
 
 31<W 
 
 80°51' 
 
 U 
 
 5 
 
 67 
 
 HH-5(2) 
 
 30°22' 
 
 80°48' 
 
 24 
 
 10 
 
 67 
 
 107G02 
 
 29°42' 
 
 114°07' 
 
 20 
 
 6 
 
 57 
 
 107G02 
 
 29°42' 
 
 114°07' 
 
 20 
 
 6 
 
 57 
 
 107G02 
 
 29°42' 
 
 114°07' 
 
 20 
 
 6 
 
 57 
 
 143G100 
 
 29°19' 
 
 109°14' 
 
 11 
 
 6 
 
 57 
 
 CC-5(1) 
 
 33°20' 
 
 78°25' 14 5 67 
 
 4.5 
 
 SW 
 
 P-5412 
 
 157.10 
 
 22°35' 
 
 109°19' 
 
 4 
 
 7 54 
 
 8.4 
 
 SW 
 
 5612-H 
 
 160G30 
 
 22°39' 
 
 107°49' 
 
 3 
 
 7 56 
 
 47.244 
 
 6°59' 
 
 1°54' 27 8 67 
 
 10a 
 b 
 
 11 
 
 Serranus sp. 
 Serranus sp. 
 Serranus sp. 
 Serranus sp. 
 
 Serranus sp. 
 
 3.7 
 
 NE 
 
 D-71-5 
 
 M-6 
 
 35°07' 
 
 74°55' 
 
 5 
 
 5 
 
 71 
 
 .VI) 
 
 NE 
 
 1 
 
 4 
 
 35°03' 
 
 75°10' 
 
 11 
 
 4 
 
 72 
 
 5.5 
 
 NE 
 
 D-66-3 
 
 M-5(2) 
 
 35°11' 
 
 75°07' 
 
 20 
 
 4 
 
 66 
 
 9.4 
 
 NE 
 
 D-68-1 
 
 NN-2(2) 
 
 27°11' 
 
 80°03' 
 
 4 
 
 2 
 
 68 
 
 9.7 
 
 NE 
 
 D-66-12 
 
 L-4(l) 
 
 35°46' 
 
 75°05' 
 
 30 
 
 9 
 
 66 
 
 12a 
 b 
 
 Hypoplectrus sp. 
 Hypoplectrus sp. 
 Hypoplectrus sp. 
 Hypoplectrus sp. 
 
 3.4 
 
 SE 
 
 4.7 
 
 SE 
 
 5.7 
 
 SE 
 
 8.5 
 
 SE 
 
 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 
 
 SE 
 
 Reared at NMFS, Miami, Fla. from eggs collected nearby 
 
 3.8 
 
 NE 
 
 D-67-16 
 
 DD-4U) 
 
 32°55' 
 
 78°51' 
 
 21 
 
 Hi 
 
 67 
 
 5.8 
 
 NE 
 
 D-67-16 
 
 DD-5(2) 
 
 32°47' 
 
 78°44' 
 
 21 
 
 10 
 
 67 
 
 55.3 
 
 NE 
 
 D-71-3 
 
 SA-2 
 
 29°51' 
 
 81°10' 
 
 3 
 
 9 
 
 71 
 
 5.5 
 
 NE 
 
 D-67-16 
 
 NN-4(1) 
 
 27°12' 
 
 79°51' 
 
 29 
 
 10 
 
 67 
 
 7.0 
 
 NE 
 
 D-68-1 
 
 PP-2U) 
 
 26°47' 
 
 79°56' 
 
 4 
 
 2 
 
 HH 
 
 3.8 
 
 NE 
 
 D-66-5 
 
 N-4(2) 
 
 34°42' 
 
 75°48' 
 
 24 
 
 5 
 
 66 
 
 5.3 
 
 NE 
 
 D-71-5 
 
 BB-7 
 
 32°53' 
 
 77°26' 
 
 1 
 
 5 
 
 71 
 
 4.7 
 
 NE 
 
 D-68-1 
 
 DD-6(1) 
 
 32°36' 
 
 78°34' 
 
 29 
 
 1 
 
 68 
 
 5.7 
 
 NE 
 
 D-71-5 
 
 M-4 
 
 35°13' 
 
 75°12' 
 
 5 
 
 5 
 
 71 
 
 8.4 
 
 NE 
 
 D-68-1 
 
 EE-6 
 
 32°08' 
 
 79°10' 
 
 30 
 
 1 
 
 68 
 
 6.7 
 
 SE 
 
 BLM 
 
 IV-2 
 
 26°10' 
 
 96°39' 
 
 30 
 
 5 
 
 76 
 
 5.1 
 
 NE 
 
 D-71-5 
 
 BB-8 
 
 32°39' 
 
 77°17' 
 
 1 
 
 5 
 
 71 
 
 5.2 
 
 SW 
 
 5706S 
 
 109G55 
 
 29°52' 
 
 113°04' 
 
 18 
 
 6 
 
 57 
 
 6.0 
 
 SW 
 
 5706S 
 
 115G40 
 
 28°52' 
 
 112°44' 
 
 17 
 
 6 
 
 57 
 
 10.2 
 
 NE 
 
 D-68-1 
 
 EE-6 
 
 32°08' 
 
 79°10' 
 
 30 
 
 1 
 
 68 
 
 49 
 
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 
 
 NE 
 
 D-68-1 
 
 PP-3(1) 
 
 26°47' 
 
 79°50' 
 
 4 
 
 2 
 
 68 
 
 6.0 
 
 NE 
 
 D-68-1 
 
 CC-7(2) 
 
 32°54' 
 
 78°07' 
 
 29 
 
 1 
 
 68 
 
 9.0 
 
 NE 
 
 D-68-1 
 
 AA-7 
 
 33°37' 
 
 76°47' 
 
 27 
 
 1 
 
 68 
 
 3.6 
 
 SW 
 
 5706S 
 
 133G40 
 
 26°16' 
 
 110°50' 
 
 12 
 
 6 
 
 57 
 
 7.8 
 
 SW 
 
 OP 
 
 052 
 
 00°04' 
 
 84°58' 
 
 19 
 
 11 
 
 67 
 
 9.5 
 
 NE 
 
 D-68-1 
 
 HH-7 
 
 30°19' 
 
 3 13' 
 
 1 2 
 
 25a 
 b 
 
 Hemanthias viuanus 
 Hemanthias uiuanus 
 
 c 
 
 Hemanthias uiuanus 
 
 26 
 
 Hemanthias peruanus 
 
 27 
 
 Hemanthias uiuanus 
 
 28 
 
 Hemanthias uiuanus 
 
 29a 
 b 
 c 
 
 Hemanthias vwanus 
 Hemanthias uiuanus 
 Hemanthias uiuanus 
 
 30 
 
 Hemanthias uiuanus 
 
 31 
 
 Paranthias furcifer 
 
 32 
 
 Epinephelus striatus 
 
 33 
 
 Epinephelus sp. 
 
 34 
 
 Epinephelus niueatus 
 
 35a 
 b 
 c 
 d 
 
 Mycteroperca microlepis 
 Mycteroperca microlepis 
 Mycteroperca microlepis 
 Mycteroperca microlepis 
 
 Mi 
 
 Mycteroperca microlepis 
 
 37a 
 b 
 
 Liopropoma sp. 
 Liopropoma sp. 
 
 38 
 
 Liopropoma sp. 
 
 39 
 
 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 
 
 NE 
 
 D-71-5 
 
 LK-5 
 
 35°59' 
 
 74°33' 
 
 5 
 
 5 
 
 71 
 
 5.3 
 
 NE 
 
 D-71-5 
 
 BB-8 
 
 32°39' 
 
 77°17' 
 
 1 
 
 5 
 
 71 
 
 6.8 
 
 NE 
 
 D-71-5 
 
 BB-8 
 
 32°39' 
 
 77°17' 
 
 1 
 
 5 
 
 71 
 
 9.3 
 
 SW 
 
 6611J 
 
 147.30 
 
 23°36' 
 
 111°42' 
 
 13 
 
 11 
 
 66 
 
 9.3 
 
 NE 
 
 D-71-5 
 
 N-7 
 
 34°13' 
 
 75°35' 
 
 29 
 
 4 
 
 71 
 
 10.3 
 
 NE 
 
 D-68-1 
 
 HH-7 
 
 30°19' 
 
 80°13' 
 
 1 
 
 2 
 
 68 
 
 4.3 
 
 NE 
 
 D-71-5 
 
 N-7 
 
 34°13' 
 
 75°35' 
 
 29 
 
 4 
 
 71 
 
 5.8 
 
 NE 
 
 D-71-5 
 
 N-6 
 
 34°23' 
 
 75°39' 
 
 29 
 
 4 
 
 71 
 
 6.6 
 
 NE 
 
 D-71-5 
 
 N-7 
 
 34°13' 
 
 75°35' 
 
 29 
 
 4 
 
 71 
 
 7.2 
 
 SE 
 
 BLM-2 
 
 1-3 Day 
 
 27°34' 
 
 96°07' 
 
 10 
 
 4 
 
 75 
 
 8.6 
 
 SW 
 
 5612 
 
 157G10 
 
 22°30' 
 
 10915' 
 
 3 
 
 12 
 
 56 
 
 7.6 
 
 NE 
 
 D-67-4 
 
 GG-7(2) 
 
 30°54' 
 
 80°00' 
 
 12 
 
 5 
 
 67 
 
 8.4 
 
 SW 
 
 OP 
 
 162 
 
 11°10' 
 
 92°00' 
 
 26 
 
 11 
 
 67 
 
 10.2 
 
 NE 
 
 D-67-16 
 
 GG-7 
 
 30°54' 
 
 80°00' 
 
 23 
 
 10 
 
 67 
 
 4.0 
 
 NE 
 
 1 
 
 4 
 
 35°03' 
 
 75°10' 
 
 11 
 
 4 
 
 72 
 
 7.4 
 
 NE 
 
 3 
 
 2 
 
 35°03' 
 
 75°08' 
 
 27 
 
 4 
 
 72 
 
 14.2 
 
 UNC 
 
 
 T-9 
 
 New Rivei 
 
 ,N.C. 
 
 26 
 
 4 
 
 72 
 
 22.6 
 
 UNC 
 
 (UNC 7934) 
 
 
 Bogue Sd 
 
 , N.C. 
 
 16 
 
 6 
 
 73 
 
 9.8 
 
 NE 
 
 D-71-5 
 
 AA-7 
 
 33°37' 
 
 76°47' 
 
 30 
 
 4 
 
 71 
 
 6.3 
 
 NE 
 
 D-67-4 
 
 NN-4(2) 
 
 27°12' 
 
 79°51' 
 
 7 
 
 5 
 
 67 
 
 7.0 
 
 NE 
 
 3 
 
 2 
 
 35°03' 
 
 75°08' 
 
 27 
 
 4 
 
 72 
 
 12.0 
 
 SE 
 
 OR II 702 
 
 0-2-13 
 
 26°47' 
 
 84°34' 
 
 13 
 
 9 
 
 70 
 
 6.6 
 
 NE 
 
 D-67-8 
 
 AA-4U) 
 
 34°11' 
 
 77°11' 
 
 1 
 
 8 
 
 67 
 
 4.7 
 
 NE 
 
 D-67-4 
 
 MM-4(1) 
 
 27°46' 
 
 79°56' 
 
 8 
 
 5 
 
 67 
 
 6.1 
 
 SE 
 
 OR II 7739 
 
 56 
 
 20°00' 
 
 80°13' 
 
 30 
 
 7 
 
 72 
 
 6.1 
 
 NE 
 
 D-66-3 
 
 N-5U) 
 
 34°33' 
 
 75°44' 
 
 20 
 
 4 
 
 66 
 
 10.2 
 
 NE 
 
 D-67-4 
 
 JJ-6U) 
 
 29°49' 
 
 80°14' 
 
 10 
 
 5 
 
 67 
 
 10.3 
 
 SE 
 
 OR II 7239 
 
 56 
 
 20°00' 
 
 80°13' 
 
 30 
 
 7 
 
 72 
 
 6.0 
 
 s\\ 
 
 6509B 
 
 118.39 
 
 28°19' 
 
 115°24' 
 
 17 
 
 9 
 
 65 
 
 6.7 
 
 SE 
 
 BLM 
 
 IV-2 
 
 26°10' 
 
 96°39' 
 
 30 
 
 5 
 
 76 
 
 7.4 
 
 NE 
 
 3 
 
 2 
 
 35°03' 
 
 75°08' 
 
 27 
 
 4 
 
 72 
 
 6.6 
 
 NE 
 
 D-67-8 
 
 AA-4(1) 
 
 34°11' 
 
 77°11' 
 
 1 
 
 8 
 
 67 
 
 10.6 
 
 NE 
 
 D-67-4 
 
 CC-5U) 
 
 33°20' 
 
 78°25' 
 
 14 
 
 5 
 
 67 
 
 10.3 
 
 NE 
 
 D-68-1 
 
 HH-7 
 
 30°19' 
 
 80° 13' 
 
 1 
 
 2 
 
 68 
 
 10.2 
 
 NE 
 
 D-67-16 
 
 GG-7 
 
 30°54' 
 
 80°00' 
 
 23 
 
 10 
 
 67 
 
 10.3 
 
 SE 
 
 OR U 7239 
 
 56 
 
 20°00' 
 
 80°13' 
 
 30 
 
 7 
 
 72 
 
 50 
 
NOAA TECHNICAL REPORTS 
 NMFS CIRCULAR AND SPECIAL SCIENTIFIC REPORT 
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