28 

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 

Thp maJor 

\"estigatlOn~ appear Ir Ihl' (Irl ultlr t-rlt" 

:'\OAA T,'rh'llcal H"jlorl "\H SCI" ul,r rt' a\ I ahl< fret n l'nlll'd r nb' 

396, 

also a\" 1,,01(' In ,'xcho'lg(' hlf othN (!Cruflc Ind I .. IJrual pull! ( 

olhprwhe nlll'd) fr'lm D:.!~ l ('r l'n) l Hrand'" E.m r nm ntal 

(ula" are 

guide tv their IdentIfication 

ernment 

00119-0 

197 :'Ilarin flora and launa 01 the n >rth,'11 tNn l natcd 

Cmdar a, :nphozoa By Iknald'; Lars. n ALl t 1 

28 'I!:" For ale b) lhe upennlendent of [) 'Jment 

"ent PnrtlO~ Office \\ lI,bmgto:l, D( 204~, 

ooa8~, 

'198, :'II anne nora and fauna of the northea lern llllted 

fung,' Ascomycete" Deutero["lycete~ and Ba IdlOrncetr 

Cavaliere :'II arch 1977 111 .. 4'1 p, l4 fig For sale b} tbe super 

intendent of Documenb, l ., l;O\ernml'nt !'nntlng Ofr " hIOgtOr., 

DC 20402 'tock. '0, 00:1 020 i.l<Jl2f)-2 

399, :'Ilanne nora lind fauna 01 the ncrthe te''l lllIted ... tat 

Copepoda: Harpactlcoida, By Bruce r ('lull \llIr h I n,1I1 + 4' p , 

100 figs, For sale by the Supermtendent 01 Docu'llrn16, l ~ (,o\Crn 

ment Printing Office. \\"a>hlngton, Dr 2040~, lock. 0 :l1)~0 125· 

4, 

400 Fishery pubhcation index, 1965 74 By ~Iary Ellen Engelt and 

Lee C Thorson, :'Ilarcb 1977 111 t 220 p, FOf ,ale b} the uper· 

intendent of Documents, U,S Government Prmtmg Office \\ a blOj(ton. 

DC 20402; Stock ).0, 003-020-00127 ! 

401 Fishenes and fishery resources of :-';ew York H,ght. l:h J L 

McHugh, March 1977, v + 50 P 4,1 figs , 49 tables, For ,ale by the 

Superintendent of Documents. L' S Government Pnntlng Office, \\'a h· 

ington. DC 20402; Stock:-';o 003-020-00129-7 

402, Guide to tbe identification of scorplOnfi,h larvae (FamIly 'cor 

paenidae) in the eastern Pacific with comparative note. on specie. 01 

Sebastes and Hellcolenw from other oceans, By H Geoffrey :'Ilo,er 

Elbert H Ahlstrom, and Elaine M, Sandknop, Apn11977, v + il p" 40 

figs,. 38 tables, For sale by the Superintendent of Documents. U S Government Printing Office. Washington, DC 20402, Stock :'\0, 003-020-

00128-9 

403, Marine flora and fauna of the northeastern United States Sipuncula. By Edward B, Cutler, July 1977, III + 7 p ., 6 figs, For sale by 

the Superintendent of Documents. U,S, Government Printing Office, 

Washington, DC 20402; Stock No. 003-012-00404-0, 

404 , Revision of the sea basses of the genus Diplectrum (Pisces: Serranidae). By Stephen A. Bortone, September 1977, v + 49 P,. 15 figs,. 

9 tables, 

405 , Marine flora and fauna of the northeastern United States, Echinodermata : Holothuroidea, By David 1. Pawson, September 1977. iii + 

15 p " 3 figs, For sale by the Superintendent of Documents, U,S, Govrnm nt P' n nK orrl 

(H ) I 

( I 

~" I 

I" 

tat 

Februa 

Brancbl 

ne'" PfCI 

i table For sale b\ the . upenntendent 

Pnlltlng Office. \\ a.hlDgton, DC • 

41J MOrine flora and launa 01 tbe northea.tern Lruted [ate 

('ru tncea Hrnnchluro Hv Roger F Cre"e) \Ia} 1!1i , },i + 10 ;J " 11 fi~" F)f ,a e b) tbe upenntendl'nt of Document,. l _ GoHrnmen 

i'nntlng Office. \\'a,hlngton, DC 20402, , tock :-.;", )3,01 i ·00419-1> 

414 ~\ nc P'" 01 blologH.'a, da[a for the winter flounder, P,eudopleuT 

'H'Ct .. , amCTlcanu.- l\\"albaum) B) Cruce Kleln-:'IlacPbee , :-';o\'embe 

Hn" iii + 43 P 21 figs" 28 table41 5, A ba'i' for c1a>sifying we.tern AtlantIC SCiaellldae \Teleostei 

Perciformesl By Labbish :-';lng Chao, ' eptember 19: , \' + 64 p .• 4 

fig> __ I table . 

416. Ocean vanability Effects on LS. marine fisbery resources ' 

1975. t2U papers I By Juhen R Goulet. Jr. and ElIzabeth D, Haynes 

Editors, December 1978, iii + 350 p , 

417 Guide to the identIfication of genera of the fish Orde 

Ophldllformes with a tentative classIficatIOn of the order. By Daniel M, 

Cohen and J~rgen G, :--lielsen. December 1978. vii + 72 p" 103 figs,. 2 

tables. 
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 

Juanita M. Kreps , Secretary 

National Oceanic and Atmospheric Administration 

Richard A. Frank, Administrator 

National Marine Fisheries Service 

Terry L. Leltzeli, Assistant Administrator for Fisheries 

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 

8erraniculus .................................................................. 12 

Serraniculus pumilio .................................................. ......... 12 

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Subfamily Anthinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 

Plectranthias ................................................................. 16 

Anthias ...................................................................... 16 

Pronotogrammus .............................................................. 20 

Hemanthias ........................................................... ..... .. 22 

Hemanthias uiuanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 

Subfamily Epinephelinae ........................................................... 26 

Paranthias ........................... . ....................................... 30 

Epinephelus .................................................................. 31 

Mycteroperca ................ . ................................................ 34 

Mycteroperca microlepts ........................................................ 34 

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 

Subfamily Grammistinae ........................................................... 38 

Ltopropoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 . . . . . . . . . . . . . . . . . . . . . . 17 

16. Larvae of Anthias sp. Type 1 from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . 18 

17. Larvae of Anthias sp. Type 2 from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . 19 

18. Larva of Anthias tenuis from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 

19. Larva of Anthias sp. Type 3 from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 

20. Larvae of Anthias gordensis from the eastern Pacific Ocean ................................. 21 

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

larva ................... ......................................................... 22 

22. Larvae of Pronotogrammus aureorubens from the northwestern Atlantic Ocean ................. 23 

23. Larvae of Pronotogrammus eos from the eastern Pacific Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 

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

aureorubens larva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 

25. Larvae of Hemanthias uiuanus from the northwestern Atlantic Ocean ......................... 25 

26. Larva of Hemanthias peruanus from the eastern Pacific Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 

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

larva ...... . ...... ... ........ . ... ................................................ 26 

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

larva ..... .. ... . ...... . ............ .. .... ........................................ 26 

29. Development of the supraoccipital crest of Hemanthias uiuanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 

30. Details of the head spination on a H emanthias uiuanus larva ................................ 29 

31. Larva of Paranthias furcifer from the eastern Pacific Ocean ................. ................ 31 

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

Ocean . ....... ............................................... ................... 32 

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

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

larva ............... .. . ..... .. ... ... ............................................. 34 

35. Young stages of Mycteroperca microlepis from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . 36 

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

microlepis larva ............................. ........ ... ........................... 38 

37. Larvae of Liopropoma sp. from the northwestern Atlantic Ocean ............................. 40 

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

sp. larva ......................... .. . .... . ........................................ 41 

39. Larva of Rypticus sp. from the northwestern Atlantic Ocean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

40 Larvae of Pseudogramma gregoryi from the northwestern Atlantic Ocean ...................... 42 

41. Spine-bearing bones of the opercular and posttemporal regions on a 10.3-mm Pseudogramma gregoryi larva ................................... .. ............... .......... 44 

42. Larvae representative of four groups of American Serranidae ..... .... ............. .......... 46 

43. Interoperculars oflarvae of representatives of four groups of American Serranidae . . . . . . . . . . . . . . . . 47 

Tables 

1. Some adult and larval characters of genera of North American serranids ....................... 3 

2. Meristic character development of larvae of Serraniculus pumilio ............... ............. 14 

3. Body proportions of larvae of Serraniculus pumilio ......... ............................... 15 

4. Meristic character development oflarvae of Hemanthias uiuanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 

5. Body proportions of larvae of Hemanthias uiuanus ........................................ 28 

6. Meristic character development of larvae of Mycteroperca microlepis ......................... 37 

7. Body proportions of larvae of Mycteroperca microlepis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 

;\leri tic character development of larvae of Pseudogramma gregoryi . . . . . . . . . . . . . . . . . . . . . . . . . . 43 

9. Body proportions oflarvae of Pseudogramma gregoryi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 

iv 
Morphological Comparisons of North American 

Sea Bass Larvae (Pisces: Serranidae) 

ARTHUR W. KENDALL, JR.1 

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 development of adult morphology. The serranine genera can be ranked in order of increasing larval specialization as follows: Serraniculus, Paralabrax, Centropristis, Diplectrum, and Serranus. 

Hypoplectrus and one type of Diplectrum, although considered serranines on the basis of adult characters, are quite different from the other serra nine 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 serrated 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 Epinephelinae 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, Pseudogramma 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 characters with the serranines. Their most outstanding larval feature is the development of one or two greatly 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 Eigenmann'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 variously 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: orthwest 

& Alaska Fisheries Center, National Marine Fisheries Service, NOAA, 

2725 Montlake Boulevard East, Seattie, WA 98112. 

1 

larvae indicate that the family may be subdivided based 

on specializations of the larvae. In this paper, subfamilial and generic characteristics of American serranid larvae are defined from material from western 

Atlantic and eastern Pacific waters and one representative of each of the four subfamilies is described in detail. 

METHODS AND MATERIALS 

Since only Hypoplectrus 2 and Centropnstis (Hoff 

1970) among American serranids have been reared 

beyond yolk exhaustion, the indirect method of assembling 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 appeared on the larvae, e. g., as spots, patches, or blotche . 

Body proportion measurements were made with an 

ocular micrometer. The larval period was divided into 

three stages: preflexion (before notochord flexion), flex-

'W. J. Richards, 0<atlonal :'.Iarine Fisherie er.·ice, outhea.;t Fl h· 

eries Center, Miami. peTS. commun. January :974 
ion (during notochord bending), and postflexion (after 

the notochord tip had completed flexion) following recommendations 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 notochord (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 (standard 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 overlying tissue in the cleared and stained specimens. 

Specimens included materials from collections at the 

National Marine Fisheries Service (NMFS) Laboratories in La Jolla, Calif.; Miami, Fla.; Narragansett, 

RI.; and Sandy Hook, N.J. 

Most of the material was collected as part of one of 

three large-seale 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 California 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 American waters with 52 species and are moderate to smallsized serranids apparently representing the base from 

which other subfamilies arose (Smith and Atz 1969; Kendall 1976). Most serranines are synchronous hermaphrodites; 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 European species of Serranus (Fage 1918; Bertolini 1933; 

Aboussouan 1972) as well as tbose of Centropristis striata 

(Kendall 1972) are the only serranines described. 

'Serranids studied to date are hermaphroditic (Smith 1965; Kendall 

1977), l.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 produced in each individual. Hermaphroditism takes on two forms in serranids: synchronous hermaphroditism, where both types of gametes are 

produced simultaneously in one individual, and protogyny, where individuals mature first as females and later in life the gonad transforms 

Into testes and produces sperm. 

2 

Larvae of all but one (Schultzea) of the Nortb American 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 specializations of other serranids. They do not have elongate preopercular 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 mostl 

along the ventral midline. Generally, there are spots a

the angle of the jaw, at the junction of the cleithra, be

tween the bases of the pelvic fins, near the anus, at th

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 th

subfamily but some is consistent within genera. Th

body shape of the larva gradually changes to that of th

adult, without abrupt changes in proportions. 

Centropristis.-Centropristis occurs only in the 

northwest Atlantic where four species are recognized 

(Kendall'). These are moderately large protogynous hermaphrodites. Centropristis striata occurs farther north 

than any other serranid on the east coast. A closely related 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 0 

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 produced. 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 pigment but afterwards nearly all larval pigment is asso-

'A. W. Kendall. 1977. Biological and fisheries data on black se 

bass, Centropristis striata (Linnaeus). Sandy Hook Lab. Tech. Se! 

Rep. No.7, 29 p. 

5Some (e.g., Miller 1959) consider C. melana a subspecies of C. striata. 
w 

Table I.-Some adult and larval characters of genera of North American serra nids. Meristic characters a nd types of sexuality were compiled from several sources cited in the text. Ramus lateralis 

accessorius (RLA) patterns from Freihofer (1963), Predorsal patterns from Kendall (1976) a nd Kendall a nd Fahay (in press). Scale morphology from McCully (1961). 

Number 

of 

species 

Adults 

u 

'';:; 

<= u 

0.::: 'u 

Ranges of meristic characters 

+3 '" Subfamily Fins 

genus ..: '" p.. Dorsal Anal Pectoral Vertebrae Sexuality 

Serraninae 

Serranieulus 

Paralabrax 

Centropristis 

Dipleetrum 

Serranus 

Hypopleetrus 

Sehultzea 

Anthiinae 

4 

3 

13 

7 

Plee tranthias 1 

Anthias 3 

Pronotogrammus 1 

Hemanthias 2 

Holanthias 

Centristhmus 

Caprodon 

IX-X,lI 

8 X-XI,12-15 

X,ll 

9 IX-X,ll-13 

5 IX-X,W-13 

1 X,14-16 

X,ll-12 

X, 

16 

14-15 

2 13-15 

13-14 

15-16 

14 

19-20 

VIII, 13 

IX,18-19 

Ill, 

7 

6-8 

7 

6-9 

7-8 

7-8 

6-8 

7 

7-8 

7-8 

7-8 

7-8 

8 

8 

Epinephelinae 

Goniopleetrus' 

Paranthias 

Epinephelus 

Myeteroperea 

15 II X-XI, 14-20 

7 

9-10 

8-10 

W-13 

Grammistinae 

Liopropoma 

Pseudogramma 

Ryptieus 

Pikea 

J eboehlkia 

8 5 XI,15-18 

5 

1 

8 

3 

1 VIII,12-13 Ill,8 

1 VII-VIll,15-24 Ill,12-20 

2 II-III,21-29 14-18 

VlII-IX,13-14 1I1,7-8 

VII,9 Ill,7 

'Dipleetrum Type 2 larvae. 

'Provisionally placed in Epinephelinae. 

14-15 

15-19 

15-20 

15-18 

13-18 

14 

15-17 

10+14 

10+14 

10+14 

W+M 

W+14 

10+14 

10+14 

synchronism 

sec. gonoch. 

protogyny 

synchronism 

synchronism 

synchronism 

13 W+16 

18-20 10+ 16 protogyny 

15-19 W+16 

18 10-ll +15-16 

17-18 10+16 

19 ll+15 

17-18 10+16 

16 

19-20 

16-20 

15-18 

12-15 

14-18 

13-17 

15 

15 

W+14 

W+ 14 protogyny 

W+ 14 protogyny 

W+ 14 protogyny 

10+ 14 sec. gonoch. 

10+ 16 synchronism 

10+ 14-15 protogyny 

W+14 

9+15 

Larvae 

<= 

... 

~ 

'" 0. 

..: 

i=2 

9 

9 

9 

9 

9 

9 

Oi 

~ E a ., .".., 

., .., 

p.. ... '" 0. 

0/0/0,2 

0/0/0,2 

0/0/0,2 

0/0/0,2 

0/0/0,2 

0/0/0,2 

0/00/2 

g 

a 

o 

..<: 

~ E-

'" a cJ; 8 

III 

III 

III 

III 

III 

Body 

shape 

Slender 

Slender 

Slender 

Slender 

Slender' 

Slender 

Slender 

Fin Spine Development 

Precocious Produced 

and/or head spines 

elongate 

- u '" .- '" > a ... -., o p.. 

x X 

3(some) X 

Serrate 

- u '" .- '" > ... a -., o p.. 

~ ~ ~ 

~ . .@ '"5.:3 ............... 0.. u s.... C,) 

co o.'u <V :u ~ ~ 

~~gp:;g...sg. 

Larvae unknown 

9 0/00/2 or III 

0/0/2 

Robust 

Robust 

3rd 

1-3 X X 

(some) 

X 

X 

X 

X 

X 

X 

9 

U/0/2 

9 0/00/2 

None 0/0/1 

None 0/0/1 

None 0/0/1 

None 0/0/1 

None 0,0/1 

None 0,0/1 

None 0/1 

None 0,0/1 

III Robust 

Robust 

2-3 

2-4 

II Robust 2-3 

II "Kite" shaped 2-3 

II "Kite" shaped 2-3 

II "Kite" shaped 2-3 

II 

II 

II 

II 

Slender with 

deep caudal 

peduncle 

2-3 

2-3 

X 

X 2-4 X X 

X 

X 

X 

X 

Larvae unknown 

Larvae unknown 

Larvae unknown 

X 1-3 X X 

X 1-4 X X 

X 1-4 X X 

X 1-4 X X 

X 

X 

X 

X 

X 

Larvae unknown 

Larvae unknown 

Pigment locations 

Ventral 

8 

~ 

... 

..<: 

~ :~ 

"-:> '" U -

<=: 

a u+3 .- ... > ., -., '" <=: p...-

X X X 

X 

X X 

X X 

X 

X 

X 

X 

X 

X 

X 

<= 

0.::: 

'" -., ~ til '" C; <=: til 

..: ":.n 

X X 

X X 

X X 

X X 

X 

X X 

X X 

X 

X 

X 

X 

~ 

Fin 

membranes 

- u_ 

~-§ ~ .~ ..!:I::: 

~." a 

c3~o 

> <=: - til ., - p.. "'" 

X X 

X X X 

X 

X X X 

X X 

X X X X 

X X X X 

X X 

X X X X 

X X X 

X X X X 

X X X 

X X X 

X X X 

X 

X 

X 
a ~-- -----

/) 

~ 

/ l.6-A- • 

Y @ '- )" ~ -~~ -- - ->.~--) j - " --

--:.-

b . 

{ @ 't \"" } 

Figure I.- Larvae of Centropri.,ti, .,triata from the north"l:,tcrn \ttHntlt ()CI:8n: a) 1.7 mm; bl .3 mm. 

posttemporol 

s l;.lroc. elt HUrT' 

Figure 2.-Spine·bearing bones of the opercular and posttemporal 

regions on a IO.6·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 peduncle spots, and at the bases of some caudal fin rays. 

Several melanophores also occur on the posterior portion 

4 

01 the ')plie lobes and on the dor opu terior lIning of the 

COelOl1lll cavity. 

Paralabrax.- Paralabrax with eight pecies i the 

onl) • orth Americanerramne lImited to the ea tern Pa· 

clfic Crable 1) It i ecologically imilar to Centropristl 

in the AtlantiC In that It is larger than mo t other erran· 

ines and occur at higher latitude than other serranines 

on the Palilic coa t. 'mith and Young (1966) indicated 

that Cf!ntrupri.,ti~ and Paralabrax are al 0 clo ely reo 

lated . The only specie that has been tudied is a ec· 

ondary gonochori ,t. while mo·t other erranine are ynchronou hermaphrodite (Smith and Young 1966). 

Paralabrax larval development i typical of erranines. Paralabrax larvae approach adult body proportlOm; oon after tail flexion is complete (Fig. 3). The 

plnOUS and soft portions of the dar al fin form at abou 

the arne rate, as they do in Centropnsti . The fins a 

Paralabrax larvae are more heavily pigmented than 

those of Centropristis but the ventral mel ana phares 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 tin. near the anus, along the base and at the insertion 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 

b \ \ 

<,-~ 7 

Figure 5.-Larvae of Diplectrum sp. Type 1 from the northwestern Atlantic Ocean: a) 5.8 mm; b) ]0.0 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 scattered 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 larvae have pigment on and between the fin rays. The preopercular 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 serranid 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 representatives 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 larvae, 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 III smaller specimens (Fig. 8). The sequence and 

extent of fin ray formation is different from that in other 

serranmes. The pectoral fin develops early, before most 

other tins, as a fan-shaped structure and extends to the 

anus. Only m grammistines is similar larval pectoral fin 

development seen. The pelvic fins form before the rest of 

6 

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. Pigment 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 outside the ranges of other American serranids except 

Schultzea beta, which has only six branchiostegal rays 

(Robins and Stark 1961) and Diplectrum, which generally has more pectoral rays (Table 1). 

Robins and Stark (1961) examined east coast Serranus 

species in some detaiL Since Serranus in the eastern Pacific is poorly known with few specimens of some species 

taken, some species may prove invalid. 

While no American Serra nus larvae have been described, eggs and early larvae of Mediterranean species 

of Serranus were among the first serranids described 
a 

b v 

• 

c 

d 

F igure G.-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. 

7 
'? 

a 

b 

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

posttemporal 

operculor 

suboperculor 

Figure 2.-Spine-bearing bones of the opercular and posttemporal 

regions on a IO.6-mm Centropristis striata larva . 

cia ted 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 peduncle spots, and at the bases of some caudal fin rays. 

Several melanophores also occur on the posterior portion 

4 

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 Pacific (Table 1), It is ecologically similar to Centropristis 

in the Atlantic in that it is larger than most other serranines 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 secondary 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). Th 

spinous and soft portions of the dorsal fin form at abou 

the same rate, as they do in Centropristis. The fins 0 

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 insertion 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 
a 

b . . 

11 Uri I niH' (lr "flrn/Clbrax " rr m Ih north a 

(\~q Il'lltmllo (hI' n 

nd t' (IWrl' re 8 It '" blunl pill 

hi tI 11111<'lh po 1 nor m rglll 

/);n/ I'ctrllm 

m IIl\ ollwr 

• 

• 

n 
b 

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 scattered 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-

\ae have pigment on and between the fin rays. The preopercular and subopercular have more armature than is 

. een in other serranine genera (Fig. 7). 

rype 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 serranid genus. This would indicate that either these larvae 

are not of Diplectrum but are of an undescribed genus, 

possi bly even in a closely related family with representative. ill both oceans, or that there are two quite distinct 

t) p ofLJiplectrum. possibly worthy of subgeneric rank. 

\\ hile Type 1 larvae are similar to other serranine larvae, 'I ypc ~ larvae show specializations not seen in other 

'ener I. The general body shape is similar to that of Type 

1 but the :-nout IS more elongate and is concave in dorsal 

profil in muller specimens (Fig. 8) . The sequence and 

t nt of III ra) tormation IS different from that in other 

I'he pectoral lin develop early, before most 

I,., H fan-"haped structure and extends to the 

nu nh 111 gr mmi:-tines i" similar larval pectoral fin 

d \ '(opm nt en. I he pelviC lin" form before the re t of 

6 

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. Pigment 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 outside the ranges of other American serranids except 

Schultzea beta, which has only six branchiostegal rays 

(Robins and Stark 1961) and Diplectrum, which generally has more pectoral rays (Table 1). 

Robins and Stark (1961) examined east coast Serra nus 

species in some detail. Since Serranus in the eastern Pacific is poorly known with few specimens of some species 

taken, some species may prove invalid. 

While no American Serra nus larvae have been decribed, eggs and early larvae of Mediterranean species 

of Serranus were among the first serranids described 
a 

b v 

• 

c 

d ... . . 

Fi!(lIrt' Ii. Lan a~ or lJipltdrum '[\. 'I \ pc I rrom th .. n rthe -t(,rn I' ,Ii 0 n 

-I 
posttemporal 

opercular ~ ,"p'0de;,h"m 

subopercular 

Figure 7.- Spine-bearing bones of the opercular and posttemporal 

regions on a lO.3-mm Diplectrum sp. Type 1 larva. 

a 

b 

Figure 9.-Spine-bearing bones of the opercular and posttemporal 

regions on a 9.9-mm Diplectrum sp. Type 2-larva. 

Figure 8.- Larvae of Diplectrum sp. Type 2 from the northea stern 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 Atlantic; however, none were found in Pacific collections. 

8 

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 beginning of the flexion stage. More detailed study may allow descriptions of the several types and their assignment to species 
" ': \ "fIi 

'\...1 .. . . :V : 

a : " .: . . . . . . , 

b 

c 

d 

Figure lO.- Larvae of S erranus sp. from the northwestern Atl a ntic Ocean : a) 3.7 mm ; b) 5.0 mm; c) 5.5 mm; d) 9.4 mm. 

9 
Serranus development shows several modifications of 

the basic serranine plan (page 2). The body is considerably 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 Serranus larvae described so far these fin spines are longest 

in S . 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 pectoral fins in some larger larvae. 

The opercular region is armed with blunt spines, which 

are generally more pronounced than those of other serranines (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-

~sttemporal 

D wp;odoith,"m 

preoperculor 

o~;rl 

'\ / E I 

~ ~ ~j WOOPO""'O; 

/~ interopercular 

Figure 11 -Spine-bea ring bones of the opercular and posttemporal 

regions on a 9.7-mm Serranus sp. larva. 

10 

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 considered valid species; however, other species of serranids 

such as Mycteroperca rosac ea 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 Serra nus and. is close to S. flauiuentrus 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 detailed 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 differences and it is difficult to consider them as modifications of serranine larval characters. These Hypoplectrus larvae are not similar to any known serranid 

larva and provide reason to doubt that Hypoplectrus belongs 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 mem branes of these early forming fins are 

pigmented. The rest of the dorsal spines are progressively 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 
a 

b 

, 

c 

,. .,. , *--:., . 'f 

~'\ , J 

- , 

d co • ·':"0' " ... ' .' 

Figure 12.-Laboratory·reared larvae of Hypop/ectrus sp. from eggs collected near ;\1iami. Fla.: a) 3.~ mm; b) ~.7 mm: c) .3.7 mm; d) 

8.5 mm. 

11 
than the longe t pines. 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 

spine developed on several bones; however, they do not 

appear on the larval drawings since they are covered by 

He h. The preopercular is edged with about 12 subequal 

pines (Fig. 13). The subopercular has about three blunt 

spines on its margin. Besides the three definitive spines 

on the opercular, there are about fo ur spines ventral to 

them, a condition not seen in serranines. 

Pigmentation consists of irregular shaped and sized 

melanophores in characteristic positions. More pigment 

i present on Hypoplectrus than on most other serranines. A series of spots along the ventral midline consists 

of a _ ot near the base of the pelvic fins, one below the 

middle of the gut, one just anterior to the anus, one near 

the in ertlOn 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 scattered on the dorsal surface of the brain posterior to the 

eyes. All the fin membranes have pigment in later larval 

. tages. By 8.5 mm, small melanophores become scattered over the dorsal portion of the anterior two-thirds of 

the fish. 

_ ... /'" > interopercular 

......-/ 

supracleithrum 

1'J::ur 1:1 !'ipine-beuring bones of the opercular and postlemporal 

,,'gions on a 12.~-mm Hypopleclrus sp. larva. 

... rhultzra.-Schultzea is monotypic represented by S. 

b ta of the \\e tern Atlantic (Table 1). It lives more peI glc II than other serranines and is adapted to feed on 

mall, fr e wimmlng prey. The mouth is modified to be 

hlghh 1Jr(,tru.lbl' through extensive changes in the 

hap 01 the premo 'illary (Robms and Stark 1961). The 

fin C( lint re lmll r t<l t hose of Serranus but there are 

unh I br n hi teg I rays instead of seven (Robins and 

t r I( 1). 

12 

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 cannot be determined. However, from adult adaptations, it 

appears that Schultzea is the most highly modified 

American serranine . 

Serraniculus.-Serraniculus contains one diminutin) ~pecies «80 mm), S. pumilia, in the northwest 

Atlantic (Table 1) . Its life history has been studied and it 

i~ a ~.\' nchronous hermaphrodite (Hastings 1973). 

A serranine larval type from the Atlantic had meristic 

characteristics of S. pumilia, 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 pumilia (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 development. Since the smallest larvae have a straight notochord and vertebrae are the only ossified meristic elements, and the largest have adult complements of all 

meristic characters except gill rakers and scales, the pattern of meristic character development can be seen in 

this series. Vertebrae ossify from anterior to posterior except 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 complement of six is seen in most specimens by 4.0 mm, when 

the notochord has completed flexion. Other meristic elements 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 subfamilies and even in Diplectrum and Serranus. The dorsal fin spines and rays each develop at about the same 

rate in S. pumilia . 

Body proportions (Table 3)-Several body proportions of 21 larvae from 2.9 to 8.3 mm were measured and 

expressed as a percentage of body length (BL) to document 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; thereafter, it increases gradually to about 125% of BL as the 

caudal fin forms. Preanal length increases from about 

60('( ofBL in larvae < 4.0 mm to about 65% ofBL in 7.0-

to 8.0-mm larvae. Head length remains fairly constant at 

about 39{'( of BL throughout larval development. Eye 

length and nout length decrease slightly during development from about 12{, to slightly less than 10% of 
a 

b 

c 

Figure 14.-Young stages of Serraniculus pumilio from the northwestern Atlantic Ocean: a) 3.11 mm; b) :,.11 mm; c) .).5.3 mm. 

BL. The greatest body depth decreases from 320(. to 

about 28C'O of BL. Depth at anus increases during early 

development up to 4.0 mm and then holds constant at 

about 25C'O of BL. Caudal peduncle depth increases from 

<6CO in BL in the 2.9-mm larvae to about 14('"0 of BL in 

4.4-mm larvae; thereafter, it changes little. A comparison of the proportions of the larvae after notochord flexion with those of the adult shows that the adult body 

shape of S. pumilia is established early in development, 

in larvae as small as 4.2 mm. 

Pigmentation-Throughout larval development, S. 

pumilia is more heavily pigmented than any other 

13 

serranine. The characteristic serranine ventral pot are 

present, but they are supplemented by many other pot. 

Pigmentation is composed of small, individual melanophores making up a pattern, rather than the pattern being comprised of large melanophore~ in characteri tic 

position as een in other erranid ~. 

The beginning of the pattern is already e. tabli"hed In 

:!.6-mm larvae with three row~ of melanophore on the 

trunk: one along the dorsal midline, one along the lateral 

line, and the third along the \·entral midline. Dor ali). 

there i a erie of about four ,poL. which are larger than 

those in the other two row'. The antenormo t of the e 
Table 2.-Meristic charaeler deH'lopmt! nt "r InrvlII' or Serranu'"/,,, (J"mtltlJ. SP"l"iml'n 

between dashed lines urc:' undergoing noto(' hord Ol'xion. J'o r tht· ('uucla/lin,"/''' and "S" 

indicate primary lind sccondllr~ lin rny_ . 

Body Branchlll- l'cc - I'~/ ( Iludlll lin 

length sIegal verte lorul qC [)","1l1 \ enlrul An«1 f)", "I 1111 (,III 

(mm) rays brae 1111 fin ~ 

3.1 NL 

:1.4 :1 

:1.5 I 

:L5 II 

:1.6 ;, 

:~.6 11 

:J.6 11 

3.7 1U 

:~.8 :! I;; 

:1.8 10 

3. t! 

:3.8 'J 

:J.t! .1 

3.8 It; 

3.ti :1 11 

LO 12 

4.0 Hl 

Hl Ii> 

4.3 Ii 

4.4 11 

1 ~ III 

4.1 17 

4.:! 17 

4.:J ~ 1 i 

1.9 Ii I~ 12 I,~ 7 

38sL fi W 11 I J 

4.4 b ~2 15 1,3 

4.6 6 24 14 I " 2 

4 i Ii '22 9 I 

5.0 6 ~4 14 1- J 

5.2 6 21 15 1.- 3 

5'; fi 2~ 15 I: .j 

5f! 6 21 6 1 4 

6.0 fi :!4 !J 15 

&.1 6 24 15 I ' t, 

6.2 6 23 ~ I·j 

7.0 5 .4 n 14 

70 6 L4 !J I'; 

7.0 6 24 14 15 

7.2 6 24 I.~ I .. ) , 

73 6 24 14 I.'; 7 

spots is just anterior to the origin of the dorsal fin and remains pronounced throughout development. Additionally, 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 association 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 surface. Superficial spots develop from the anus to the termination of the dorsal fin between the dorsal and lateral 

P 

6 

9 

9 

B 

c. 

9 

J 

9 

U 

14 

I' s lin Pin" Hay 

\ 1 

11- \ III 

III" IX ~. 2 

l 111- II 

! 

lIl- 10 J 

!. Il!- II 

4 Il!- ).. 11 " ;, 

II 1\ 

2 II 1 \([[ 5 

~ 1117 II) ;, 

II 1 \11 ,') 

II I \ll 

6 II I 1\ 4 

i !II 7 ~ II ,') 

b ill , X 11 t 

Ii b !Ill X 11 6 

series of da he . In larger lan:ae imilar pot al 0 extend 

ventrally to a Ie er extent from the lateral da he . These 

uperficial spots are mall but variable in ize and po ition and appear a a ingle pigmented blotch. uperficia 1 pigment also develops a a line on the head pa ing 

from the snout po tenorly at the level of the eye and 

another line on the opercular region in line with the 

lateral serie of da hes on the trunk. A patch of small 

spots al 0 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 cleithra. A larger spot is present on the ventral midline between the junction of the cleithra and the base of the 

pelvic fins. Along the gut the ventral spots diverge slightly to form a series on each side of the midline. A few spots 

are also present on the midline ventral to the posterior 

third of the gut. The anteriormost of these is the largest 
Table 3.-Body proportions of larvae of Serraniculus pumilio. Specimens 

bet.ween dashed lines are undergoing notochord fl exion. 

Percent of standard length 

Body 

length Total Body Eye Head 

(mm) length depth length length 

2.9NL 103 32 13.0 39 

3.7 104 31 11.0 38 

3.9 107 33 12.0 38 

4.0 107 34 11.0 39 

4.0SL 115 33 11.0 36 

4.2 118 34 12.0 39 

4.2 115 36 12.0 42 

4.4 119 33 11.0 39 

5.5 117 31 10.0 38 

5.7 122 31 11.0 38 

6.0 123 31 11.0 38 

6.1 123 29 10.0 36 

6.7 117 29 9.4 37 

6.8 120 31 11.0 38 

7.0 120 29 9.8 36 

7.7 125 23 9.8 32 

7.8 124 29 10.0 39 

7.9 117 29 11.0 38 

7.9 124 28 10.0 39 

8.3 125 27 9.9 39 

Adlllt ' 27-31 8.5-10 34.5-36 

Ginsburg (195:2). 

and is ventral to the deepest point of the gut. Posterior to 

the gut, a series of spots extends along the ventral midline 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 development. Other small spots present during development are on the membranes associated with the branchiostegal rays, pectoral and pelvic fins, and internally 

on the dorsolateral surface of the gut cavity. 

Late in larval development the superficial spots intensify 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 examined. Larval material for five of these genera has been 

gleaned from plankton collections, and their development shows a similar pattern. Laboratory-reared material 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 presently understood affinities are based, need to be reexamined. 

Two distinct types of larvae were found that had meristic characters of Diplectrum. Both types of larvae oc15 

Pre- Caudal Depth 

Snout anal peduncle at 

length length depth anus 

10.0 58 5.7 19 

12.0 59 8.0 25 

10.0 63 10.0 27 

11.0 61 9.8 27 

12.0 61 11.0 28 

10.0 63 13.0 28 

11.0 62 12.0 30 

10.0 63 14.0 29 

10.0 66 14.0 27 

11.0 59 13.0 25 

11.0 62 15.0 26 

9.3 65 14.0 29 

9.4 62 14.0 26 

9.0 67 14.0 27 

9.8 65 14.0 25 

7.4 61 13.0 28 

10.0 64 15.0 26 

9.5 64 14.0 25 

9.5 62 13.0 25 

9.9 66 14.0 26 

8.5-10.5 12-13.fi 

curred in both oceans and more than two species of Diplectrum are known from each ocean. Diplectrum Type 2 

larvae are not similar to those of other known serranines

They do not have the characteristic serra nine 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 modifications of the basic pattern of development may indicate relationships. Among American serranids, the 

serranine genera Serraniculus, Centropristis, and Paralabrax 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 elongate 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 positions primarily along the ventral midline. Diplectrum 

Type 1 and Serranus show modifications of the basi() 

serranine pattern in that the spinous dorsal fin develops 

before the rest of the fin, and in some species of Serranus 
some of the dorsal spines are elongate and tipped with 

pigmented "flags." The pelvic fins are also early developing 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.6 This is a cohesive group of 

small, brightly colored fishes that occur over hard bottoms n~ar continental slopes. They are distinguished by 

several morphological modifications from serranine characteristics (Table 1). They have large scale (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 specie of 

anthiines in American waters grouped in even genera. 

Four genera have representatives in both oceans but no 

species are common to both oceans. 

Ten types of larvae assignable to four genera (Anthlas. 

Pronotogrammus, Plectranthias, and Hemanthia) 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 seem 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 western 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. 

·P. C. Heemstra, CSIRO, Cronu\la, N.S.W., Australia, pers. commun . 

April 1973. 

16 

Only a few P. garupellus larvae were found, so a detailed description is not possible. However, the larval 

characters indicate relationships of the species within 

the subfamily and will be discussed briefly. Plectranthias garupellus shows little development of characteristic anthiine larval feature (Fig. 15). one of the fin 

pines are serrate. The third dor al spine in small larvae 

is elongate but not stout. The head has spines in area 

characteristic of other anthiine larvae but they are weakly developed and not serrate . A imple spine protrudes 

from the frontal ridge above the eye. Too few specimen 

were available for staining and illu trating the opercular 

erie but the preopercular ha two ridge of blunt spine 

With an elongate, thin pine at the angle of the posterior 

ridge. The interopercular ha an elongate thin pine lying 

proXimal to the long preopercular spine and the other 

opercular bone have orne pine protruding. The po ttemporal and upracleithral pine protrude bluntly. 

There i no protruding upraoccipital cre t. 

The pigment pattern of larval P. garupellu.s i distinctive. There are three characteri tic blotche of pigment on the trunk. Two blotche are dor ai, one below 

the sixth and eventh dor al fin pine, and the other 

below the four po teriormo t dor aloft ray . The third 

hlotch IS on the ventral midhne ju t po tenor to the insertion of the anal fin. Other pot occur on orne pecimens. uch as one ventrally on the caudal peduncle. 

Anthias. Anthla I circum tropical with everal 

species In the we tern Atlantic and one m the ea tern PaCIfic Ocean (Table 3). They are rather mall, brightly 

colored. deep-bodied fi he . Anthlas anthta· larvae have 

been de cnbed (Fage 191 ; Roule and Angel 1930; Bertolml 193:3) and they clo ely re emble Anthias larvae I 

have examined. I found four type of anthiine larvae in 

western Atlantic material and one type in Pacific 

material. which were quite imilar and larger pecimen 

had meristic feature of Anthias pecie (Figs. 16-20). 

Only one western Atlantic type could be Identified to 

species (Anthta tenui) on the basi of meri tic characters of the larvae. Anthias nlcholsi seems to be the most 

common adult of thi genu in the we tern Atlantic and 

Anthtas Type 1 i the mo t common larval type collected; therefore, Anthtas Type 1 may be Anthias 

nichols!. Since there is only one Anthias pecies in the 

eastern Pacific (Anthias gordensis), the larvae are presumed to be that species (Fig. 20). 

Anthia 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 appearance. 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 
a 

b 

Figure 15.-Larvae of Plectranthias garupellus from the northwestern Atlantic Ocean: a) 5.5 mm; b) 8.0 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 

17 

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 serration varies among the species, some having 3 to 6 serrae 

while others have up to 12. The most conspicuous spine 

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 po terior ridge ha 4 to 8 
a 

b 

Figure 16.-Larvae of Anthias sp. T)pe I from the north"e.tcrn Atlantic Ot'ea n : 8) :1. 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 preopercular 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 

18 

fin, usually a few spot are on the kull above the optic 

lobes, and in some pecie there i a spot at the ymphySIS of the lower jaw. Dorsal trunk pigment varies among 

the specie . In the Atlantic there are four types of 

Anthzas larvae ba ed on dorsal trunk pigment and other 

characters. Anthta Type 1 has a pot on the trunk musculature 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). 
a 

b 

c 

Figure 17.-Larvae of Anthias sp. Type 2 from the northwestern Atlantic Ocean: a) .1.7 mm ; b) .'>.i mm: c) \ . .J mm. 

19 
~ 

.' '"., 

/. . .. 

~ " . ' ~ ::::-...... iI 

v 

, 

, 

\ 

\ 

f 

\ 

> / 

-f I i' 

I~~ ,/7 ~ 

I 

I 

/ 

Figure IS.-Larva of Anthias tenuis from the northwestern Atlantic Ocean. 6.7 mm. 

' ... '<: ..• '-:/ '. .~.' 

~"'-'. ~ .. -, .~ 

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 preopercular 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 protruding supraoccipital crest, and a moderate number of 

spines in the ridge above the eye. Thus, Anthias 

gordensis larvae correspond most closely to Anthias Type 

20 

3 larvae among the Atlantic types. As more becomes 

known about the relationships within this genus, clues 

about the identity of Anthws 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 American 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 
a 

-,\ 

-® .-

b "1 '-", S 1'10 

/ , 1 

'\-4'-- >- I " 

./ @ ~ 

;-

, 

} .. 

-=: =--

• 

Figure 2U . l.an at' of ,\nlhia.\ J:flnl .. n.,i · frum Ih., ,',"I"cn I' co flc ()tt' n ,I .. mm b) m 

has meristic fe t ure, of P. aurcuru ben, and t he Pacific 

type those of P. co" . The two lar\'ae share common lharaders that differentiate them from larvae 01 ther 

gen ra ~one 01 the fin ray, are serrate (Ir mUl'h e1 n~ate 

but the second Hnd third dor, al spine::. form b tor tht> 

ot her elemellt, and the peh It fin b lorm d ar1~ '1 h 

opercular region 1- h a\'ll~ armed with ~e\ r I "errat 

spll1e::. (Fig, ~-t l. The pr opercular ha... I Ill! rr t 

spll1e nt its angl and the l11tt'r 'P reu < r h . a imilar 
posttemporol 

opercular 

0( \c I { 

,~ J ( "bop",,~, 

preopercular I '---~ «,-~ Inleropercular 

Figurt' 21 Splnt'-ht."arin~ hOI1(" of th(l Up('f('ulnr und po..,lfplTlpor 

rel'ion, on a 11I.2·mlO ,\nlh,as ,po ') ~ pt' I 111"" 

the plg-menl c(lnsi~ts of a group III mel!lnoph()re~ that 

form a blotch 01 plg-ment laterally on the trunk 0\ er th • 

last lew rays of the anal fin. Other pigment con I 1'< of 

se\ eral melanophore. on the 'urface at thl optiC lobe:- of 

the brain. A fe\\ spot.; at pigment are at the b e of the 

caudal fin on both species, and on /'. e'i:' orne PII::ment i 

on the pehi( fin membrane. Lower Jaw pIgment i pre· 

ent on P eos and two ventral midline ~pob lire on I). 

oureorubens one at the Insertion of the lInal fin and om' 

midway on the caudal peduncle 

Hemanthias.-Hemanthla L represented by three 

AmenLan specie: H. UIL'onus and H.ieptLl in the tlan· 

tic and H. peruonus in the Pacific. 

I found one type of Hcmanthw larvae 111 eaeh ocean, 

the Atlantic species With 47 lateral line scale~ (Fig. 25) i 

presumably H l'il'anu8, the most northerly species 

From larval catche , H. L iuanu,\ IS the most abundant 

anthiine off the east coast of the mted 'tates. 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 ha\e 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 senes of heavy spines ventral to the 

larger one. Both species have a protruding supra· 

occipital crest which is serrate, resembling a "cock· 

comb." There are numerous ridge 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 t hese 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 

22 

al ~o crrl1!1'. I'ignwnt on II. VW {JrlU con i t of OPP<} ing 

() s on 1111' [rllnk Jnt:dlUl til I hi' po t£'rior extent ()f the 

s('('()nrl elm. ullllld Hllal fill . 1Ilf e i a l 0 u pr)[ ventrally 

IIIl Ill(' (,il url a l JJI'duIll'I ' flnel (mc ut [hI' ('8udal fin base 

SpillS Ill. (J ()( (,lI r Oil the rl llf nl urlul' of th . (Iptic lobe 

II I'mf/lllh/(J~ !)('rIJlU ll.IS 11 11 PI KITl nl (In the membran' fI 

Ill!' pillflllS do r ill IIlld pI·lvlC 1111 (r ig. :lfi). Both pecie 

b( ('IIIll!' (ulerl with IIrrlCltl' ctenOld cnlc whllp th 'Y till 

JlII t' S lafval ('Ilflf[l('tl'r 

J/C' fflrwlhiw; Cit'(UlU 

mm ther a re 1'\ to ~3 verte· 

I pme,Ito phlcray,and tol~ 

gdl r.k rs. I he wnd dors I, n I, pectora!, no c udal 

fin ar d \ 1 pm rapidly durmg thl penod hov.ing 

\\Ie!" rang' 01 Ilumbe of elem nt formed B twe n 6.6 

and 7.j mm, th m p !tern of WIde ran e, of formed 

element I' e 'Il. II1dlC tmg rapid form tion. orne "pecImen, hu\ dult cum pi ment of vertebr , econd dor-

, ,d. un,)!. audal, nd peh ic fin ra} . By 1.6· .0 mm, the 

ddult omplelmnh of II har cter examined except gill 

raker" arl' prl~,enl. 'uto hord nexion ta e place at 5.5 

mm and ... pe imen ... >1).0 mm are developing cale . That 

I • ' Lilies IOTOl when th only rueri. tic character wit h it -

,J(!dlt eomplcment i. the branchir tegal ray, 

~l'mle are \'ell de\'eloped on the third dor al pine and 

the pel\'ll' , pine. The "cL'ond and tou rth do r. al and the 

11T"t nnd "econd anal spll1e .. aLa de \'elop , errae. 

Hod! proportions Table ,'))-Larvae lrum 3.1 to 9.2 

mm were mea"ured to trace the pro portional develop· 

ment 01 se\ eral body dl mensions. T he only proportion 

that show" Il()ticeable di ffere nce with ize i total length, 

which increases from itO' ( oj BL to l:wr ( of BL a the 

larvae grll\\. ThiS difference renec t the formation and 

growth of the audal tin, Grea te t body depth remam 

fairly constant ra ngmg Jrom ~9 to 44 ( of BL; the depth 

at the an us is :W ~ of BL; and caudal peduncle depth i 

l~' ( of BL. Head lengt h is about equal to greate t depth 

ranging from 34 to 46' ( of BL, snout length i 12(( of BL 

as is eye diamet er. The preanal length ranges from 53 to 

6f)( ( of BL. Thus, H . l'it'anus larvae are large headed and 

deep bodied with a trunk that tapers quickly from the 

anus to the caudal peduncle . 
a 

b 

c 

.. ~ ... -~(. 

~'J: ~ ~. ; J ~/ 

~ 

II 

ilr--<: 

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

9.0 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 showing the greatest change (Figs. 29, 30). Th~ supraoccipital crest remains the largest cranial spine and is 

strongly serrate. The ridges on the cranium form a complex pattern covering the area between the eyes and the 

supraoccipital region (Fig. 28). The serrate supraorbital 

23 

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 interopercular 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 subopercular 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 posteriorly directed, elongate serrate spine. Apparently 
a 

b 

opercular 

interopercular 

24 

Figure :!:1. L.lnae of Pronol0J.lrammu eO/l from the eastern 

Pacific Ocean: a) .1.6 mm: bl i. mm. 

these spine are resorbed in the late larval or early 

juvenile period butpecimen' to de cribe the e events 

are lacking. 

Pigmentation-Pigmentation i' present on the malle t larvae examined (3.1 mm) and remain e sentially 

unchanged during larval development. Two spots are 

pre ent on the ventral midline of the caudal peduncle, 

one just po terior 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 extend along the bases of 

several rays. Dor ally 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 posttemporal regions on a 9.5-mm Prontogrammus aureorubens larva. 
a 

b 

c 

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 26.-La r va of Hemanthias peruanus from the eastern Pacific Ocean, 9.3 mm. 

posttemporal 

opercular 1 supracleithrum 

~ subopercular 

preopercular 

f,:ope rculor 

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 surface 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 fi n 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 serrate, but the interopercular spine is not serrate. There is 

no protruding supraoccipital crest, neither the frontal 

26 

posttemporoi 

suprocieithrufTj 

subopercuior 

interopercuior 

Figure 28.-Spine-bearing bones of the opercular and postterrlporal 

regions on a IO.3-mm Hemanthias vivanus larva . 

ridge over the eye nor the fin spines is serrate. Anthias 

develops more extreme larval characters. The preopercular and interopercular spines are stronger and 

both are serrate. The ridge above the eye has several serrae. In some species there IS a nonserrated protruding 
Table 4.-Meristic character development of larvae of Hemanthias vivanus. Specimens 

between dashed lines are undergoing notochord flexion. For the caudal rm, "P" and "S" 

indicate primary and secondary fin rays. 

Body Branehio- Pee- Pel- Caudal fin 

length stegal Verte- toral vie Dorsal Ventral Anal Dorsal fin Gill 

(mm) rays brae fin fin S P P S fin Spines Rays rakers 

3.2NL I 

3.3 I 

3.5 I 

3.6 I 

3.6 I 

3.7 I 

3.7 I 

3.7 I 

3.7 I 

3.8 I 

3.8 I 

3.8 I 

3.8 2 III 

3.9 I 

3.9 I 

3.9 I 

4.0 I 

4.0 I 

4.1 II 

4.1 2 III 

4.1 I II 

4.1 I II 

4.2 I II 

4.2 I II 

4.2 I II 

4.2 I II 

4.3 3 I III 

4.4 I III 

4.4 I III 

4.4 3 3 I III 

4.5 3 3 I III 

4.7 2 3 I III 

4.7 3 2 III 

4.8 4 3 III 

4.9 4 3 III 

4.9 4 4 IV 

4.9 5 4 IV 

4.9 4 3 IV 

4.9 5 5 1,1 IV 2 

4.9 6 3 I III 

5.0 6 6 I I V 2 

5.0 6 6 I II V 2 

5.2 6 5 I) I V 

5.3 6 5 1,1 II VI 3 

5.3 6 5 1,1 I IV 

5.4 6 12 I) II V damage 

---------_.---- ----------------------------------------------------------------------

5.5 6 15 I) II IX 5 

--------------- ----------------------------------------------------------------------

5.6SL 7 20 1,1 II IX 9 

5.6 7 14 1,2 II VIII 7 

5.7 7 18 1,2 II,l IX 8 

5.8 7 16 1,2 II,l IX 8 

5.8 7 16 1,2 II VIII 8 

5.9 7 15 1,1 II IX 7 

6.1 7 14 1,1 II VIII 8 

6.3 7 18 1,2 II) X 9 

6.3 7 18 1,1 II X 8 

, 6.3 7 23 11 1,4 8 6 II,9 X 12 

'6.4 7 18 1,2 II,l X 8 

6.5 7 20 .1,2 II.1 IX 9 

, 6.5 7 22 8 1,3 8 6 II,9 X 9 10 

'6.8 7 23 1,2 II,l X 11 

'6.8 7 20 1,2 2 3 II,l X 11 

'6.8 7 26 14 1,5 9 8 II,9 X 14 13 

, 7.0 7 18 1,2 II) X 9 

27 
a 

b 

c 

Table 4.-Continued. 

Body Branchio- Pec- Pel- Caudal fin 

length stegal Verte- toral vic Dorsal Ventral Anal Dorsal fin Gill 

(mm) rays brae fin fin S P P S fin Spines Rays rakers 

'7.1 7 26 14 1,5 8 7 1l,9 X 14 13 

'7.3 7 26 16 1,5 9 8 1l,9 X 14 14 

'7.3 7 23 2 1,2 6 4 1l,1 X 12 

'7.7 7 26 17 1,5 9 8 1l,9 X 14 15 

, 7.7 7 26 17 1,5 9 8 n ,9 X 14 15 

'Scales present. 

Table 5.-Body proportions of larvae of Hemanthias uiuanus. Specimen 

between dashed lines are undergoing notochord flexion . 

Percent of standard leng:!h 

Body Pre- Caudal Depth 

length Total Body Eye Head nout 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 32 

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 ll6 41 15 

5.0 ll6 38 13 

5.0 ll3 41 15 

5.2 ll6 37 12 

5.2 118 39 13 

5.4 123 40 15 

5.5 117 44 16 

5.7 121 38 13 

6.0 118 43 16 

6.0 122 42 14 

6.3 122 39 13 

6.5 124 40 14 

6.8 117 33 13 

7.0 124 42 14 

7.8 124 37 14 

8.0 113 40 11 

8.1 122 40 15 

9.2 121 35 12 

43 

39 

48 

40 

41 

43 

43 

40 

46 

44 

44 

42 

38 

43 

42 

38 

41 

37 

28 

16 61 12 35 

13 58 11 30 

15 60 14 38 

12 55 12 31 

16 60 13 31 

15 63 14 30 

15 62 14 36 

15 58 13 31 

13 65 16 35 

15 60 14 36 

13 60 14 31 

14 61 14 32 

14 61 13 29 

14 61 14 32 

14 59 1:3 29 

11 54 11 30 

14 60 11 32 

14 55 12 28 

supraoccipital crest. The fin spines are strong and t 

third dorsal spine and the pelvic spine are elonga 

Some species develop ctenoid scales while retaining Is 

val characters. Pronotogrammus develops more extren 

larval characters including a strong protruding SUpli 

occipital crest and heavy opercular armature. These I 

vae, however, are not scaled. Among American anthiim 

Hemanthias exhibits the most extreme development 

larval characters. The head is armored with a serra 

supraoccipital crest, numerous serrate ridges on the fro 

tals and parietals, and heavy serrate spines on bones 

the opercular series. Ctenoid scales develop during tJ 

larval period. In Hemanthias uiuanus larval anthii 

characters reach their apex with the development of s€ 

rations on several dorsal and anal fin spines. 

Figure 29.-Development of the supraoccipital crest of 

Hemanthias vivanus: a) 4.3 mm; b) 5.8 mm; c) 6.6 mm. 
Figure 30.-Details of the head spination on a Hemanthias vivanus larva, 7.2 mm. 

29 
Subfamily Epinephelinae 

The Epinephelinae is represented by three genera in 

Ameri can waters from both oceans. These are Paranthias 

.. \lith 1 species, Mycteroperca with 13 species, and Epinephelus 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 (Kendall 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 

subge.lera. 

Larvae of several species of Epinephelus have been illu trated and described. The species are outstanding in 

having the econd 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; 

mith 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. 

parta (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 Epinephelus 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 epinepheline. The other two do not have elongated fin 

spine ; in fact, the spinous dorsal and pelvic fins develop 

after the oft 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 posiblv aciaenid and the other a haemulid, judging from 

the number of fin elemen ts and general larval 

morphology. Presley (1970) described Epinephelus 

nil eatll.' larvae from the Florida Straits. Smith (1971) illu. trated a larval Epmephelus from off the Carolinas. 

Fowler (19-1-1) described a new genus and species (Serriha taperea exu/) from a larval fish from the eastern Pacific. As Heemstra (197-1) ha shown, this is probably a 

I nal Epmepheiu." po ibly Epinephelus panamensis. 

~ ourmnnOlr (1976) illu ·trated and briefly described three 

pllwphehne larvae from the outhea tern Pacific . Kend II nd rahay (in pre's) described a larval Go'tio30 

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 junction 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 characteristics 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. Separation 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 
.. I r 

. 

, . 

-.. 

• 
characters include characteristic skull crests and the 

presence of a postocular process (Smith 1971). 

As discussed earlier, larvae of several species of Epinephelus from around the world have been described 

(e.g .. Bertolini 1933; Mito et a1. 1967; Fourmanoir 1976). 

Presley (1970) described larvae he assigned to Epinephelus niueatus from Florida. His description is rather 

brief, and considering the similar appearance of all epinepheline 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 until 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 Xl, 

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. moria or E. guttatus. The other specimen (Fig. 33) is from the eastern 

Pacific with XI, 19 dorsal fin elements and ill, 9 anal fin 

elements making it most likely either E . dermatolepis 

dermatolepis or E. alphestes multiguttatus. Although 

these specimens are in different subgenera, no characters 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 elongate, 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 proFigure 32.-Larva of a member of the Epinephelus striatus species group from the northwestern Atlantic Ocean, 

7.6 mm. 
, . 

, 

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

8.4 mm. 

truding, serrate spines. The pigment pattern consists of a 

few melanophores on the surface of the optic lobes, some 

33 

on the lateral surface of the gut, and a large blotch on the 

caudal peduncle. The caudal peduncle blotch is var-
iously placed, from along the ventral midline to laterally 

on the lateral line. It tends to become internal on larger 

specimens. The elongate dorsal and pelvic spines are variously pigmented but the pigment is absent on many netted 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 posterior spines of the dorsal fin undergo similar transformation. 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 larval 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 transformation may occur. 

The bones in the opercular region of a 10-mm Epinephelus sp. (possibly E. niueatus) 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. 

pesttemperal 

Figure ;l~ .Spine-bearong bone, of the opercular and posttempora l 

regions on a 1 O.2-mm Epinephe ius niveatus larva . 

Mycteroperca -This is the second most speciose epinepheline genus with eight species in the Atlantic and 

five in the Pacific Ocean (Table 1). They are distinguished from the other two epinepheline genera by having 10 to 1:3 anal rays and straight, parallel lateral skull 

crests that join the rim of the orbit anterior to the midorhital area (Smith 1971). 

34 

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 M ycteroperca. These larvae had the full suite of characters of epinepheline larvae and, on the basis of larval 

characters, could not be separated from other epinepheline 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 ridg 

above the eye and protruding, serrate spines in the post 

temporal region. They had a few pigment spots on thl 

dorsal surface of the optic lobes of the brain, some on th 

lateral surface of the gut, and a large blotch on th 

caudal peduncle. Some pigment was present on th 

membranes of the elongate fin spines. 

Mycteroperca microlepis (Figs. 35, 36).-The meristic characteristics of Mycteroperca microlepis fall 

within the ranges for several species of M ycteroperca 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. 

mlcrolepis have been caught in the Middle Atlantic 

Bight. 

In plankton samples from as fa r north as Cape Hatteras, 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 epinepheline larvae taken off North Carolina. Larger larvae 

and juveniles (12-35 mm) from nearshore and estuarine 

collections along the Atlantic coast from Florida to ew 

,Jersey were examined to note the development of 

juvenile pigment and the regression of the elongate 

spines of early larvae . 1 0 other species of Mycteroperca 

could be identified in the samples with certainty. 

Meristic element development (Table 6)-Development 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 ossified, the pelvic fin has one spine and one ray, and there 

are three branchiostegal rays and two gill rakers. Between 5 and 6 mm, the notochord is flexing and most 

meristic features are forming ossified elements. The ossit'jed vertebrae increase from 4 to 18, forming from anterior to posterior except for the urostyle . Several vertebrae 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 notochord becomes fully flexed at 6 mm. When notochord 

t1exion is complete, by 7 mm, most meristic features are 

well formed, approaching adult counts. Vertebrae increase to 22 with the urostyle forming prior to the 2 or 3 

vertebrae anterior to it. The pectoral fin increases to 12 

rays and the pelvic fin attains its adult complement of 1 
spine and 5 rays. The anal fin increases rapidly having 2 

spines and 11 rays. The first soft ray of the anal fin 

becomes the third spine later in development, as is characteristic 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 addition 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 peduncle depth also increase between 4 and 6 mm. The greatest 

body depth remains between 29 and 35% of BL throughout 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 decreasing 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 somewhat 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 riear 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 Mycter35 

operca microlepis appear similar to those of other epinephelines. The pre opercular 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 preopercular 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 supracleithrum 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 American 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 distinctions 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 larval characters with epinephelines but has less variation 

in dorsal spine length and a stouter body, both reminiscent 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 armature 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. 
a 

b 

--- ----- • 

------------.. 

J 

o ~ , 

~ 't" 

.. 

o· ..... ) 

------ ~/ 

36 

----
c 

d 

Figure 35.-¥oung stages of Mycteroperca microlepis from the northwestern Atlantic Ocean: a) ~.O mm; b) 7.~ mm; c) 11.2 

mm; d) 22.6 mm. 

Table 6.-Meristic character development of larvae of Mycteroper"a microlepis. Specimens 

between dashed lines are under going notochord flexion. For the caudal fin, "P" and "S" 

indicate primary and secondary fin rays. 

Body Branchio- Pec- Pel- Caudal fin 

length stegal Verte- toral vic Dorsal Ventral Anal Dorsal fin Gill 

(mm) rays brae fin fin S P P S fin Spines Rays rakers 

4.0NL 3 1,1 ill 2 

4.6 4 1,2 ill 3 

5.1 5 4 1,2 1 2 ill 6 

5.6 6 6 1,3 5 5 III 7 

----------------------------------------------------.----------------.---------------

5.3 5 4 1,3 4 4 ill 6 

5.3 6 6 2 1,3 6 6 IV 8 

5.3 6 8 2 1,3 7 7 1\ 10 

5.5 7 18 6 damage 9 8 1,6 V 9 

-------------------------------------.---------.------.------------------------------

6,OSL 7 20 9 1,4 9 8 IT,9 X 11 1+9 

6.1 7 19 6 1,4 9 8 1,6 \1 9 

7.0 7 22 12 1,5 9 8 2 IT,ll X 15 1+10 

7.4 7 24 12 ' 1,5 1 9 8 2 IT,ll X 15 ~~ 9 

7.9 7 24 14 1,5 3 9 8 3 n.13 X 19 3+10 

8.1 7 24 15 1,5 3 9 8 2 IT.12 X 1b 3+10 

9.2 7 24 17 1,5 4 9 8 5 n,ll X 1w 1+12 

9.8 7 24 16 1,5 5 9 5 IT.12 X 1 4+11 

37 
Table 7.- Body proportions of larvae of Myctoperca microlepis. Specimens between dashed lines are undergoing 

notochord flexion . 

Percent of standard length 

Body Caudal Depth Second Pelvic Preoperlength Total Body Eye Head Snout Preanal peduncle at spine of fin cular 

(mm) length depth length length length length depth anus dorsal fin spine spine 

4.SNL 104 30.4 10.9 32.S 9.6 52.2 7.6 15.2 39.1 0.43 

----------------------------- -----------------------------------------.-------------.---------------------

5.3 lOS 32.1 11.3 32.1 9.4 50.9 8.5 18.9 49.1 0.58 

5.3 109 34.0 11.9 35.8 10.6 56.6 8.3 20.8 69.2 54.7 0.75 

5.3 113 34.0 12.1 35.8 12.1 54.7 9.4 26.4 58.5 0.94 

5.5 110 36.4 12.7 40.0 13.6 56.6 10.2 25.5 0.91 

5.6 107 32.1 ILl 35.7 10.9 55.4 7.9 19.6 0.55 

5.9 112 32.2 10.8 42.4 64.4 10.5 22.0 1.02 

---------_.--.--- -----------------------------------------------------------------------------------------

6.0SL 120 

6.1 120 

7.0 120 

7.4 122 

7.9 120 

8.1 121 

/l.3 119 

9.2 123 

9.4 121 

9.8 123 

12.0 123 

13.6 125 

14.0 131 

16.8 123 

17.6 122 

19.3 115 

2Ll 122 

21.5 120 

24.4 125 

35.4 120 

opercu,ar0 " 

,,~ 

.J 

\J 

~/ 

u 

36.7 

34.4 

34.3 

33.8 

32.9 

34.6 

33.7 

33.7 

35.1 

33.7 

35.0 

33.1 

33.6 

32.7 

31.8 

3Ll 

31.8 

30.2 

32.8 

28.5 

13.0 38.3 11.5 

12.3 41.0 13.0 

12.1 37.1 11.6 

11.9 39.2 11.5 

11.9 38.0 10.8 

9.9 37.0 11.6 

12.0 38.6 13.3 

12.0 38.0 12.0 

12.8 38.3 10.6 

11.2 39.8 14.3 

11.7 38.3 10.8 

9.6 37.5 11.8 

10.7 37.9 11.4 

10.1 39.3 9.5 

11.4 35.3 8.5 

10.4 36.3 9.3 

10.9 31.3 13.7 

10.7 37.2 10.7 

ILl 42.6 11.1 

9.0 36.2 9.0' 

posttemporal 

(~ 

\ { supracleithrum \ 

'v 

subopercular 

Figure 36.-Spine-bearing bones of the opercular and posttemporal 

regions on a 9.8-mm Mycteroperca microlepis larva. 

Subfamily Grammistinae 

Larvae of three genera, Liopropoma (larvae of Pikea 

may be included here), Rypticus, and Pseudogramma, 

38 

60.0 11.5 23.3 61.7 61.7 0.93 

67.2 10.2 26.2 59.0 0.92 

61.4 11.6 24.3 54.3 61.4 0.99 

60.8 ILl 21.6 62.2 1.15 

60.8 11.9 24.1 60.8 1.04 

59.3 13.6 24.7 59.3 58.0 0.99 

57.8 12.0 25.3 65.1 67.5 1.33 

57.6 12.0 23.9 59.8 64.1 1.02 

59.6 12.8 25.5 58.5 6.3.8 1.06 

66.2 112 26.5 60.2 65.3 1.02 

57.5 13.3 28.3 49.2 55.0 1.00 

58.1 11.8 27.9 39.7 42.6 0.92 

62.9 10.7 25.7 39.3 47.9 

61.9 11.9 25.6 40.5 39.3 0.83 

58.0 10.2 24.4 40.3 46.0 0.91 

58.5 10.9 24.9 21.3 20.2 0.62 

60.7 10.9 27.0 20.4 29.4 0.52 

52.1 11.2 256 20.5 17.2 0.65 

63.5 ILl 32.8 13.5 14.8 0.33 

61.6 10.7 27.1 110 12.1 0.14 

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 re lationships 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 synonymized with Liopropoma. Two genera, Joboehlkia 

and Flagelloserranus, described from preadult material, 

apparently belong with this group. Pikea has three northwestern 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 

a 

/ 

b 

\. 

/ 

--... 

.. 

Figure 37.-Lanae of Liopropoma sp. from the north"l',tern Atlantic Ocean: a) 1i.:1 mm; b) 7.n 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 larval spination to those of Liopropoma, although specimens of the same size as those observed of Liopropoma 

40 

were not available for tudy, so compari on may be invahd. 

Pseudogramma.-Rhegma has been synonymized 

with Pseudogramma (Go line 1960) and at times considered a serranid, a grammistid, or a pseudogrammid 

(Kendall 1977). Larval and other characters indicate 

that this genus is closely related to Rypticus and 
posttemporal 

opercular 

suprocleithrum 

P'""-" o, ) 

O,Ie'OP"",'O, 

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 epinepheline serranids. Pseudogramma contains one species 

from each of the following areas: the central Pacific, the 

eastern Pacific, and the western Atlantic. They have incomplete 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 Pseudogramma 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 several species, none of which could be identified using larval 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 synonymized 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 

pre caudal and 2 caudal), 4 branchiostegal rays, 15 pectoral 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 dorsal 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 RypticU8 sp. from the northwestern Atlantic Ocean, 6.6 mm. 

41 
a 

\ 

b 

c 

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 II. .\1t'ri.,tic charucter de\elopmt'nt or l<in U(' or P ('ud"Jlrumml/ Rr I:on I, JI( 1m n 

bet"'een dUbhed line., are undl'rgoing notothord n(, ion ~ or thl' ( ud I fin,"P" nd .. . inditate primar.\ and .,econdar) fin ru~., . 

Body Branehio· Pee· Pel. Cauda' fin 

length .,legal Verte· toral vtc Dorsal \'entral nal I rsalfin GIll 

(mm) ray., brae lin lin P I' s lin plm Hn ra 

3.3 NL 4 10+2 If) 2 

-------------------------------- ..... --_._ .................... -_. 

:1.6 6 10+ 15 II 

4.1 6 10+8 16 4 6 II 4 

4.1 6 10+9 16 6 6 II <4 

4.2 6 10+9 16 6 6 II 4 

--------------------- ........................................................ ~ .... _----_ ........ __ .. 

4.28L 7 10+14 16 

4.3 7 10+11 15 

4,4 7 10+16 15 

4.5 7 10+12 15 

4.6 7 10+14 15 

4.7 7 10+16 15 1,3 2 

5.3 7 10+16 1.5 3 2 

5.6 7 10+16 15 1,4 3 

5.9 7 10+16 15 1,4 3 

6.7 7 10+16 15 1,5 3 

7.R 7 10+16 15 1,5 3 

9.6 7 10+16 15 1,5 3 

10.3 7 10+16 15 1,Ii 3 

5·mm specimens (branchiostegal rays, vertebrae, pee· 

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 Pseudogramma gregoryi is quite different from that of other 

erranids in that the pectoral fin forms before other fins 

and the pelvic fin forms last. The third anal pine 

develops as a spine rather than as a ray that transform 

into a spine as it does in other serranids. 

Body proportions (Table 9)-A sene of larvae from 3.9 

to 11.4 mm was measured to trace changes in body proportions during larval development. Due to the limited 

number of specimens available. some partly damaged 

larvae had to be u.ed for these measurements; therefore. 

some measurements may not represent actual body 

• hape . As the notochord f1exes. total length increases 

from about 110', to about 125' ( of BL. caudal peduncle 

depth increases from 6-10' ( to about l·V ( of EL. and the 

preanal length increase~ from about 45{r to about 55 cot 

BL. The rest of the proportion::; remain rather con~tant 

mer the ize range at hand. The body depth is about 25 ~ 

of BL (23.2-:n.7'cl and the depth at the anus is only 

slightl: less (22.5-29.f>',). The measurement: a~soclated 

with the head are: eye. 8.3 to 11.6'( of BL; .nout. 4.-; to 

lO.O', of BL; and head length. 26.6 to :~-;.:' c of BL. P,eudogramma gregory; then. i~ shallower bodied and has a 

' horter head and preanal length than ot her ,erranid~. 

PigmentatIOn-Larvae of PscudoRramma [!rt! 0 I are 

notable. as are tho~e of LlOpropoma and R\ptlcu , tor 

their lack of pigment. rhe only Ulldy pl!,!lJ1ent 01 P 

grl'''Or,' I dunng lan'al de\'elopment )Cl'ur. on the dOf 

lateral surtace of the gut cant\ ~'on1l' pi 'ment de\el p 

on the membrane a:'t-ll'iated With the elon at d nd 

Spine 01 the dOft-al lin .md al. 0 on the tip~ ofth p toral 

9 

9 

Ii 

!J 

9 

9 

9 

9 

9 

9 

9 

1.13 1\ l7 7 

Ii !J II i 

Ii 2 I,It> \111 II< I ~ 

10 Il I +' 

1,16 \ III l,k , 2 IlI,15 \111 ." 7 

'2 III,lb \ III HI 

8 :3 IIJ, IS \III !!f 3+7 

3 lII,15 \ !II I~ +7 

3 1I1,1.~ \ III I (+.101 

8 a Ill,I5 \ !II I" I , 3 111,15 \ !II I G 

8 :1 111.15 \ III III +\0 

lin rays. ::,mce many 'pe<,imt'n~ were bleB hed, hud 

broken dorsal ~pine, or were In othem I poor DC \tlOn, 

it was not possible to determine exactly when thl pi . 

ment first appeared. It wa~ only pre cnt In lar er I r\ a 

among tho~e examined. 

\ d urlnc!: 
Table 9.-Body proportions of larvae of Pseudogramma gregoryi. Specimens 

between dashed lines are undergoing notochord flexion. 

Percent of standard leng!h 

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 

-- --- ------------------------------ - --- - - -- ----- --------- ------------- - ----

p 'eopercular 

4.3 110 

4.3 112 

4.6 112 

4.5 SL 120 

4.5 118 

4.8 113 

4.8 116 

4.8 118 

5.0 118 

5.2 ll7 

5.3 ll2 

5.7 122 

5.9 ll9 

6.0 121 

6.1 120 

6.0 119 

7.2 121 

8.0 ll8 

9.7 125 

9.8 125 

10.4 122 

11.4 124 

25.0 

26.1 

25.7 

26.8 

23.9 

27.6 

27.6 

26.3 

26.3 

31.7 

27.4 

26.4 

26.6 

26.0 

26.8 

25.0 

26.7 

25.2 

26.0 

6.3 

28.3 

25.4 

~mpOCOI 

10.2 

11.6 

9.5 

ll.3 

9.9 

10.5 

9.2 

9.2 

10.0 

9.8 

10.7 

9.9 

8.5 

10.4 

9.3 

8.3 

9.5 

9.4 

.8.4 

9.0 

9.0 

8.8 

\j supracl eithrum 

Fi ur.· II ·Spine-bearing bones of the opercular a nd posttemporal 

rrj(ions on a IIU-mm Pseudogramm a g rego'1'i la rva. 

held-collee ted specimens; but the bases of these elong te pll1e~ are larger than those of other spines and this 

I u,., Il.1l ill recognizmg these larvae. 

l. I he pectoral fins develop rays earlier and are larger 

t h f1 111 other ,.,erranids. In other serranid genera the pec30.9 

30.4 

31.1 

32.4 

29 .6 

27.5 

27.6 

30.3 

31.3 

37.8 

31.0 

29.7 

26.6 

34.4 

29.9 

28.1 

29.5 

29.9 

31.8 

28.8 

30.7 

32.0 

44 

8.8 44.1 8.8 25.0 

8.7 46.4 10.1 26.1 

50.0 9.5 23.0 

7.0 52.2 14.1 26.8 

9.9 46.5 11.3 22.5 

6.6 42.1 13.2 26.3 

46.0 13.2 26.3 

9.2 50.0 14.5 23.7 

10.0 47.5 12.5 23.8 

9.8 59.8 11.0 24.4 

7.1 50.0 16.7 26.2 

7.7 50.5 15.4 25.3 

5.3 46.8 13.8 23.4 

8.3 45.8 14.6 25 .0 

8.2 48.5 15.5 25.8 

6.3 44.8 12.5 24.0 

6.7 50.5 16.2 27 .6 

4.7 50.4 14.2 27.6 

9.1 53.2 13.0 27.3 

7.1 53.8 13.5 28.2 

6.6 56.0 14.5 29 .2 

7.7 53.6 14.4 26.5 

toral fins form last; however, in these genera (exceptLI 

propoma) they form while only a few dorsal spines al 

caudal fin rays are developing. 

4. The body shape is little elevated and the caudal j: 

duncle is deep, producing a tubular-shaped body simil 

to that seen in some labrid and scarid larvae. Serran 

larvae in the subfamilies Anthiinae and Epinephelin 

are deeper bodied than grammistines. Only larvae 

serranines have the depressed condition of grammistillE 

These three genera have been considered members 

separate families or subfamilies; but on the basis of la 

val morphology and other characters (Kendall 197€ 

they appear more closely related to each other. Sever 

other genera have been aligned with those described he 

but their larvae are unknown, so an accurate assessmel 

of the relationships within this group based on larval d 

velopment is not possible at present. However, Hubl 

and Chu (1934) illustrated late larvae of Diploprion b 

fasciatum, a genus considered a grammistid by Rands 

et a1. (1971). Their specimens had greatly elongated fle 

ible second and third dorsal spines. Jeboehlkia gladife 

described from a small specimen by Robins (1967), hI 

an elongated but stiff first dorsal fin spine and is pro I 

ably a small specimen of a species closely related to tho: 

described here. Fourmanoir (1976) illustrated and brief 

described larvae of Grammistes sexilineatus an 

Aporops bilinearis, both with an elongate dorsal spir 

and preopercular spines similar to the grammistines d, 

scribed here . The unique first pterygiophore of gran 

mistine and related fishes (Kendall 1976) may help SUI 

port the elongate larval dorsal spines, as the enlarg€ 
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 between 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 described (Mansueti 1958, 1964). Since they develop in estuarine 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 melanophores scattered over the body, and the lack of heavy armature in the opercular area. 

Serraninae larvae demonstrate various degrees cf 

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 pigment 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 increased 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, indicating 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. Epi45 

nepheline larvae possess a suite of specialized character 

that apparently enable them to have a long pelagic existence. They have extremely elongate and serrate second dorsal and pelvic spines. The preopercular also has a 

long serrate spine at its angle and there is a serrate supraorbital crest. Pigment is generally confined to the ventral 

area of the caudal peduncle, the hindbrain, t he lateral 

surface of the gut cavity, and the membranes of the elongate 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 epinepheline awaits further analysis of adult morphology. 

Since there are no other larvae with only some of the epinepheline 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 indicate that they are part of a single lineage. The most outstanding 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 Pseudogramma. In summary, it appears that these fishes 

could have developed from fishes with serranine-like larvae 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 epinephelinegrammistines I. Kendall 1976) so its larvae may show partial 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, Anthimae, 

and Epinephelinae and the genera Lwpropoma, Rypticus, and Pseudogramma, here grouped as the subfamily Grammistinae. Within the serramnes and anthllne . 

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 wa seen 

The four groups of American erranid larvae can be 

partially characterized as follows (Figs. 42, 43). 

Serraninae- Body proportions how rather direct development. There are no elongate pine in the opercular region. rather a series of blunt pOints. The fin pine. 

are thin and only slightly elongated in orne. ;\10 t larval 
a 

/ 

b 

\ > 

.@ .' 

~ -

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 microlepis, 7.4 mm); d) Gr ammistinae (R ypticus sp., 6.6 mm). 

46 
a 

b 

c f7 

dO 

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 positions along the ventral midline. 

Anthiinae- These are deep-bodied larvae with produced 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 serrate. 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 I>igment 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 

47 

engaging in useful discussions about this work: Phillip C. 

Heemstra, CSIRO, Cronulla, .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 J ersey Marine Science 

Consortium, and H. Geoffrey Moser, National Marine 

Fisheries Service, La Jolla , gave much encouragement 

and helpful advice. 

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BARLOW, G. W. 

1975. On the sociobiology of some hermaphroditic serranid fishes, 

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1977. Revision of the sea basses of the genus Diplectrum (Pisces: 

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CLOTHIER, C. R 

1950. A key to some southern California fishes based on vertebral 

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COURTENA Y, W. R, JR 

1967. Atlantic fishes of the genus Rypticus (Grammistidae). Proc. Acad. Nat. Sci. Phila. 119:241-293. 

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1918. Shore-fishes. Rep. Dan. Oceanogr. Exped. 1908-1910. 

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1963. Patterns of the ramus lateralis accessorius and their 

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GINSBURG, I. 

1952. Eight new fishes from the Gulf coast of the United States, 

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1960. A new Hawaiian percoid fish, SuttonLa lmeata, with a discussion of its relationships and a definition of the family Grammistidae. Pac. Sci. 14:28-38. 

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HASTINGS, R W. 

1973. Biology of the pygmy sea bass, 'erraniculus pumllio (Pi_ ces: 

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1974. On the identity of certain eastern Pacific and Caribbean 

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1970. Artificial spawning of black sea bass, Centropnstes striatus 

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HOLLISTER, G. 

1934. Clearing and dyeing fish for bone study. Zoologica (N.Y.) 

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HUBBS, C. L. , and Y. T. CHU. 

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1960. Fauna Japonica Serranidae. (Pisces). Tokyo News Servo 

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KENDALL, A. W., JR. 

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1970. Flagelloserranus a new genus of serranid fishes With the deSCription of two new species (Pisces, Percomorphl). Dana Rep. 

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MANSUETI, R. J. 

1958. Eggs, larvae, and young of the striped bass, Roccus saxatilus. Md. Dep. Res. Educ. Contrib. 112:1-36. 

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1961. The comparative anatomy of the scales of the serranld 

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48 

RAFFAELE, F. 

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Appendix Table I.- Data associa ted with lar val fish used for illustrations. Institution abbreviations: NE: NMFS, Northeast 

Fisheries Center ; SW: NMFS, Southwest Fisheries Center ; SE: NMFS, Southeast F isheries Center; UNe: University of North 

Carolina . 

Figure 

l a 

b 

2 

3a 

b 

4 

5a 

b 

6a 

b 

7 

c 

d 

8a 

b 

9 

lOa 

b 

11 

c 

d 

12a 

b 

13 

c 

d 

14a 

b 

c 

15a 

b 

16a 

b 

17a 

b 

c 

18 

. 19 

20a 

b 

21 

Species 

Centropristis striata 

Centrapristis striata 

Centropristis striata 

Paralabrax sp. 

Paralabrax sp. 

Paralabrax sp. 

Diplectrum sp. Type 1 (AtL) 

Diplectrum sp. Type 1 (AtL) 

Diplectrum sp. Type 1 (Pac.) 

Deplectrum sp. Type 1 (Pac.) 

Diplectrum sp. Type 1 (Pac.) 

Diplectrum sp. Type 1 (Pac.) 

Diplectrum sp. Type 1 (AtL) 

Diplectrum sp. Type 2 

Diplectrum sp. Type 2 

Diplectrum sp. Type 2 

Serranus sp. 

Serranus sp. 

Serranus sp. 

Serranus sp. 

Serranus sp. 

Hypaplectrus sp. 

Hypaplectrus sp. 

Hypaplectrus sp. 

Hypaplectrus sp. 

Hypaplectrus sp. 

Serraniculus pumilia 

Serraniculus pumilia 

Serraniculus pumilia 

Plectranthias garupellus 

Plectranthias garupellus 

Anthias sp. Type 1 

Anthias sp. Type 1 

Anthias sp. Type 2 

Anthias sp. Type 2 

Anthias sp. Type 2 

Anthias tenuis 

Anthias sp. Type 3 

Anthias gardensis 

Anthias gardensis 

Anthias sp. Type 1 

Length 

(mm) 

4.7 

8.3 

10.6 

6.0 

9.0 

9.5 

5.8 

10.0 

4.3 

5.0 

6.1 

13.0 

10.3 

4.5 

8.4 

9.9 

3.7 

5.0 

5.5 

9.4 

9.7 

3.4 

4.7 

5.7 

8.5 

12.5 

3.8 

5.8 

55.3 

5.5 

7.0 

3.8 

5.3 

4.7 

5.7 

8.4 

6.7 

5.1 

5.2 

6.0 

10.2 

Institution 

NE 

NE 

NE 

SW 

SW 

SW 

NE 

NE 

SW 

SW 

SW 

SW 

NE 

SW 

SW 

SW 

NE 

NE 

NE 

NE 

NE 

SE 

SE 

SE 

SE 

SE 

NE 

NE 

NE 

NE 

NE 

NE 

NE 

NE 

NE 

NE 

SE 

NE 

SW 

SW 

NE 

49 

Cruise 

D-67-4 

D-71-5 

D-67-4 

6509B 

S-5509 

C-51 

D-67-4 

D-67-16 

5706S 

5706S 

5706S 

5706S 

D-67-4 

P-5412 

5612-H 

ETP 

D-71-5 

1 

D-66-3 

D-68-1 

D-66-12 

Station 

FF-5(1) 

N-2p 

CC-5(1) 

118.39 

Lat. 

(N) 

Long. 

(W) 

Date 

Day/Mo/Yr 

79°56' 12 5 67 

75°55' 29 4 71 

115°24' 17 10 65 

(CaICOFI sample - no precise data) 

130.30 

GG-4(1) 

HH-5(2) 

107G02 

107G02 

107G02 

143G100 

CC-5(1) 

157.10 

160G30 

47.244 

M-6 

4 

M-5(2) 

NN-2(2) 

L-4(1) 

29°42' 

29°42' 

29°42' 

29°19' 

35°07' 

35°03' 

35°11' 

27°11' 

113°49' 10 8 53 

114 °07' 

114°07' 

114 °07' 

109°14' 

74 °55' 

75°10' 

75°07' 

80°03' 

II 5 67 

24 10 67 

20 6 57 

20 6 57 

20 6 57 

II 6 57 

14 5 67 

4 7 54 

3 7 56 

27 8 67 

5 5 71 

II 4 72 

20 4 66 

4 2 68 

30 9 66 

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

Reared at "lMFS, Miami, Fla. from eggs collected nearby 

Reared at f\'MFS, Miami, Fla. from eggs collected nearby 

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

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

D-67-16 

D-67-16 

D-71-3 

D-67-16 

D-68-1 

D-66-5 

D-71-5 

D-68-1 

D-71-5 

D-68-1 

BLM 

D-71-5 

5706S 

5706S 

D-68-1 

DD-4(1) 

DD-5(2) 

SA-2 

NN-4(1) 

PP-2(1) 

N-4(2) 

BB-7 

DD-6(l) 

M-4 

EE-6 

IV-2 

BB-8 

109G55 

115G40 

EE-6 

32°55' 

32°47' 

29°51' 

32°36' 

35°13' 

32°08' 

78°51' 

78°44' 

81 °10' 

78°34' 

75°12' 

79°10' 

21 10 67 

21 10 67 

3 9 71 

29 10 67 

4 2 68 

24 5 66 

5 71 

29 1 68 

5 5 71 

30 1 68 

30 5 76 

5 71 

113°04' 18 6 57 

112°44' 17 6 57 
Appendix Table I.-Continued . 

Figure 

22a 

b 

c 

23a 

b 

24 

25a 

b 

c 

26 

27 

28 

29a 

b 

c 

30 

31 

32 

33 

34 

35a 

b 

c 

d 

36 

37a 

b 

38 

39 

40a 

b 

41 

c 

d 

42a 

b 

c 

d 

43a 

b 

c 

d 

Species 

Pronotogrammus aureorubens 

Pronotogrammus aureorubens 

Pronotogrammus aureorubens 

Pronotogrammus eos 

Pronotogrammus eos 

Pronotogrammus aureorubens 

Hemanthtas vivanus 

Hemanthtas VLVanus 

Hemanthtas VLVanus 

Hemanthias peruanus 

Hemanthias VLVanus 

Hemanthias vivanus 

Hemanthias vwanus 

Hemanthias VLVanus 

Hemanthias VLVanus 

Hemanthtas vivanus 

Paranthias furcifer 

Epinephelus stnatus 

Epinephelus sp. 

Epinephelus niveatus 

Mycteroperca microlepts 

Mycteroperca microlepls 

Mycteroperca microlepis 

Mycteroperca microlepls 

Mycteroperca microlepis 

Liopropoma sp. 

Liopropoma sp. 

Ltopropoma sp. 

Rypticus sp. 

Pseudogramma gregoryi 

Pseudogramma gregoryi 

Pseudogramma gregoryi 

Pseudogramma gregoryi 

Pseudogramma gregoryi 

Paralabrax sp. 

A nthias tenuis 

Mycteroperca microlepis 

Rypticus sp. 

Centropristis striata 

Hemanthias vivanus 

Epinephelus niveatus 

Pseudogramma gregoryi 

Length 

(mm) 

4.6 

6.0 

9.0 

3.6 

7.8 

9.5 

4.2 

5.3 

6.8 

9.3 

9.3 

10.3 

4.3 

5.8 

6.6 

7.2 

8.6 

7.6 

8.4 

10.2 

4.0 

7.4 

14.2 

22.6 

9.8 

6.3 

7.0 

12.0 

6.6 

4.7 

6.1 

6.1 

10.2 

10.3 

6.0 

6.7 

7.4 

6.6 

10.6 

10.3 

10.2 

10.3 

Institution 

NE 

NE 

E 

SW 

SW 

NE 

NE 

NE 

E 

W 

NE 

NE 

NE 

NE 

NE 

SE 

S\V 

NE 

SW 

NE 

NE 

NE 

UNC 

U C 

NE 

NE 

NE 

SE 

NE 

NE 

SE 

NE 

NE 

SE 

SW 

SE 

NE 

NE 

NE 

NE 

NE 

SE 

50 

Cruise 

0-68-1 

0-68-1 

0-68-1 

5706S 

OP 

0-68-1 

0-71-5 

0-71·5 

0-71-5 

6611J 

0-71·5 

0-68-1 

0-71-5 

0-71-5 

0-71-5 

BLM-2 

5612 

0-67-4 

OP 

0-67-16 

1 

3 

(UNC7934) 

0-71-5 

0-67-4 

3 

OR II 702 

0-67-8 

0-67-4 

OR II 7739 

0-66-3 

0-67-4 

OR II 7239 

6509B 

BLM 

3 

0-67-8 

0-67-4 

0-68-1 

0-67-16 

OR II 7239 

Station 

PP-3(1) 

CC-7(2) 

AA-7 

133G40 

052 

HH-7 

LK·5 

BB-8 

BB-8 

147.30 

N-7 

HH-7 

N-7 

-6 

N-7 

1-3 Day 

157GlO 

GG-7(2) 

162 

GG-7 

4 

2 

T-9 

AA-7 

NN-4(2) 

2 

0-2-13 

AA-4(1) 

MM-4(1) 

56 

N-5(1) 

JJ-6(1) 

56 

118.39 

IV-2 

2 

AA-4(1) 

CC-5(1) 

HH-7 

GG-7 

56 

Lat. 

(N) 

26°47' 

32°54' 

33°37' 

30°19' 

35°59' 

32'39' 

32°39' 

34°13' 

34°23' 

34°13' 

Long. 

(W ) 

Date 

OayfMolYr 

79°50' 4 2 68 

78°07' 29 68 

76°47' 27 68 

74 °33' 

77°17' 

77°17' 

75°35' 

75°39' 

75°35' 

96°07' 

12 6 57 

19 11 67 

2 68 

5 5 71 

5 71 

5 71 

13 11 66 

2 68 

29 4 71 

29 4 71 

29 4 71 

10 4 75 

3 12 56 

12 5 67 

92°00' 26 11 67 

80°00' 23 10 67 

11 4 72 

27 4 72 

New River, N.C. 

Bogue d., N.C. 

26 4 72 

16 6 73 

27°46' 

20°00' 

34°33' 

29°49' 

28°19' 

26°10' 

35°03' 

34°11' 

33°20' 

30°19' 

30°54' 

20°00' 

79°56' 

80°13' 

75°44' 

80°14' 

115°24' 

96°39' 

75°08' 

77°11' 

78°25' 

80°13' 

80°00' 

80°13' 

30 4 71 

7 5 67 

27 4 72 

13 9 70 

8 67 

8 5 67 

30 7 72 

20 4 66 

10 5 67 

30 7 72 

17 9 65 

30 5 76 

27 4 72 

8 67 

14 5 67 

1 2 68 

23 10 67 

30 7 72 

,