Epinephelus marginatus ( LOWE, 1834 )

Heemstra, P. C., 1991, A Taxonomic Revision Of The Eastern Atlantic Groupers (Pisces: Serranidae), Bol. Mus. Mun. Funchal 43 (226), pp. 5-71 : 38-46

Epinephelus marginatus ( LOWE, 1834)

Figs. 12 & 13 , Plate I, Fig. 1

? Perca gigas BRÜNNICH, 1768: 65 (type locality, Marseilles, France; holotype lost?)

? Holocentrus merou LACEPÈDE, 1802: 376 (replacement name for Perca gigas BRÜNNICH, 1768 ).

Serranus gigas : VALENCIENNES, in Cuv. & VAL., 1828: 270, Pl. 33; GÜNTHER, 1859: 132; STEINDACH NER, 1877: 175.

Serranus marginatus Lowe, 1834: 142 (type locality, Madeira; no type specimens indicated; neotype MMF 3388, 200 mm. SL, Madeira).

Serranus fimbriatus Lowe, 1836: 195 , PL I, Fig. 1 (unnecessary replacement name for Serranus marginatus LOW , thought to be preoccupied by Serranus marginalis VALENCIENNES, 1828 [which was based on Holocentrus marginatus LACEPÈDE, 1802 , an incorrect spelling of Epinephelus marginalis BLOCH, 1793 ]).

Cerna gigas : DODERLEIN, 1882: 177, Pl. 1, Fig. 1 .

Serranus aspersus JENYNS, 1843: 6 (type locality, Cape Verde Islands; holotype BMNH 1917.7.1436).

? Serranus cernioides CAPELLO, 1868: 156 , Pl. 4, Fig. 1 (type locality, Lisbon; holotype lost).

Epinephelus brachysoma COPE, 1871: 466 (type locality, Rio de Janeiro; holotype ANSP 13372).

Epinephelus gigas : JORDAN & SWAIN, 1885: 388 (synonymy based on STEINDACHNER, 1877; no description); JORDAN & EIGENMANN, 1890: 359; BOULENGER, 1895; 232; BARNARD, 1927: 482; CADENAT, 1951: 191, Fig. 125.

Epinephelus guaza (non LINNAEUS): JORDAN & EVERMANN, 1896: 1154; J. L. B. SMITH, 1949: 195, Pl. 18; RIVAS, 1964: 29; C. L SMITH, 1971: 137, 1981; SÉRET, 1981: 156, fig., HEEMSTRA & RANDALL, 1984, 1986: 525, Pls. 36 & 39', TORTONESE, 1986: 785, fig.: BIANHI, 1986: 43, fig; BAUCHOT, 1987: 1311, fig; MANZONI, 1987: 67, Fig; BELLEMANS et al., 1988: 96, Pl. 11, Fig. 83; SCHNEIDER, 1990: 105. Pl. 11, Fig. 82 (labelled " Epinephelus marginatus ").

Diagnosis: Dorsal fin XI,14- 16; anal fin III,8 (1 of 89 fish counted had 9 anal fin rays); pectoral fin rays 17- 19; lateral-line scales 62-73; lateral scale series 98- 113; gill-rakers 7-10 + 14-16 (including 2- 5 rudiments on each limb). Body depth less than head length, contained 2.6-3.1 times in SL (for fish 8-62 cm. SL); head length contained 2.3-2.5 times in SL; body moderately compressed, the greatest width contained 1.7- 2.3 times in the depth. Eye diameter contained 4.3-7.9 times in head length, equal to snout length of fish 8- 10 cm. SL and about half snout length of fish 50 cm.; eye diameter greater than or subequal to interorbital width for fish 10-30 cm. SL, less than interorbital width of fish over 40 crn. SL; interorbital width 5.0-7.3 times in head length; caudal peduncle depth contained 7.5-9.5 times in SL. Caudal fin rounded (juveniles) or truncate with rounded corners (adults, Fig. 12 ); 3rd or 4th dorsal fin spines longest, longer than longest dorsal rays and contained 22-28 times in head length, the interspinous membranes distinctly incised; anal fin margin rounded, the 2nd and 3rd spines subequal in juveniles, but the 3rd spine longest in adults, 2.2- 3.3 times in head; pectoral fins fleshy, the fin length contained 1.6-2.0 times in head length; pelvic fins not reaching anus, shorter than pectorals, their length contained 1.8-2.4 times in head length; pelvic fin origin below or slightly posterior to base of pectorals.

Interorbital area convex; dorsal profile of head slightly convex. Preopercle rounded or subangular, the rear edge finely serrate, with the serrae at the "angle" slightly enlarged; adults with a shallow indentation on rear edge just above the angle; no serrae on ventral edge of preopercle; opercle with middle and lower spines distinct, the upper spine hidden by skin and scales; upper edge of operculum convex; edge of subopercle and interopercle smooth. Nostrils rounded, subequal, or rear nostril slightly larger; anterior nostril set in a short tube, the posterior edge produced into a flap. Maxilla naked, reaching to or slightly past a vertical at rear edge of eye; ventral edge of maxilla smoothly curved from distal expansion to narrow middle part (no step-like discontinuity on ventral edge of maxilla). Mid-lateral part of lower jaw with 2-4 rows of subequal teeth, the inner teeth depressible medially; premaxilla with a band of small, depressible, conical teeth; juveniles with a pair of small fixed canines (enlarged conical teeth) at front of both jaws, but in large adults (> 60 cm. SL) there are no apparent canines in the jaws. Gill-raker at angle subequal to adjacent raker on lower limb and also subequal to longest gill filaments. Mid-lateral body scales ctenoid (at least in area covered by pectoral fins); adults with numerous auxiliary Scales. Pyloric caeca 26-50.

Colour: Head and body dark reddish brown or greyish dorsally, usually yellowish gold ventrally; irregular white, pale yellowish-green or silvery-grey blotches usually visible on body and head; a black, more or less distinct "moustache" streak along maxillary groove below eye (mostly covered by upper part of maxilla when the mouth is closed); lower edge of anal fin and rear edge of caudal fin narrowly white; pelvic fins blackish distally; pectoral fins dark reddish brown or grey; margin of spinous part of dorsal fin and basal part of paired fins often golden yellow. First gill arch and gill-rakers densely covered with minute melanophores.

Description of neotype: (Measurements are given in Table 1 .) Body depth contained 2.6 times in SL; head length contained 2.5 times in SL; body width equals half of its depth; snout length 3.7 times in head; orbit 6.2 times in head; interorbital 5.3 times in head; preorbital depth 10.5 times in head; upper jaw 2.2 times in head; maxilla width 8.6 times in head; lower jaw 1.8 times in head; caudal peduncle depth 3.1 times in head; peduncle length 2.7 times in head. Caudal fin rounded. Dorsal fin with 11 spines and 15 rays; anal fin with 3 spines and 8 rays; pectoral fin rays 19 (both sides); pectoral fin length contained 1.5 times in head length; pelvic fin length contained 1.9 times in head length. Midlateral part of lower jaw with 2 rows of subequal teeth. Gill-rakers 10 + 16 (including 5 rudiments on upper limb and 3 on lower limb). Lateral-line scales 69; lateral scale series 110.

Colour in alcohol: Head and body brown, with irregular pale silvery grey spots and blotches, those on rear part of body arranged in vertical series; fins darker than body; median fins with a few indistinct pale spots; maxillary streak blackish brown.

GEOGRAPHICAL DISTRIBUTION

Epinephelus marginatus occurs on both sides of the Atlantic Ocean, throughout the Mediterranean Sea, along the west coast of Africa, and round the south coast of Africa to southern Mozambique. I have examined specimens from Madeira, the Azores, Spain, France, Italy, Greece, Lebanon, Israel, Algeria, Cape Verde Islands, Angola, South Africa, Mozambique and Brazil. Based on identifiable records of " Epinephelus guaza ", the species is also known from Egypt, Tunisia, Morocco, Mauritania, Sénégal, Ivory Coast and the Congo. According to WHEELER (1969) E. marginatus (identified as " Epinephelus guaza ") is rare in British seas. Reported from India by REDDY (1984; as Epinephelus guaza ), but I have not seen any Indian Ocean specimens from north of 24° S. In the western Atlantic, E. marginatus is known from southern Brazil, and it was reported from Uruguay and Argentina by RINGUELET and ARAMBURU (1960).

REMARKS

In the literature on Mediterranean groupers, two similar species (the yellowbelly grouper, E. marginatus , and the dusky grouper, E. haifensis BEN-TUVIA, 1953 ) have been confused under the names Epinephelus (or Serranus ) guaza or gigas . E. marginatus differs in having 8 anal fin rays (9 in haifensis ), more elongate body (depth 2.6 - 3.1 versus 2.4-2.8 in SL), pelvic fins distinctly shorter than pectorals and not reaching the anus (pelvic fins subequal to pectorals and reaching to or beyond anus in haifensis ), 17- 19 pectoral fin rays (19-21 in haifensis ) and the head and body usually showing irregular pale blotches (no pale blotches in haifensis ).

JORDAN and EVERMANN (1896), in their influential and comprehensive work, The Fishes of North and Middle America, were the first to use the Linnaean name guaza [sic] for the species that currently bears this name. Previously, the species had been identified as Serranus gigas ( BRÜNNICH, 1768) by VALENCIENNES (1828), GÜNTHER (1859) and STEINDACHNER (1877) or Cerna gigas by DODERLEIN, 1882 or Epinephelus gigas by JORDAN and SWAIN (1885), JORDAN and EIGENMANN (1890), BOULENGER (1895) etc.; or it was described as a new species ( Serranus marginatus LOWE, 1834 and Epinephelus brachysoma COPE, 1871 ). After JORDAN and EVERMANN's (1896) publication, E. marginatus and E. haifensis were Confused under the names E. guaza or E. gigas .

Unfortunately, the species name " Epinephelus guaza " (originally Labrus Gvuza LINNAEUS, 1758 ) cannot be used for this well-known species, because the original description clearly applies to a species of the genus Mycteroperca from the coast of Venezuela. Linnaeus’s description of Labrus Gvaza (1758. 285) was taken verbatim from the travel diary of his student PEHR LÖFLING (spelt "LOEFLING“ on the title page). This diary was published in 1758, two years after the death of LÖFLING and in the same year as the tenth edition of LINNAEUS’s Systema Naturae. LÖFLING spent two years in Spam waiting for the Spanish to organize the expedition to South America in which he was to participate (WHEELER, 1980). While he was in Spain, LÖFLING collected plants and animals, recording descriptions of the various species in his travel diary. In South America, LÖFLING added descriptions of more plants and animals to his journal, but he died not long after his arrival. In the published version of this diary (LOEFLING, 1758) the page with the description of Labrus guaza is headed with the rubric "CUMANA", which is the name of a port on the Caribbean coast of Venezuela; and all of the animals described on this page are from this locality. Although most of the species descriptions by LÖFLING that were incorporated in the Systema Nature (indicated by the reference "Loefl. epist.") are of plants and animals that he observed in Spain, that of Labrus guaza is clearly not from Spam. For some reason or perhaps as an oversight, LINNAEUS gave as the type-locality of this species " in pelago ", rather than the more explicit mention of Cumana or South America or the Caribbean.

/

The original description of Labrus Gvaza , as given by LINNAEUS (1758) is typically brief:

" L. [ Labrus ] fuscus , cauda ratundata, radiis caudatus membranam superantibus. Loefl. epist. D.11 / 27 P. 16. V.6. A. 13. C. 15. Habitat in Pelago." ("Dusky Labrus , caudal fin rounded, the rays projecting past the membrane. Dorsal fin with 27 rays, of which the first 11 are spines and the last 16 soft-rays; pectoral fin rays 16; pelvic fin rays 6 [i. e., I,5]; anal fin with 13 rays [= 3 spines + 10 soft-rays]. Habitat: in the open ocean")

This description does not fit the well-known amphi-Atlantic/Mediterranean yellowbelly grouper that is commonly identified as Epinephelus guaza . In fact, it cannot apply to any species of Epinephelus , as they all have 7-9 anal fin rays (one specimen of 29 E. morio that were counted has 10 anal rays), and no Epinephelus species has the caudal fin rays projecting beyond the membrane. The description does, however, fit Mycteroperca cidi CERVIGON, 1966 and M. interstitialis (POEY, 1860) ; and these two species are common in the vicinity of Cumana (the type locality given by LÖFLING for Labrus guaza ). Since the description applies equally well to both of these species of Mycteroperca and there is no extant type-specimen, the name Labrus gvaza LIN-Plate I NAEUS must be considered a doubtful name (nomen dubium) and is thus not available as the valid name of any species.

The next oldest name that has been listed in the synonymy of " Epinephelus guaza " is Perca gigas BRÜNNICH, 1768: 65 (type locality, Marseilles). The fin-ray counts given by BRÜNNICH do not fit E. marginatus or E. haifensis , although this description was apparently based on one or (perhaps) both of these species. The anal fin count ("3/ 12 "; i. e, 3 spines and 9 soft-rays) would apply to haifensis , rather than marginatus , but the pectoral fin count of 16 is too low even for marginatus , which has fewer pectoral rays. The brief colour notes fit marginatus better than haifensis . Since there is no existing type-specimen of Perca gigas , this name must also be considered a nomen dubium.

Serranus Mentzelii VALENCIENNES, 1828 was considered a synonym of " Epinephelus guaza " by C.L. SMITH (1971) and BAUCHOT et al. (1984: 31), and they listed three supposed "syntypes" collected in Brazil by DELALANDE The two alcoholic "syntypes" (MNHN 137 & 7368) Were recently examined and found to be specimens of Mycteroperca (the species not readily apparent) with 11 anal fin rays. In the original description of S. Mentzelii ( CUVIER and VALENCIENNES, 1828: 291) , VALENCIENNES gives an anal fin count of III,8; and he mentions only a single specimen "qui est long de deux pieds huit pouces" (2 feet, 8 inches long). Consequently, neither of these "syntypes" (MNHN 137 & 7368) nor the dry specimen (MNHN A. 5777, 256 rnm. total length) also listed as a "syntype" by BAUCHOT et al. (1984) can be considered a type specimen, as they are much shorter than the only specimen (holotype) mentioned in the original description; and they do not agree with the description of this species. The holotype is apparently lost. Dr M. L. BAUCHOT sent a photograph of the large charcoal drawing from the collection of Prince Maurice, which VALENCIENNES mentioned in the original description. Although this drawing is crude, it looks like Epinephelus itajara ( LICHTENSTEIN, 1822) , and it shows the dark cross-bars on the base of the caudal fin and peduncle that are typical of large specimens of this species. The description of the colour pattern given by VALENCIENNES also fits E. itajara much better than E. marginatus .

Serranus dichropterus VALENCIENNES, 1828 was listed as a synonym of " Epinephelus guaza " by C. L. SMITH (1971) and BAUCHOT er al. (1984: 27). But VALENCIENNES proposed Serranus dichropterus as a replacement name for Holocentrus ongus BLOCH, 1790 ; consequently (according to Article 72[e] of the International Code of Zoological Nomenclature, 1985 edition) Serranus dichropterus is automatically an objective synonym of Epinephelus ongus (BLOCH) .

Perca robusta COUCH (1832) was also listed in the synonymy of Epinephelus guaza by C. L. SMITH (1971). In his original description, COUCH (1832: 21) gave the length of his holotype (which was apparently not preserved) as 3 feet and the body depth as 7 inches. Although the fin counts given by COUCH (dorsal fin with ll spines and 16 rays; anal fin with 2 spines [the small first spine was probably overlooked] and 8 rays; pectoral fin rays 19) fit E. maıginatus , the body depth of "7 inches" is much too small for a marginatus of 3 ft. total length (the depth would be at least 9 inches in a fish this size). A more likely candidate for P. robusta is Epinephelus aeneas which, is more elongate than marginatus (body depth 3.0-3.6 times in SL), and it has virtually the same fin counts. Unlike E. marginataus , E. aeneus also has 3 or 4 oblique, pale blue stripes across the operculum, and COUCH mentions "two slightly marked lines on the gill covers running obliquely downward, one on each plate." The original illustration of Perca robusta is too crude to be recognizable as any of the grouper species that are known from the eastern Atlantic; the body depth of the illustrated fish is contained 3.4 times in the total length. In view of the deficiencies of the original description and illustration of P. robusta , this species is regarded as unidentifiable.

The oldest available name that can definitely be ascribed to the yellowbelly grouper is Serranus marginatus LOWE (1834) . This original description is also brief:

" Serranus marginatus . Sem nigrescens , lateo maealatas; pinnis dorsali, anali, caadaliqae nigris, albo marginatis; pinna dorsali filamentoso. D. 11 + 17. P. 18. V. 1 + 5. A.3 + 9. C.18. This fish is very nearly related to Sem Gigas, CUV. & VAL. ; but appears to be distinguished by the greater number of the soft rays of its dorsal and anal fins, as well as by the white margin of these and the caudal. "

In 1836 LOWE redescribed his species with a replacement name, Serranus fimbriatus , and he gave slightly different fin-ray counts (D 11 + 15 - 16; A 3 + 8) for the dorsal and anal fins. These revised counts fit exactly the species here recognized as E. marginatus ; and, combined with the brief colour notes and LOWE’s illustration (Plate I, Fig. 1 , labelled Serranus marginatus ) leave no doubt of the identity of his species. A stuffed was specimen of Serranus marginatus (29cm. SL, BMNH 1858.42.81) from Madeira donated to the Zoological Society of cm. London by the Rev. R.T. LOWE and later incorporated in the fish collection of the British Museum. This specimen ( Fig. 3 ) probably was one of those exhibited at the December 24, 1833 meeting of the Zoological Society at which the description of Serranus rnarginatus was read. Although LOWE (1834) mentioned that the species "attains a length of 2 feet, and the weight of 8 pounds", he did not give a length for the fish on which his description was based; hence, it is not possible to definitely identify a type specimen.

Serranus aspersus JENYNS , based on a fish 4.25 inches from the Cape Verde Islands, was listed as a synonym of Epinephelus adscensionis by BOULENGER (1895) and C.L. SMITH (1971). But JENYNS’ (1843) counts of 15 dorsal and 17 pectoral fin rays for S. aspersus are too low for E. adscensionis (with 16- 18 dorsal and 18-20 pectoral rays), and the colour description (which is unlike the colour pattern of E. adscensionis ) matches juveniles of E. marginatus . Furthermore, E. adscensionis is not known from the Cape Verde Islands.

MATERIAL EXAMLNED

EASTERN ATLANTIC: Madeira: Neotype, MMF 3388 (200 mm.); BMNH 1858.42.81 (290 mm); BMNH 1862.4.22.30 (430 mm); BMNH 1922.1.13.30 (168 mm.); BMNH 1928.1.21.30 (212 mm.); BMNH 1935.3.5.14 (205 mm.); BMNH 1962.6.25.10 (233 mm. SL); MMF 24939 (84 mm.); MMF 3118 (118 mm.); MMF 3884 (162 mm.); MMF 3885 (151 mm.); MMF 7580 (150 mm.); RUSI 35667 (51 & 63 mm.). Azores: BMNH 1962.6.25.10 (233 mm.); RUSI 36079 (97 mm.); RUSI 36102 (3, 104- 181 mm.); USNM 94487 (262 mm.). Cape Verde Islands: MB 503 (109 mm.).

MEDITERRANEAN: Spain: IIPB 1052/1987 (158 mm.); IIPB 1053/1987 (175 mm.); IIPB 1054/1987 (232 mm.); IIPB 1055/1987 (239 mm.); IIPB 1056/1987 (254 mm.); IIPB 1057/1987 (194 mm.). France: MNHN 4215 (171 mm.); MNHN 1898-554 (106 mm.). Italy: IRSNB 2013 (310 mm.); MNHN 7229 (208 mm.). Greece: BMNH 192812130 (212 mm.); MNHN 1975-641 (152 mm). Lebanon: USNM 198881 (4, 56-88 mm.); USNM (uneat.) (6, 61-106 mm.). Israel: BMNH 1935.35.14 (205 mm); MNHN 1958- 15 (154 mm.); TAU 61 (166 mm.). Algeria: MNHN 264 (211 mm.).

ANGOLA: RUSI 2598 (195 mm.); MB 414 (89 mm.); MB 2053 (340 mm); MB 2063 (354 mm.); MB 2342 (445 mm.); MB 2545 (147 mm.).

SOUTH AFRICA: BMNH 1906.11.19.48 (316 mm.); BMNH 1922.1.13.30 (168 mm.); RUSI 2656 (339 mm); RUSI 11952 (155 mm); RUSI 12922 (451 mm. SL); RUSI 13143 (620 mm.); RUSI 13309 (370 mm.); RUSI 13410 (303 mm.); RUSI 17330 (280 mm); RUSI 17487 (3, 98-204 mm.); RUSI 74- 345 (6, 57- 134 mm.); RUSI 76- 15 (3, 85-139 mm.); RUSI 77-16 (6, 77-98 mm.); RUSI 76-20 (228 mm); RUSI 77-3 (7, 96-165 mm.); RUSI 77-6 (4, 83- 131 mm.); SAM 13803 (8, 54- 185 mm.); BPBM (uncat.) (209 mm.).

MOZAMBIQUE: RUSI 16304 (443 mm.); RUSI 17150 (378 & 390 mm.); RUSI 17597 (115 mm.); RUSI 17599 (165 mm).

WESTERN ATLANTIC: Brazil: ANSP 13372, 149 mm ., holotype of E. brachysoma COPE, 1871 ; CAS/SU 68129 (247 mm.); CAS/SU 68130 (223 mm.); MCZ 10147 (194 mm.); MNHN 7403 (168 mm.); MZUSP (uncat) (4, 124- 208 mm.); USNM 100823 (161 mm.); USNM 100886 (178 & 187 mm.).

Figures

Fig. 12 - Epinephelus marginatus, 45 cm. SL. Cape Vidal, South Africa, RUSI 12922. Drawn by E. HEEMSTRA.

Fig. 13 - Epinephelus marginatus, 29 cm. SL, Madeira, BMNH 1858.42.81; presented to the Zoological Society of London by the Rev. R. T. LOWE. Photograph courtesy of the British lt/ ......, lııcøıım ....... / Nqtııı--ıl (uawra. ....

Fig. 1 - Diagramatic illustration of a grouper's head to show method of measuring maxilla width (MW); UL =upper lip; SM=supramaxilla.

Fig. 3 - Cephalopholis taeniops, 16 cm. SL, Sénégal, from STEINDACHNER, 1882.