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Iwatsuki, Yukio, Al-Mamry, Juma M., Heemstra, Phillip C. (2016): Validity of a blue stripe snapper, Lutjanus octolineatus (Cuvier 1828) and a related species, L. bengalensis (Bloch 1790) with a new species (Pisces; Lutjanidae) from the Arabian Sea. Zootaxa 4098 (3): 511-528, DOI: http://doi.org/10.11646/zootaxa.4098.3.5

Lutjanus octolineatus (Cuvier 1828) Proposed English name: Whitebelly snapper Figures 1 A���B, 2 A, & 3 A; Table 1 Diacope octolineata Cuvier in Cuvier & Valenciennes 1828: 418 (in part, Type locality: Red Sea; Tahiti; Mauritius, Mascarenes, southwestern Indian Ocean; Ambon Island, Molucca Islands, Indonesia; Taxonomic identities of other syntype specimens are discussed under Remarks). Diacope octovittata Valenciennes in Cuvier & Valenciennes 1830: 528 (Type locality: Mauritius, southwestern Indian Ocean). Lutjanus bengalensis (non Bloch); Bleeker 1874: 27 (R��union); Kyushin et al. 1977: 68, unnumbered photograph (in part, Saya de Malha Bank); Allen & Talbot 1985: 21, pl. 1 E, F (in part, Mauritius, and Amirante Islands [the Amirantes Islands are in Seychelles]); Allen 1985: 61, pl. XII, 45 (in part, northern Indian Ocean); Kuiter 1998: 106, unnumbered underwater photograph (Maldives); Debelius 1999: 102 (in part, Mascarenes); Kuiter & Debelius 2006; 449, unnumbered photograph (Maldives); Allen & Erdmann 2012: 445, unnumbered photograph (Mauritius). Lutjanus octovittatus; Bleeker 1874: 35 (R��union). Lectotype: MNHN 0000-A 5446 (blue glass plate attached with one dried specimen at each for both sides; the designated lectotype with snout skin and the other without snout skin as paralectotype below), dried, 122 mm SL, Mauritius. Paralectotype: MNHN 0000-A 5446, dried, presumably ca. 119 mm SL (with snout region damaged on the side of blue glass plate attached with a label of MNHN 5446), other data same as the lectotype. Non-type specimens (n = 6): MNHN 0000- 7046 (holotype of Diacope octovittata), 137 mm SL, Mauritius, Mascarenes, southwestern Indian Ocean; NSMT 121733 ��� 121734, 2 specimens, 154���162 mm SL, Seychelles, Indian Ocean; HUMZ 90013, 209 mm SL, Saya de Malha Bank, Indian Ocean (10 ��47.0'S, 62 ��13.0'E); WAM 26780 -001, 2 specimens, 67���72 mm SL, Mauritius, Indian Ocean. Photographic voucher: SAIAB 81620, 136 mm SL, Nosy Be, Madagascar. Diagnosis. A species of Lutjanus with the following combination of characters: Dorsal fin usually XII, 12 or 13; anal fin III, 8; pectoral fin 16 or 17; lateral-line scales 47���50: horizontal scale rows above lateral line 8 or 9; scale rows on cheek 4���6; subocular extension of cheek scales irregularly present in 1 or 2 row(s) anteriorly and 3 or 4 rows posteriorly, respectively below eye (Fig. 2 A); total gill-rakers on first arch 8 or 9 + 18 or 19 = 26���27; no blue stripe on upper part pf opercle, the third blue stripe on body (upper supracleithrum) beginning from dorsoposterior margin of opercle, extending to below the last soft dorsal-ray base. Description. Meristic counts and proportional measurements as percentages of standard length (SL), plus two diagnostic characters in coloration and squamation of subocular extension for the lectotype, the paralectotype and non-type specimens of Lutjanus octolineatus are shown in Table 1. Body oblong, greatest depth 2.8 ��� 3.1, head length 2.5���2.8 in SL; snout length 3.1���3.8, eye diameter 2.9���3.7, interorbita 1 width 4.3���5.5, maxilla length 2.6���2.7, and preorbital depth 5.6 ���7.0 in head length; head relatively small with pointed snout; snout-forehead profile almost straight; pair of small rounded and somewhat ovate nostrils, anterior and posterior openings on each side of snout, with no flap on outer edge of anterior nostril; body scales rising obliquely above lateral line; body, opercle and preopercle with finely ctenoid scales; parietal, frontal, interorbital and rear edge of preopercle scaly; preopercular margin finely serrate (serrae increasing in size ventrally); sharp, flattened, flexible projection at middle of opercular margin, with blunt bony spine (mostly covered by scales) immediately above its base; margins of opercle, interopercle and subopercle smooth; preopercular notch and interopercular knob well developed; vomerine teeth in a crescentic band without median posterior extension (becoming a larger and randomly scattered group of 10���13 similar canine teeth in a 209 mm SL specimen, 2 or 3 irregular rows of canines on both right and left branches of specimens 10���15 cm SL); tongue smooth without granular teeth; upper jaw with very short canine on each side anteriorly, and a smaller canine between it and symphysis; remaining lateral portion of upper jaw with a series of ~ 3���10 smaller canines becoming embedded, lower jaw with ~ 5���10 small canines on each side, becoming embedded with growth, 2 or 3 canines visible, but not the anterior canine tooth when mouth closed; dorsal-fin margin slightly incised, fourth or fifth spines longest, remaining spines gradually decreasing in length; profile of soft portion of dorsal-fin relatively low and weakly rounded; pectoral fins pointed, caudal fin emarginate. TABLE 1. Counts and proportional measurements of Lutjanus octolineatus L. bengalensis and L. sapphirolineatus n. sp. Lutjanus octolineatus Lutjanus bengalensis L. sapphirolineatus n. sp. ������continued on the next page TABLE 1. (Continued) Color in life, based on J.E. Randall���s photograph (10.8 cm SL, Mauritius; http://www.fishbase.org/photos/ PicturesSummary.php?StartRow= 1 &ID= 1409 &what=species&TotRec= 6; 14 Dec. 2015) and G. Allen���s underwater photograph (12 cm TL, St. Joseph, Seychelles; http://www.fishbase.org/photos/ PicturesSummary.php?StartRow=0&ID= 1409 &what =species&TotRec= 6; 14 Dec 2015): generally bright yellow on upper half, abruptly white (as indicated by the proposed common name), the belly coloration sometimes becoming reddish, often post mortem; upper side of body with four dark-edged blue stripes, uppermost extending from interorbital to below middle of spinous dorsal fin, second from upper rear edge of eye to below front part of soft portion dorsal-fin, third from behind upper rear edge of opercle (no stripe on upper part of opercle) to area just below base of the last soft dorsal-fin ray, and the lowest (fourth) from just above middle of maxilla across lower edge of eye, extending through middle preopercle and opercle to anterior third of caudal peduncle just below lateral line; median fins yellow, paler distally in soft portion of dorsal and anal fins; pectoral and pelvic fins whitish; iris yellow. ������continued on the next page TABLE 1. (Continued) * The lectotype and the paralectotype specimens of Mesoprion pomacanthus, 30 (37 +) and 36 (43 +) mm, are designated and registered in RMNH PISC. 38327 and RMNH PISC. 38328, respectively. ** Subocular extension of cheek scales irregularly present in 1 or 2 row(s) anteriorly and 3 or 4 rows posteriorly, respectively below eye. Color in alcohol: generally yellowish tan (including fins), with four brown or bluish stripes as noted above. Distribution. Lutjanus octolineatus is confined on the basis of voucher specimens to the Western Indian Ocean: currently known from South Africa (Sodwana Bay and Cape Vidal), Mauritius, Seychelles, Saya de Malha Bank (southwest of the Chagos Archipelago) and Maldives (Kuiter 1998). Photographic vouchers are from Mozambique, Madagascar and R��union (E. Heemstra, pers. comm.). Remarks. One of two syntypes (MNHN 0000-A 5446, 122 mm SL, Fig 1 A, explained above under Lectotype and Paralectotype) is designated as the lectotype of Diacope octolineata Cuvier in Cuvier & Valenciennes 1828, the other (without snout skin) becoming the paralectotype. Allen & Talbot (1985) noted ���We have examined a syntype of Diacope octolineata, the holotype of Mesoprion etaape (both MNHN 7973, 155 mm SL)��� in Remarks in L. kasmira. The suggested syntypes of Diacope octolineata Cuvier are as follows: MNHN 0000-A 5448 (as L. kasmira [Forssk��l 1775], 1, not 2 as erroneously listed in CAS database, C. Ferrara, R. Causse, G. Zora, pers. comm.), MNHN 0000- 7971 (1, as L. kasmira), MNHN 0000- 7972 (1, as L. kasmira), all from Mauritius, plus MNHN 0000- 7973, the last, was subsequently used for the holotype of Mesoprion etaape Lesson 1831 from Tahiti. All these specimens have ���dorsal-fin rays X, 14 and four clear stripes on the body in dried and preserved specimens���, here shown as diagnostic characters of L. kasmira among the blue-striped snapper complex (Allen & Talbot 1985; p. 48, Table 3 as blue-striped complex). Unfortunately, one syntype from the Red Sea has been lost (C. Ferrara, R. Causse, G. Zora, pers. comm, MNHN database). Lastly, two other syntypes (MNHN 0000-A5547, 2 dried specimens, ~ 122���133 mm SL, Indian Ocean, collected by Commerson) were identified as L. notatus (Cuvier 1828) with six recognizable dark stripes on the body. Both have a narrow dark stripe from the lower pectoral-fin base (visible with transmitted illumination) to the lower caudal peduncle (see Table 3 of Allen & Talbot [1985]; L. bengalensis is without such a stripe) and dorsal-fin rays presumably XI, 12 for both (counted using dorsal-fin pterygiophores because of the missing fin are diagnostic characters of L. notatus among the blue-striped snapper complex (Allen & Talbot 1985), and do not apply to L. quinquelineatus with X, 13���15 (usually 14). Lutjanus octolineatus is distinguished from the closely related L. bengalensis by a dorsal-fin spine count of XII (vs. XI in L. bengalensis). It has no blue stripe on the upper part of opercle, i.e. third stripe beginning from dorsoposterior margin of opercle (vs. blue stripe present on upper part of opercle in L. bengalensis, i.e. third stripe beginning before posterior margin of opercle). Furthermore, L. octolineatus has a subocular extension of 1 or 2 scale rows on the cheek to below anterior margin of eye, but L. bengalensis has no subocular extension. Hence, the two species are distinct, and are sympatric in the Western Indian Ocean. Subtly different niches of the two species might be observed with further research. Analysis of the mitochondrial cytochrome co-oxidase subunit 1 (CO 1, 603 bp) strongly supports these as two distinct species, L. octolineatus and L. bengalensis, as well as distinguishing the other blue-striped snapper complex: L. kasmira, L. notatus, L. quinquelineatus, and L. sapphirolineatus n. sp. (See each species, pl. VI, A���C, pl. VIII, C���D, pl. VIII, E���F of Allen & Talbot [1985], and Fig. 4, respectively, the last noted below). Lutjanus sapphirolineatus n. sp., described below, can be differentiated by the following combination of characters: dorsal-fin XI, 13, and the third stripe from upper rear edge of eye, through upper part of opercle to upper part of caudal peduncle. Lastly, Kyushin et al. (1977) included a color photograph (p. 70, 198 mm SL) that was identified as L. kasmira, as well as one of the true L. kasmira (p. 70, 206 mm SL). Both specimens were collected from the Chagos Archipelago. We identified the 198 mm SL specimen as L. octolineatus (missing from HUMZ), with 12 dorsal-fin spines, a diagnostic character. This presumably L. octolineatus specimen has a subtly different stripe pattern (usually there is no blue stripe on the upper opercle in specimens collected from South Africa, Mauritius, R��union, Seychelles and Maldives), but shows a small, faint blue patch on the rear upper margin of the opercular membrane, which continues to the third blue stripe on the body (Kyushin et al. 1977; p. 70, unnumbered figure, 198 mm SL)., Published as part of Iwatsuki, Yukio, Al-Mamry, Juma M. & Heemstra, Phillip C., 2016, Validity of a blue stripe snapper, Lutjanus octolineatus (Cuvier 1828) and a related species, L. bengalensis (Bloch 1790) with a new species (Pisces; Lutjanidae) from the Arabian Sea, pp. 511-528 in Zootaxa 4098 (3) on pages 513-519, DOI: 10.11646/zootaxa.4098.3.5, http://zenodo.org/record/264482, {"references":["Cuvier, G. & Valenciennes, A. (1828) Histoire naturelle des poissons. Tome second. Livre troisieme. Des poisons de la famille des perches, ou des percoides. F. G. Levrault, Paris, xxi + 490 pp.","Cuvier, G. & Valenciennes, A. (1830) Histoire naturelle des poissons. Tome sixieme. Livre sixieme. Partie I. Des Sparoides; Partie II. Des Menides. F. G. Levrault, Paris, xxiv + 559 pp.","Bleeker, P. (1874) Poissons de Madagascar et de l'ile de La Reunion. In: Bleeker, P. & Pollen, F. P. L. (Eds.), Recherches sur la faune de Madagascar et de ses dependances, d'apres les decouvertes de Francois P. L. Pollen et D. C. van Dam. Part 4. Brill, Leiden, 104 + 2 unnumbered pp., 21 pls.","Kyushin, K., Amaoka, K., Nakaya, K. & Ida, H. (1977) Fishes of Indian Ocean. Japan Marine Fishery Resource Research Center, Tokyo, 392 pp., 179 pls. [in Japanese and English]","Allen, G. R. & Talbot, J. H. (1985) Review of the snappers of the genus Lutjanus (Pisces: Lutjanidae) from the Indo-Pacific, with the description of a new species. Indo-Pacific Fish, 11, 1 - 87, pls. 1 - 10.","Kuiter, R. H. (1998) Photo guide to Fishes of Maldives. Atoll Editions, Apollo Bay, Victoria, 256 pp.","Debelius, H. (1999) Indian Ocean Reef Guide. IKAN-Unterwasserarchiv, Frankfurt, 321 pp.","Kuiter, R. H. & Debelius, H. (2006) World Atlas of Marine Fishes. Second edition. IKAN- Unterwasserarchiv, Frankfurt, 728 pp.","Allen, G. R. & Erdmann, M. V. (2012) Reef Fishes of the East Indies. Vol. II. Tropical Reef Research, Perth, 856 pp.","Forsskal, P. S. (1775) Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit. Post mortem auctoris edidit Carsten Niebuhr. Molleri, Copenhagen, xxxiv + 164 pp."]}

Lutjanus sapphirolineatus n. sp. Proposed English name: Arabian blue-striped snapper Figures. 2 C, 4; Table 1 ? Genyoroge bengalensis; G��nther 1859: 178 (Red Sea). Lutjanus bengalensis (non Bloch): Dor 1984: 138 (?in part, Red Sea); Allen & Talbot 1985: 21 (in part, Middle East); Allen 1985: 61 (in part, Middle East); Goren & Dor 1994: 37 (?in part, Red Sea); Randall 1995: 197 (Oman); Manilo & Bogorodsky 2003: S 108 (?in part, Red Sea); Heemstra & Heemstra 2004: 201 (?in part, Middle East); Fricke et al. 2009: 60 [?in part, Western Indian Ocean]; Iwatsuki et al. 2013: 32 (Oman). Holotype: MUFS 33724, 153 mm SL, Muttra, Gulf of Oman, Oman. Paratype specimens (n = 9): MUFS 33521 ���33522, 2 specimens, 79���144 mm SL, Muttra, Oman; OMMSFC 760 (formerly MUFS 33723), 145 mm SL, same as holotype SAIAB 201838 (formerly MUFS 33523), 143 mm SL, same as holotype; USNM 305002, 4 specimens, 137���145 mm SL, Somalia (10 �� 44 ' 30 "N, 051�� 14 ' 30 "E). Diagnosis. A species of Lutjanus with the following combination of characters: Dorsal fin XI, 13; anal fin III, 8; pectoral fin 17; lateral-line scales 47���49: horizontal scale rows above lateral line 5 or 6; scale rows above lateral line rising obliquely toward dorsal profile; predorsal scales extending forward just above anterior edge of orbit; scale rows on cheek 6 or 7 without subocular extension of smaller scales; subocular extension of cheek scales irregularly present in 1 or 2 row(s) anteriorly and 3 or 4 rows posteriorly below eye, respecitvely; gill-rakers on first arch 8 or 9 + 18 or 19 = 26���28; tongue smooth, without teeth; preopercular notch developed just above angle; interopercular knob inconspicuous; belly white with yellow spot on each scale forming narrow, longitudinal, yellow lines; four dark-edged blue stripes present, the first blue stripe almost running from above posterior nostril to the tenth dorsal fin spine base; the third just from upper rear edge of eye, through upper part of opercle, tapering posteriorly and extending to upper part of caudal peduncle below last soft dorsal-ray base; fourth blue stripe from just above midpoint of lower jaw, below eye and continuing through opercle, above upper pectoral-fin base to caudal peduncle. Description. Meristic counts and proportional measurements as percentages of standard length (SL), plus two diagnostic characters in coloration and squamation of subocular extension for the holotype, nine paratype and nontype specimens of Lutjanus sapphirolineatus n. sp. are shown in Table 1. Characters presented in the diagnosis are not repeated. Body slender and elongate to moderately deep, laterally compressed, greatest depth 2.7 ��� 3.1 (3.0 in holotype) in SL; head length 2.5���2.7 (2.7 in holotype) in SL; snout length 3.1���3.7 (3.6 in holotype), eye diameter 3.3���3.7 (3.3 in holotype), interorbita 1 4.2���5.1 (4.2 in holotype), maxilla length 2.5���3.3 (2.5 in holotype), and preorbital depth 6.7���8.9 (8.5 in holotype), in head length; head small with pointed snout; snout-forehead profile straight; body, opercle and preopercle with finely ctenoid scales; parietal, frontal, interorbital and rear edge of preopercle scaly; pair of small rounded anteriorly and somewhat ovate nostrils posteriorly, respectively, and anterior and posterior openings on each side of snout, with very low flap on outer edge of anterior nostril; vomerine with minute crescentshaped tooth patch without posterior extension; preopercular margin finely serrate (serrae increasing in size ventrally); sharp, flattened, flexible projection at middle of opercular margin, with blunt bony spine (mostly covered by scales) immediately above its base; opercular, interopercular and subopercular margins otherwise smooth; upper jaw with dagger-like short canine on each side anteriorly, a smaller canine between it and symphysis; remaining lateral portion of upper jaw with a series of 8 ��� 10 smaller canines, lower jaw with 10 ��� 12 small canines on either side, almost embedded, not visible except anterior canine slightly visible when mouth closed; dorsal-fin margin weakly incised, fourth to fifth spines longest, remaining spines gradually decreasing in length; profile of soft dorsal-fin relatively low and weakly rounded; pectoral fins pointed; caudal fin emarginate. Color in life, based on photographs (MUFS 33724, 153 mm SL, holotype, photographed by Y. Iwatsuki); G.V. Hermosa, Jr. photograph (http://www.fishbase.org/photos/PicturesSummary.php?StartRow= 2 &ID= 1409 &what=s pecies&TotRec= 6 [accessed in 18 Dec. 2015] and http://www.fishbase.org/photos/UploadedBy.php?autoctr = 7077 &win=uploaded [accessed in 18 Dec. 2015], Gulf of Oman): bright yellow on upper half, white ventrally; belly with yellow spot on each scale forming narrow, longitudinal yellow lines; upper body with four dark-edged blue stripes, uppermost extending from interorbital to tenth spinous dorsal fin, second from upper edge of eye to front of soft dorsal-fin base, third from upper rear edge of eye, through upper part of opercle, tapering posteriorly and extending to upper part of caudal peduncle below last soft dorsal-ray base; the fourth stripe beginning from just above midpoint of upper jaw, continuing just beneath eye, over middle opercle, area above upper pectoral fin base, nearly to middle of caudal peduncle just below lateral line; median fins yellow; pectoral and pelvic fins whitish; iris golden yellow. Color in alcohol: generally yellowish tan (including fins), with four brown or bluish stripes as noted above. Distribution. Lutjanus sapphirolineatus n. sp. seems to be confined to the Middle East, and is currently known only from Somalia and Oman. However, we guess that this species is likely to be distributed in the southern Red Sea and the Gulf of Aden as well, because Dor (1984, Red Sea) and Randall (1995, Oman), reported L. bengalensis from there. If these reports are accurate, we presume they refer to L. sapphirolineatus n. sp. Manilo & Bogorodsky (2010) included L. bengalensis in their checklist based on specimens collected Karaman Island, Yemen, in southern Red Sea. On request of the first author of this article, specimens deposited in SAIAB collection, were examined and reidentified as L. kasmira (E. Heemstra pers. comm.). Our current distributional information is that L. sapphirolineatus n. sp. might be distributed in southern Red Sea but L. bengalensis is not likely to occur in southern Red Sea although G��nther (1859) also reported the name, Genyoroge bengalensis, from Red Sea (this needs further confirmation). Etymology. The species is named ��� sapphirolineatus ��� with reference to the diagnostic character of four blue stripes on the body of this species. Remarks. Lutjanus sapphirolineatus n. sp. from Somalia and Oman has long been confused with L. bengalensis, as noted above, but careful examination of the blue-striped snapper complex from the Indo-Pacific shows that specimens from the Middle East should be considered as new to science. Lutjanus sapphirolineatus n. sp. differs from L. octolineatus and L. bengalensis in having the following combination of characters: a stripe on upper preopercle and upper opercle, continuing as the third blue stripe, 11 dorsal-fin spines, and the presence of a suborbital extension of cheek scales (often as 1���4 scale row[s] posteriorly); horizontal scale rows above lateral line 5 or 6; vs. the third stripe beginning from rear edge of opercle, 12 dorsal-fin spines, and the presence (1 or 2 scale row[s]) of a suborbital extension of cheek scales (1 or 2 scale row[s]) and 8 or 9 horizontal scale rows in L. octolineatus; and vs. presence of a blue stripe on upper opercle, 11 dorsal-fin spines, 8 or 9 horizontal scale rows and no anterior suborbital extension of cheek scales in L. bengalensis. Comparisons. The blue-striped snapper complex includes the following species: Lutjanus bengalensis, L. coeruleolineatus, L. kasmira, L. notatus and L. quinquelineatus (Allen & Talbot 1985). Two others are added to the complex in this report: L. octolineatus and L. sapphirolineatus n. sp., bringing the total to seven species from the Indo-Pacific. Lutjanus kasmira differs from the six other known blue-striped snappers in having four blue stripes on the body and preopercular flange naked or with some embedded scales (Allen & Talbot 1985). Furthermore, this species often has narrow gray stripes on a white abdomen and dusky upper pectoral fin and snout region (Allen & Talbot 1985; this study). Lutjanus coeruleolineatus, L. quinquelineatus, and L. notatus can be differentiated from the four other species in having 7 or 8 (rarely 9), 6, and 5 blue stripes on the body, respectively. Such stripes are usually retained in preserved specimens (vs. see diagnosis above at each and Table 1). Lutjanus coeruleolineatus is easily diagnosed from the other six species in having preopercular flange naked and clear black oval spot below junction between spinous and soft dorsal-fin rays as noted in Introduction. Furthermore, L. quinquelineatus and L. notatus resemble each other but can be differentiated in having X, 13���14 (usually 14) and XI or XII, 12���13 (usually 13) in dorsal-fin rays, respectively (Allen & Talbot 1985; this study)., Published as part of Iwatsuki, Yukio, Al-Mamry, Juma M. & Heemstra, Phillip C., 2016, Validity of a blue stripe snapper, Lutjanus octolineatus (Cuvier 1828) and a related species, L. bengalensis (Bloch 1790) with a new species (Pisces; Lutjanidae) from the Arabian Sea, pp. 511-528 in Zootaxa 4098 (3) on pages 523-525, DOI: 10.11646/zootaxa.4098.3.5, http://zenodo.org/record/264482, {"references":["Gunther, A. (1859) Catalogue of the acanthopterygian fishes in the collection of the British Museum. Gasterosteidae, Berycidae, Percidae, Aphredoderidae, Pristipomatidae, Mullidae, Sparidae. Catalogue of the fishes in the British Museum, 1, i - xxxi + 1 - 524.","Dor, M. (1984) CLOFRES. Checklist of the Fishes of the Red Sea. Israel Academy of Sciences and Humanities, Jerusalem, xxii, one map + 437 pp.","Allen, G. R. & Talbot, J. H. (1985) Review of the snappers of the genus Lutjanus (Pisces: Lutjanidae) from the Indo-Pacific, with the description of a new species. Indo-Pacific Fish, 11, 1 - 87, pls. 1 - 10.","Goren, M. & Dor, M. (1994) An Updated Checklist of the Fishes of the Red Sea. CLOFRES II. The Israel Academy of Sciences and Humanities, Jerusalem, xii + 120 pp., 2 maps.","Randall, J. E. (1995) Coastal fishes of Oman. Crowford House Publishing Pty Ltd, Bathurst, xvi + 439 pp.","Manilo, L. G. & Bogorodsky, S. V. (2003) Taxonomic composition, diversity and distribution of coastal fishes of the Arabian Sea. Journal of Ichthyology, 43 (1), 75 - 149","Heemstra, P. C. & Heemstra, E. (2004) Coastal fishes of southern Africa. NISC and SAIAB, Grahamstown, xxiv + 488 pp.","Fricke, R., Mulochau, T., Durville, P., Chabanet, P., Tessier, E. & Letourneur, Y. (2009) Annotated checklist of the fish species (Pisces) of La Reunion, including a Red List of threatened and declining species. Stuttgarter Beitrage zur Naturkunde A, Neue Serie, 2, 1 - 168."]}

Lutjanus bengalensis (Bloch 1790) English name: Bengal snapper Figures 1 C, 2 B, & 3 B���C; Table 1 Holocentrus bengalensis Bloch 1790: 102, pl. 246, fig. 2 (Type locality: Bay of Bengal, eastern Indian Ocean). Mesoprion pomacanthus Bleeker 1855: 407 (in part, Type locality: Ambon Island, Molucca Islands, Indonesia).? Genyoroge bengalensis; G��nther 1859: 178 (Red Sea). Lutjanus bengalensis; Kyushin et al. 1977: 68, unnumbered photograph (in part, Andaman Sea); Allen & Talbot 1985: 21, pl. I, E���I, F. (in part, as L. octolineatus in plate, Andaman Sea and Indonesia); Allen 1985: 61, pl. XII, 45 (in part, as L. octolineatus in plate, Andaman Sea and Indonesia); Iwatsuki et al. 1999: 316, fig. 1 A (Iye Isl., Okinawa, Japan): Iwatsuki & Satapoomin 2009: 136, unnumbered underwater photograph (Andaman Sea). Lectotype: ZMB 303, 94 mm SL, Bengal, Bay of Bengal, eastern Indian Ocean. Non-type specimens (n = 17): MUFS 10452 ���104568, 7 specimens, 165���190 mm SL, Nago, Okinawa Island, Ryukyu Islands, Japan; MUFS 12267, 139 mm SL, Malaysia; MUFS 18621, 129 mm SL and MUFS 18886, 2 specimens, 129���132 mm SL, Makassar (Ujung Pangdang), South Sulawesi, Indonesia; MUFS 42884 ���42885, 2 specimens, 51���54 mm SL, Indian Ocean (purchased by aquarium shop but presumably Seychelles); MUFS 40234, 112 mm SL, Meitsu, Miyazaki, Japan; RMNH PISC. 38327 (lectotype of Mesoprion pomacanthus, designated herein), 30 mm SL (37 + mm TL), Ambon Island, Molucca Islands, Indonesia, removed from RMNH 5521; RMNH PISC. 38328 (paralectotype of M. pomacanthus), same locality as the lectotype of M. pomacanthus, removed from RMNH 5521; SU 26900 at CAS, 112 mm SL, Philippines. Photographic voucher specimens. KAUM-I. 12266 ���12267, 2 specimens, 130���142 mm SL, Malaysia; KAUM-I. 54316, 96 mm SL, Kagoshima, Japan. Diagnosis. A species of Lutjanus with the following combination of characters: Dorsal fin XI, 13 or 14 (usually 13); anal fin III, 8; pectoral-fin 16 or 17; lateral-line scales 47���50; horizontal scale rows above lateral line 8 or 9; scale rows above lateral line rising obliquely toward dorsal profile; predorsal scales extending forward nearly to level of just above anterior edge of orbit; scale rows on cheek 8���11; no subocular extension of cheek scales; gill rakers on first arch 7 or 8 + 15���17 = 22���25; tongue smooth, without teeth; preopercular notch developed just above angle; interopercular knob inconspicuous; the first blue stripe running from above anterior edge of eye ending below the 7���9 th dorsal-fin spine base; the third stripe usually running from middle of upper part of opercle or anterior edge of upper opercle to below posterior fourth of soft portion of dorsal fin, reaching the last soft dorsalray base. Description. Meristic counts and proportional measurements as percentages of standard length (SL), plus two diagnostic characters in coloration and squamation of subocular extension for the lectotype and non-type specimens of Lutjanus bengalensis are shown in Table 1. Body oblong to moderately deep, greatest depth 2.4 ��� 2.8, head length 2.5���2.8 in SL; snout length 2.6���3.4, eye diameter 3.0��� 4.2, interorbita 1 width 4.5���5.4, maxilla length 2.2���2.9 and preorbital depth 5.1���7.3, in head length; head relatively small with pointed snout; snout-forehead profile weakly rounded; pair of small nostrils rounded anteriorly and somewhat ovate posteriorly, in anterior and posterior openings on each side of snout, with very low flap on outer edge of anterior nostril; body scales rising obliquely above lateral line; body, opercle and preopercle with finely ctenoid scales; parietal, frontal, interorbital and rear edge of preopercle scaly; preopercular margin finely serrate (serrae increasing in size ventrally); sharp, flattened, flexible projection at middle of opercular margin, with blunt bony spine (mostly covered by scales) immediately above, its base; margins of opercle, interopercle and subopercle smooth; preopercular notch and interopercular knob well developed; vomerine tooth patch crescent-shaped without a posterior extension (similar small-sized lateral canine teeth present in 1 row on both right and left branches); tongue smooth without granular teeth; upper jaw with dagger-like very short canine tooth, on each side anteriorly, with smaller canines between it and symphysis; remaining lateral portion of upper jaw with a series of ~ 10 visible smaller embedded canines, lower jaw with ~ 10 small embedded canine teeth on each side of jaws, most not visible when mouth closed; dorsal-fin outline very weakly incised, fourth or fifth spines longest, remaining spines gradually decreasing in length; profile of soft dorsal-fin relatively low and weakly rounded; pectoral fins pointed, caudal fin emarginate. Color in life, based on SAIAB���s Indian Ocean specimens (2 specimens, SAIAB 82483 and uncatalogued 1815 #6, 94 and 105 mm SL, respectively, Sodwana Bay, South Africa) and Pacific specimens (MUFS 10452 ���10458, 7 specimens, 165���190 mm SL and uncatalogued photographs from Japan): head and body generally bright yellow on upper half and white ventrally, sometimes with reddish tinge, often after death; upper side with four dark-edged blue stripes, uppermost stripe extending from interorbital to middle of spinous dorsal fin, the second from upper rear edge of eye to anterior part of soft dorsal-fin base, third stripe from posterodorsal margin of preopercle or middle of upper part of opercle to area posteriorly on body a short distance anterior to base of last soft dorsal-fin rays, and lowest stripe from just above of middle of maxilla across lower edge of eye, extending through middle preopercle and opercle to anterior third of caudal peduncle just below lateral line; median fins yellow; pectoral and pelvic fins whitish; iris yellow. Color in alcohol: generally yellowish tan (including fins), with 4 brown or bluish stripes as noted above. Distribution. Lutjanus bengalensis is widely distributed in the Indo-West Pacific (Allen & Talbot 1985; this study) and is currently known from South Africa, Seychelles, Indonesia, Malaysia, Philippines and Japan, on the basis of specimens examined. The species is also confirmed from the following areas by photographic vouchers and references (Kyushin et al. 1977; Allen & Talbot, 1985; Shen 1993; Chen 1996; Randall & Lim 2000; E. Heemstra pers. comm.; U. Satapoomin pers. comm.): Mozambique, Madagascar, Kenya, Saya de Malha Bank (southwestern area off the Chagos Archipelago), Andaman Sea (Similan Island), Taiwan and China. Furthermore, photographic vouchers (E. Heemstra, pers. comm.) show the species from the following localities: southern and northern Mozambique, Madagascar and South Africa (Sodwana Bay and Cape Vidal), plus a photograph that might be the species from Zanzibar, Tanzania. However, records of the distribution of L. bengalensis from the Middle East, including the Red Sea, may have been erroneous (Allen & Talbot 1985), because L. bengalensis was confused with L. sapphirolineatus n. sp., described below in this study. In occurrence notes on L. bengalensis from the western Pacific, Allen & Talbot (1985) and Allen (1985) reported one record of this species from Ambon, Indonesia. An underwater photograph clearly showing a school of snappers (Allen & Erdmann 2012), as L. kasmira, was taken at Mios Kon, Raja Ampat Islands, West Papua, Indonesia (by M. Allen), and could be the easternmost record of L. bengalensis (A. Ryanskiy, pers. comm.); there are no gray (or narrow reddish) stripes on the abdomen, and no dusky upper pectoral-fin rays. In the northwestern Pacific, the species was firstly reported from Okinawa Island, Japan (Iwatsuki et al. 1999). Currently this species is very common from southern Japan (Ryukyu Islands, and ranging as far north as the Kii Peninsula (Kimura et al. 2008), Tokai District, and Kanto District near Tokyo, observed by the first author), as well as Taiwan and southern China. However, during the period 1983-1995, the first author visited southwestern Asia (Philippines, Indonesia, Singapore and Taiwan) several times, but identified only one specimen (ca. 15 cm TL, not kept) of this species at the Chinese fish market in Singapore; it seemed a rare species from the Pacific at that time. The distributional range of L. bengalensis as well as L. xanthopinnis is likely to extend northwards, southwestern Japan from the northwestern Pacific presumably due to global warming (Iwatsuki et al. 2015). Remarks. A lectotype, ZMB 303, was designated by Paepke (1999) and we confirm this specimen as the lectotype (Fig. 1 C, 94 mm SL, Bengal, Bay of Bengal, India) of Holocentrus bengalensis Bloch 1790. Unfortunately, we did not have the opportunity to examine the paralectotypes, ZMB 300 (1 specimen) and ZMB 33057 (1, right side skin). Lutjanus bengalensis has long been confused with L. octolineatus, but careful examination of the lectotype of this species shows a blue stripe on the upper part of opercle, 11 dorsal-fin spines and no suborbital extension of cheek scales (vs. no stripe on upper part of opercle, 12 dorsal-fin spines and the presence of a suborbital extension of cheek scales in L. octolineatus; Figs. 1���3), discussed above under Remarks for L. octolineatus. Bleeker (1855) described Mesoprion pomacanthus on the basis of five specimens (38���48 mm in total length, Bleeker���s measurement, from snout tip to tip of caudal fin). However, Bleeker (1874; Bleeker 1876���1877) subsequently recognized M. pomacanthus as a junior synonym of Lutjanus bengalensis (Bloch 1790). Bleeker���s (1873) description of L. bengalensis was based on 20 specimens, 43���265 mm TL. The smallest of the five syntypes of M. pomacanthus (38 mm TL) was apparently already lost in 1873. According to M. van Oijen (pers. comm.), there are presently 28 of the 29 specimens of L. bengalensis listed/recorded in Bleeker (1873) in the collection of Naturalis Biodiversity Center (RMNH). The original Bleeker jar bought at the auction of the Bleeker collection (in December 1879) now contains 24 specimens. At the time of the auction, it contained 25 specimens (Hubrecht 1879). Later, three specimens from the Series B���E must have been added as 28 specimens are recorded in the RMNH register. Two specimens identified by G.R. Allen (WAM) as L. kasmira (Forssk��l 1775) were removed from RMNH 5521 in 1978 and registered as RMNH 27825 in this report. One of these two specimens is Bleeker���s largest specimen of L. bengalensis from which Bleeker removed the pharyngeal jaws for the description (Bleeker 1873). Moreover, RMNH 5521 contained four syntypes of M. pomacanthus; two specimens 35 (42 +) and 37 (44 +) mm, identified by Allen & Talbot (1985) as L. kasmira (now registered as RMNH PISC. 38326), and two specimens 30 (37 +) and 36 (43 +) mm, identified by Allen & Talbot (1985) as L. bengalensis (now registered as RMNH PISC. 38327 and RMNH PISC. 38328, respectively). Allen & Talbot (1985) identified these specimens solely on the basis of the number of dorsal-fin spines. In order to avoid further confusion, we here designate the specimen of 30 mm SL (37 + mm TL), RMNH PISC. 38327 as the lectotype of Mesoprion pomacanthus; the three other syntypes becoming the paralectotypes. In addition, Fricke (1999) placed Labrus octovittatus Lacep��de 1801 (p. 430, 477, pl., 22, fig. 2; Type locality: Grand Ocean) in synonymy of L. bengalensis although Allen & Talbot (1985) did not note this nominal species. The figure clearly shows presumably?X, probably 14 in dorsal-fin rays (as counted as IX, but the first short dorsalfin spine may have been missed or overlooked and not drawn) and presence of the third stripe, beginning from the middle of upper part of opercle. Such characters strongly suggest L. kasmira (Forssk��l 1775), (see diagnostic characters of Allen & Talbot 1985). The two characters could not be applied to L. octolineatus (with XII, 12 or 13 dorsal-fin rays) and L. sapphirolineatus n. sp. (with the third stripe beginning just from upper rear edge of eye, through upper part of opercle, to upper part of caudal peduncle; see each diagnosis of the two species)., Published as part of Iwatsuki, Yukio, Al-Mamry, Juma M. & Heemstra, Phillip C., 2016, Validity of a blue stripe snapper, Lutjanus octolineatus (Cuvier 1828) and a related species, L. bengalensis (Bloch 1790) with a new species (Pisces; Lutjanidae) from the Arabian Sea, pp. 511-528 in Zootaxa 4098 (3) on pages 521-523, DOI: 10.11646/zootaxa.4098.3.5, http://zenodo.org/record/264482, {"references":["Bloch, M. E. (1790) Naturgeschichte der auslandischen Fische. Vol. 4. Icthyologie, ou Histoire naturelle des Poissons, Berlin, xii + 128 pp., pls. 217 - 252.","Bleeker, P. (1855) Zesde bijdrage tot de kennis der ichthyologische fauna van Amboina. Natuurkundig Tijdschrift voor Nederlandsch Indie, 8, 391 - 434.","Gunther, A. (1859) Catalogue of the acanthopterygian fishes in the collection of the British Museum. Gasterosteidae, Berycidae, Percidae, Aphredoderidae, Pristipomatidae, Mullidae, Sparidae. Catalogue of the fishes in the British Museum, 1, i - xxxi + 1 - 524.","Kyushin, K., Amaoka, K., Nakaya, K. & Ida, H. (1977) Fishes of Indian Ocean. Japan Marine Fishery Resource Research Center, Tokyo, 392 pp., 179 pls. [in Japanese and English]","Allen, G. R. & Talbot, J. H. (1985) Review of the snappers of the genus Lutjanus (Pisces: Lutjanidae) from the Indo-Pacific, with the description of a new species. Indo-Pacific Fish, 11, 1 - 87, pls. 1 - 10.","Iwatsuki, Y., Yoshino, T. & Shimada, K. (1999) Comparison of Lutjanus bengalensis from the western Pacific with a related species, L. kasmira, and variations in both species (Perciformes: Lutjanidae). Ichthyological Research, 46 (3), 314 - 317. http: // dx. doi. org / 10.1007 / BF 02678519","Iwatsuki, Y. & Satapoomin, U. (2009) Lutjanidae, Snappers. In: Kimura, S., Satapoomin, U. & Matsuura, K. (Eds.), Fishes of Andaman Sea: West coast of southern Thailand. National Museum of Nature and Science, Tokyo, pp. 135 - 143.","Shen, S. C. (Ed.) (1993) Fishes of Taiwan. Department of Zoology, National Taiwan University, Taipei, 960 pp.","Chen, C. - H. (1996) Checklist of the fishes of Penghu. FRI Special Publication, 4, 1 - 175.","Randall, J. E. & Lim, K. K. P. (Eds.) (2000) A checklist of the fishes of the South China Sea. Raffles Bulletin of Zoology, 8 (Supplement), 569 - 667.","Allen, G. R. & Erdmann, M. V. (2012) Reef Fishes of the East Indies. Vol. II. Tropical Reef Research, Perth, 856 pp.","Kimura, S., Sakakibara, H., Tsumoto, K. & Iwatsuki, Y. (2008) A lutjanid fish, Lutjanus bengalensis (Actinopterygii: Perciformes), from Owase: the first record from Mie Prefecture and northernmost records for the species. Bulletin of Faculty of Mie University, 35, 65 - 69.","Iwatsuki, Y., Tanaka, F. & Allen, G. R. (2015) Lutjanus xanthopinnis, a new species of snapper (Pisces: Lutjanidae) from the Indo-west Pacific, with a redescription of Lutjanus madras (Valenciennes 1831). Journal of the Ocean Science Foundation, 17, 22 - 42.","Paepke, H. - J. (1999) Bloch's fish collection in the Museum fur Naturkunde der Humboldt Universitat zu Berlin: an illustrated catalog and historical account. Ruggell (Liechtenstein). Theses Zoologicae, 32, 1 - 216, pls. 1 - 32 (unpaginated).","Bleeker, P. (1874) Poissons de Madagascar et de l'ile de La Reunion. In: Bleeker, P. & Pollen, F. P. L. (Eds.), Recherches sur la faune de Madagascar et de ses dependances, d'apres les decouvertes de Francois P. L. Pollen et D. C. van Dam. Part 4. Brill, Leiden, 104 + 2 unnumbered pp., 21 pls.","Hubrecht, A. A. W. (1879) Catalogue des collections formees et laissees par M. - P. Bleeker. De Breuk & Smits, Leiden, v + 71 pp.","Forsskal, P. S. (1775) Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit. Post mortem auctoris edidit Carsten Niebuhr. Molleri, Copenhagen, xxxiv + 164 pp.","Fricke, R. (1999) Fishes of the Mascarene Islands (Reunion, Mauritius, Rodriguez). An annotated checklist with descriptions of new species. Koeltz Scientific Books, Koenigstein, viii + 759 pp.","Lacepede, B. G. E. (1801) Histoire naturelle des poissons (Lacepede). Histoire naturelle des poisons, 3, 1 - 558."]}

Polysteganus coeruleopunctatus (Klunzinger 1870) English name: Blueskin Seabream Figures 1 A, 2 A; Table 1 Dentex (Polysteganus) coeruleopunctatus Klunzinger 1870: 763 (Al-Qusayr [Kosseir], Egypt, Red Sea); Fricke 1992: 18 (same as original description); Fricke 2005: 60 (same as original description). Polysteganus coeruleopunctatus; Sato 1976: 98, unnumbered color sub-adult and large adult photographs (East Africa); Dor 1984: 160 (Red Sea); Baranes & Golani 1993: 308 (Red Sea); Goren & Dor 1994: 44 (Red Sea); Manilo & Bogorodsky 2003:S 109 (Red Sea). Lectotype. ZMB 7989, 258 mm SL, [Kosseir] Al-Qusayr, Egypt, Red Sea (Fig. 2 A). Paralectotype. SMNS 1873, 223 mm SL (X), same data as for the Lectotype. Other material examined. HUJF 13901, 2 specimens, 126���127 mm SL (X), Eilat, Red Sea; HUJF 20120, 226 mm SL, Eilat, Israel; SAIAB 13464, 223 mm SL, off Ras Ngomeni, Ras Ngomeni, Kenya; SAIAB 42677, 3 specimens, 161���184 mm SL (X), Eilat, Red Sea; SAIAB 42816, 332 mm SL, off Kilifi, Mombasa, Kenya; SAIAB 48180, 148 mm SL, off Mozambique (23.47 ��S, 35.72 ��E). Diagnosis. Distinguished from all congeners of Polysteganus by the following combination of characters: scale rows above lateral line 6 ��; scales rows between 5 th dorsal-fin spine base and lateral-line scale 5 �� or 6 ��; pored lateral-line scales 51 or 52; suborbital depth subequal to or greater than orbit diameter; dorsal-fin rays reddish pink as well as similar reddish pink on upper margin dorsal-fin ray (no clear yellow or faint yellowish); 7 or 8 faint irregular blue spotted lines on body, not visible from all angles; reddish membranes on caudal-fin lobes and caudal fin with clear white lowest margin of lower caudal fin lobe; no blue iridescence on each scale of dorsal and lateral sides of whitish pink body; canines large, both upper and lower canines on both jaws usually visible even when mouth closed; deep body, 44���46 % (mean 44 %) of SL; anal- and pelvic- fin rays whitish or reddish hyaline; no yellow (white) rear margin of branchiostegal rays and membrane; weak bony bulge before and above eye, forming a low median ridge in larger specimens of over 30 cm SL; caudal fin moderately forked. Description. Counts and proportional measurements as percent of standard length (SL) of the type and other specimens of Polysteganus coeruleopunctatus are given in Table 1. Body depth relatively deeper; head robust, rostro-occipital profile gently curved; interorbital fully scaly, scales extending to just above anterior and posterior nostrils, anterior nostrils small rounded and posterior nostrils oval; orbit diameter greater than interorbital width; maxilla naked, its posterior end not reaching vertical at front edge of eye, rear margin of maxilla not concealed by infraorbitals; jaw robust, especially upper jaw; canines generally stout, 4 in upper and 6 in lower jaw (two outermost canines in upper jaw greater in size than 2 medial canines, and 2 outermost canines of lower jaw greater than 4 medial canines, but one or two canines often missing or broken in both jaws); an outer lateral row of conical teeth and a series of villiform teeth on inner side of outer lateral teeth row; mandibular profile straight; rear margin of preopercle rounded and serrated, preopercle flange scaly, a narrow naked area (preopercle with keel- like ridge), separating preopercle flange scaly area and cheek scales; first 3 dorsal-fin spines progressively longer, fourth spine longest and slightly greater than snout length; no sheath scales on dorsal and pelvic-fin rays; predorsal length slightly greater than body depth, and clearly less than dorsal-fin base length; lateral line gently curved, generally parallel to curved dorsal profile between dorsal outline and curved lateral line, from nape to peduncle, but somewhat deeper anteriorly than posteriorly, giving an impression of a deeper body than that of the similar species, P. lineopunctatus (Figs. 1���2); pectoral-fin base scaly and rounded posteriorly, two or three lowest rays slightly extended posteriorly, compared with rays above (Figs. 1���2, being similar to that of Fig. 2 B of P. lineopunctatus), the fin tip reaching beyond vertical at anus, and almost reaching vertical at first anal fin spine base, its length greater than head length; first pelvic-fin ray length clearly greater than fourth dorsal-fin spine length, fin tip not reaching beyond vertical at pectoral-fin tip and rear margin of anus; soft dorsal and anal fins with no scales; anal fin with low scaly sheath, anal-fin spines slender, second slightly greater than third, subequal to orbit diameter; caudal fin lobes broad and wide, caudal fin moderately forked (Figs. 1 A, 2 A); scales ctenoid; gill-rakers lanceolate; vertebrae 10 + 14. Color when fresh, based on color photographs of specimens (HUJF 20120, 227 mm SL (Fig. 1 A); J.E. Randall���s photograph (Gulf of Aqaba, Red Sea, BPBM 39500, 167 mm SL): head and body uniform bright red with silvery-golden reflection on each scale, upper head and dorsolateral part of body deeper yellowish red than lower head and abdominal part of body; iris golden and pupil black; all fins uniform reddish hyaline, especially in membrane of dorsal-fin and very pale reddish and whitish hyaline in other fins except caudal fin reddish. Color of preserved specimens: upper sides of head and trunk uniform yellowish-tan. Polysteganus coeruleopunctatus Polysteganus lineopunctatus ZMB 7989, Lectotype; SMNS Other specimens: HUJF BMNH 1903.2. 6.5 SAM 9908, 1 of 2 Other specimens: SAIAB 1873, Paralectotype 13901, 2 specimens, Holotype syntypes of Dentex 17470, 318 mm SL, SAIAB 125���126 mm SL, HUJF natalensis * 29416, 189 mm SL, SAIAB 20120, 226 mm SL, 48180, 148 mm SL, SAIAB SAIAB 13464, 223 mm SL, 82139, 3 specimens, 92-196 SAIAB 42677, 150 mm SL, mm SL and SAIAB 42816, 332 mm SL n = 2 n = 6 n = 1 n = 1 n = 7 Suborbital depth 8.2; 7.7 6.5���9.2 (7.8) 10.7 6.1 6.4���9.4 (7.9) First dorsal-fin spine length 6.7; 6.4 6.4���7.6 (7.1) 7.4 8.7 5.7���7.8 (7.0) Second dorsal-fin spine length 10.5; 9.9 9.7���12.7 (11.1) 10.0 12.2 8.8���12.1 (10.7) Third dorsal-fin spine length 14.1; 13.2 12.4���15.1 (13.9) 13.0 16.6 12.2���15.8 (13.8) Fourth dorsal-fin spine length 14.2; 14.6 13.6���15.6 (14.6) 14.9 18.1 13.1���16.3 (14.4) First dorsal-fin ray length 10.5; 10.4 9.5���11.7 (10.6) 7.9 9.6 10.2���13.5 (11.6) First anal-fin spine length 7.7; 6.8 6.4���8.7 (7.3) 6.7 6.4 6.0��� 12.4 (8.3) Second anal-fin spine length 12.5; 12.3 11.3���15.6 (13.5) 12.3 11.7 10.8���16.3 (13.4) Third anal-fin spine length 10.7; 10.8 10.3 ���13.0 (11.7) 9.1 11.1 10.3���14.1 (11.8) First anal-fin ray length 9.3; 10.7 8.9 ���13.0 (11.1) 9.8 12.8 9.5���14.1 (11.5) Pelvic-fin spine length 16.6; 16.8 15.7���17.5 (16.4) 16.1 17.8 14.5���16.7 (15.5) First pelvic-fin ray length 19.8; 22.2 21.1���23.4 (22.4) 20.9 23.3 20.8���26.1 (22.7) Caudal peduncle depth 12.4; 12.6 11.3���12.1 (11.7) 9.8 13.4 10.6���12.2 (11.6) Caudal peduncle length 18.5; 24.0 19.9���23.1 (20.9) 19.5 20.6 20.0��� 22.3 (21.0) Predorsal length 43.1; 42.1 39.1���44.3 (42.1) 44.4 41.7 39.2���45.7 (42.7) Preanal length 64.3; 63.6 60.4���64.6.(62.1) 67.4 65.0 61.6���64.8 (63.2) Prepelvic length 40.2; 38.6 39.6���45.3 (41.0) 41.4 42.2 38.9���45.9 (40.7) Dorsal-fin base 50.9; 51.1 48.1���55.2 (52.3) 47.9 53.4 50.3���55.2 (52.2) Anal-fin base 17.4; 16.1 15.7���20.3 (17.8) 16.5 18.4 17.9���20.3 (18.8) One of two syntypes of Dentex natalensis was lost according to M. Bougaardt (SAM, pers. comm.). Distribution. Polysteganus coeruleopunctatus has been reported from the Western Indian Ocean including the Red Sea (type locality). The species was photographed in the East African Fish Faunal Research (probably Kenya) by Sato (1977: 98, unnumbered color sub-adult and large adult photographs) and might thus occasionally occur there. However, specimens formerly identified as P. coeruleopunctatus from Mozambique and South Africa are reidentified as P. lineopunctatus (Boulenger 1903) in this study. Remarks. Polysteganus coeruleopunctatus (Klunzinger 1870) has long been recognized as a valid species from the Western Indian Ocean and its distribution was considered as from the Red Sea and East Africa, to KwaZulu-Natal south coast and Eastern Cape Province, South Africa (Smith & Smith 1986; van der Elst 1981; Heemstra & Heemstra 2004). However, another species in synonymy, P. lineopunctatus (Boulenger 1903), is recognized by us as valid. Specimens formerly identified as P. coeruleopunctatus from South Africa and Mozambique were confirmed as P. lineopunctatus in this study (see next section of P. lineopunctatus). There are two syntype specimens of Dentex (Polysteganus) coeruleopunctatus Klunzinger 1870: Al-Qusair, Egypt, Red Sea. ZMB 7989 (258 mm SL) is designated as the lectotype because of its good condition and the other specimen (SMNS 1873, 223 mm SL) is designated as paralectotype in order to avoid further confusion with Polysteganus taxonomy., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2015, Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean, pp. 133-150 in Zootaxa 4059 (1) on pages 135-140, DOI: 10.11646/zootaxa.4059.1.7, http://zenodo.org/record/246088, {"references":["Klunzinger, C. B. (1870) Synopsis der Fische des Rothen Meeres. I. Theil. Percoiden-Mugiloiden. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 20, 669 - 834.","Fricke, R. (1992) Types in the fish collection of the Staatliches Museum fur Naturkunde in Stuttgart. Part 2. The Klunzinger collection. Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie), 473, 1 - 25.","Fricke, R. (2005) Types in the fish collection of the Staatliches Museum fur Naturkunde in Stuttgart, described in 1845 - 2004. Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie), 684, 1 - 95.","Sato, T. (1976) East African Deep Sea Fishes. E. Afr- 72 Dentex coeruleopunctatus. page 98 In: Far Seas Fisheries Research Laboratory (Ed.), Colored Illustrations of Bottomfishes Collected by Japanese Trawlers. Vol. II. Japan Deep Sea Trawlers Association, Tokyo, pp. 79 - 112.","Dor, M. (1984). CLOFRES. Checklist of the Fishes of the Red Sea. Israel Academy of Sciences and Humanities, Jerusalem, xxii, one map + 437 pp.","Baranes, A. & Golani, D. (1993) An annotated list of deep-sea fishes collected in the northern Red Sea, Gulf of Aqaba. Israel Journal of Zoology, 39, 299 - 336.","Goren, M. & Dor, M. (1994). An Updated Checklist of the Fishes of the Red Sea. CLOFRES II. The Israel Academy of Sciences and Humanities, Jerusalem, xii + 120 pp., 2 maps.","Manilo, L. G. & Bogorodsky, S. V. (2003). Taxonomic composition, diversity and distribution of coastal fishes of the Arabian Sea. Journal of Ichthyology, 43, S 75 - S 149.","Boulenger, G. A. (1903) Descriptions of six new perciform fishes from the coast of Natal. Annals of the South African Museum, 3 (3), 63 - 67, pls. 2 - 7.","Smith, J. L. B. & Smith, M. M. (1986) Family No. 183: Sparidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smiths' Sea Fishes. Macmillan, Johannesburg, pp. 580 - 594.","van der Elst, R. (1981) A Guide to the Common Sea Fishes of Southern Africa. C. Struik Publishers (PTY) Ltd, Cape Town, 398 pp.","Heemstra, P. C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. NISC and SAIAB, Grahamstown, xxiv + 488 pp."]}

Polysteganus flavodorsalis new species Proposed English name: Yellowfin Seabream Figure 3 A���C; Table 2 Holotype. SAIAB 84079, 143 mm SL (X), sex not determined, Nazareth Bank, Mascarene Plateau (11.15 �� 41.11 'S, 61 ��04.54'E), Trawl No. Mascarene- 7, depth 302���305 m, coll. by D. Tweddle & O. Alvheim, 18 October 2008. Paratypes. SAIAB 84022, 3 specimens, 247���265 mm SL (X), sex determined only for 265 mm SL specimen examined, off Cargados Carajos, Mascarene Plateau, Trawl No. Mascarene- 7 (16 �� 27.62 'S, 60 �� 16.84 'E), coll. by D. Tweddle & O. Alvheim, depth 214 ��� 219 m, 14 October 2008; SAIAB 84072, 223 mm SL (X), Nazareth Bank, Mascarene Plateau, Trawl No. Mascarene- 10 (15 �� 55.43 'S, 60 �� 13.38 'E), depth 232 ��� 236 mm (X), coll. and photographed by D. Tweddle & O. Alvheim; SAIAB 84097, 218 mm SL (X), Nazareth Bank, Mascarene Plateau, Trawl No. Mascarene- 12 (15 �� 23.32 'S, 61 �� 13.41 'E), coll. by D. Tweddle & O. Alvheim, depth 258 ��� 288 m, 19 October 2008. Diagnosis. Distinguished from all congeners of Polysteganus by the following combination of characters: scale rows above lateral line 6 ��; scales rows between 5 th dorsal-fin spine base and lateral line 5 ��; pored lateralline scales 52 or 53; suborbital depth clearly less than orbit diameter (OD), 72���75 % (mean 74 %) of OD; dorsal-fin rays reddish pink with bright yellow margin, anal-fin rays pinkish; no obvious lines of blue spots on body, but usually a blue iridescence in each body scale around dorsal and lateral sides of body from some angles; yellow and red membranes only on upper and lower caudal-fin lobes, respectively (the two colors are strongly contrasted; Fig. 3 A���C); somewhat deeper body, 39���43 (mean 41 %) of SL; no bony bulge before and above eye; caudal fin moderately forked. Polysteganus cerasinus n. sp. expressed as percentages of standard length (mean in parenthesis). Description. Counts and proportional measurements as percent of standard length of the holotype and five paratype specimens of Polysteganus flavodorsalis n. sp. are given in Table 2. Body depth relatively deeper; head robust, rostro-occipital profile gently convex; interorbital fully scaly, scales extending to nearly above anterior nostrils and just above posterior nostrils, anterior nostrils small and rounded, posterior nostrils slit-like; orbit diameter clearly greater than interorbital width; maxilla naked, its rear end nearly reaching vertical at front edge of eye; rear margin of maxilla not concealed by infraorbitals; jaw robust, especially upper jaw; canines slender and short, 4 in upper and 6 in lower jaw (2 outermost canines in upper jaw somewhat greater than 2 medial canines, and 2 outermost canines of lower jaw considerably larger than 4 medial canines; an outer lateral row of conical teeth and a series of villiform teeth on inner side of outer lateral teeth row; mandibular profile nearly straight; rear margin of preopercle rounded and feebly serrate; a narrow naked area (preopercle with keel-like ridge), between cheek scales and preopercle flange scaly area; first to third dorsal-fin spines progressively longer, fourth dorsal-fin spine longest, but clearly greater than snout length; no sheath scales on dorsal and pelvicfin rays; predorsal-fin length subequal or less than body depth, and clearly less than dorsal-fin base length; lateral line gently curved, subequally parallel to curved dorsal profile from nape to peduncle; pectoral-fin base scaly and rounded posteriorly, 2 or 3 lowest rays slightly extended posteriorly, compared with rays above, creating a concave rear margin for pectoral fins, the fin tip reaching beyond vertical at anal fin origin, its length subequal to or slightly greater than head length; pelvic fin length subequal to fourth dorsal-fin spine length, fin tip not reaching beyond vertical at pectoral-fin tip, but often beyond rear margin of anus; soft dorsal and anal fins with no scales; anal fin with moderate scaly sheath, anal-fin spines slender, second slightly greater than third and clearly less than orbit diameter; caudal fin lobes broad and wide; scales ctenoid; gill-rakers lanceolate; vertebrae 10 + 14. Color when fresh, based on color photographs of the holotype and 5 paratypes, photographed by O. Alvheim; head and body uniform pinkish red or orange with blue iridescence at center of each scale, upper head and dorsolateral part of body with more blue iridescence than lower head and abdominal parts of body, and color of snout and upper part of head somewhat yellowish with growth (Fig. 3 A���C); iris golden and pupil black; all fins generally uniform pinkish. Color of preserved specimens: upper sides of head and trunk uniform yellowish-tan. Distribution. Polysteganus flavodorsalis n. sp. is currently known only from Cargados Carajos and Nazareth Bank (11.15 �� 1 'S, 61 �� 13.41 'E) at a depth of 214 ��� 305 m. Etymology. This fish is named for the bright yellow upper margin of the dorsal-fin membranes. Remarks. Specimens were initially identified as P. coeruleopunctatus because of the uniform pink or red body and similar body proportions of the two species. However, we noticed that this species differs in the dark blue iridescence on the body and the yellow and pink color of the upper and lower caudal fin lobes (Figs. 3 A���C)., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2015, Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean, pp. 133-150 in Zootaxa 4059 (1) on pages 141-144, DOI: 10.11646/zootaxa.4059.1.7, http://zenodo.org/record/246088

Iwatsuki, Yukio, Heemstra, Phillip C. (2015): Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean. Zootaxa 4059 (1): 133-150, DOI: http://dx.doi.org/10.11646/zootaxa.4059.1.7

Key to species of the Polysteganus from Indian Ocean 1 a Large oval dark blotch on lateral line below 5 th and 6 th dorsal-fin spines in adults, but no dark blotch in 9 cm juveniles; body with 4 or 5 wide blue stripes (more than 1 / 3 width of black pupil diameter).............................. P. undulosus 1 b No dark blotch on lateral body; 7 or 8 narrow blue stripes (less than 1 / 4 width of black pupil diameter) or lines of blue spots on body if present....................................................................................... 2 a 2 a Greatest body depth at preopercle vertical edge; lower-limb gill-rakers 14���16......................... P. praeorbitalis 2 b Greatest body depth at pectoral-fin base; lower-limb gill-rakers 10���13............................................ 3 3 a Body with 7 or 8 longitudinal blue stripes.......................................................... P. baissaci 3 b No longitudinal blue stripes or lines of blue spots on body..................................................... 4 4 a Caudal fin weakly forked, nearly truncate if widely spread; distinct bony bulge before and above eye in specimens about 60 cm SL (bulge probably not apparent in juveniles); body reddish orange, without lines of blue spots........... P. mascarenensis 4 b Caudal fin forked; no strong bony bulge both before and above eye; body reddish yellow often with clear irregular lines of blue spots................................................................................................ 5 5 a Scale rows between 5 th dorsal fin spine and lateral line 4 ��; no irregular lines of blue spots; rear margin of membrane of branchiostegal rays yellow, plus yellow color on a few body scales just above upper pectoral-fin base..................................................................................................... Polysteganus cerasinus n. sp. 5 b Scale rows between the 5 th dorsal fin spine and lateral line 5 ����� 7 ��; no or 7���8 irregular lines of blue spots; rear margin of membrane of branchiostegal rays white (not yellow).......................................................... 6 6 a Blue iridescence in each body scale on dorsal and lateral sides of body visible at some angles, and no irregular lines of blue spots; upper caudal-fin lobe yellow, lower red..................................... Polysteganus flavodorsalis n. sp. 6 b No blue iridescence in body scales on dorsal and lateral side; 7 or 8 irregular lines of blue spots on body, visible at some angles; both caudal-fin lobes with pink or red membranes...................................................... 7 7 a Scale rows above lateral line 6 ��; pored lateral-line scales 51 or 52; 7 or 8 faint irregular lines of blue spots on body, visible at some angles; dorsal-fin rays reddish pink but with no bright yellow upper margin; canines large, both upper and lower canines on both jaws usually visible even when mouth closed; body deeper, 44���46 % (mean 44 %) of SL (Figs. 1 A, 2 A)............................................................................................... P. coeruleopunctatus 7 b Scale rows above lateral line 7 ��; pored lateral-line scales 53���55; 7 or 8 distinct irregular lines of blue spots on body (often present even in preserved specimens); dorsal-fin rays reddish pink or orangish red with bright yellow upper margin; canines somewhat smaller, generally only upper canines visible when mouth closed; body somewhat slender, 40���43 % (mean 42 %) of SL (Figs. 1 B, 2 B)........................................................................ P. lineopunctatus, Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2015, Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean, pp. 133-150 in Zootaxa 4059 (1) on page 147, DOI: 10.11646/zootaxa.4059.1.7, http://zenodo.org/record/246088

Polysteganus lineopunctatus (Boulenger 1903 ) Proposed English name: Bluespotted Seabream Figures 1 B, 2 B; Table 1 Dentex lineopunctatus Boulenger 1903: 66, pl. 6 (type locality: KwaZulu-Natal coast, South Africa). Dentex natalensis Gilchrist & Thompson 1908: 156 (type locality: KwaZulu-Natal, South Africa). Polysteganus coeruleopunctatus (non Klunzinger); Smith & Smith 1986: 590 (in part, southwestern Africa); Heemstra & Heemstra 2004: 232 (in part, Red Sea to the Eastern Cape, South Africa, Western Indian Ocean); van der Elst 1985: 359, unnumbered color figure (in part, South Africa). Holotype. BMNH 1903.2. 6.5, 320 mm SL, KwaZulu-Natal coast, South Africa (Fig. 2 B). Other material examined. SAIAB 17470, 318 mm SL, 20 miles off Port Durnford, KwaZulu-Natal, South Africa (29 ��09' 36 ''S, 32 ��03' 49 ''E); SAIAB 29416, 189 mm SL, Coffee Bay, Eastern Cape Province, South Africa; SAIAB 37126, 288 mm SL, 20 miles off Port Durnford, KwaZulu-Natal, South Africa; SAIAB 48180, 148 mm SL, Mozambique; SAIAB 82139, 3 specimens, 92���196 mm SL (X), Mozambique (21 ��35.0'S, 35 �� 32.3 'E ��� 21 �� 33.6 'S, 35 �� 32.2 'E), bottom trawl, 17 October, 2007, collected by P.C. & E. Heemstra; SAM 9908 (1 of 2 syntypes of Dentex natalensis; 1 missing, M. Bougaardt pers. comm.), 283 mm SL, KwaZulu-Natal, South Africa, southwestern Indian Ocean, collected by R. Robinson. Diagnosis. Distinguished from all congeners of Polysteganus by the following combination of characters: scale rows above lateral line 7 ��; scales rows between 5 th dorsal-fin spine base and lateral-line 5 ��; higher counts of pored lateral-line scales (53���55); suborbital depth clearly less than or subequal to orbit diameter; dorsal-fin rays reddish pink or orangish red but with deep yellow upper margin of dorsal-fin rays; 7 or 8 distinct irregular lines of blue spots on body (often present even in preserved specimens); pale reddish yellow membranes on caudal-fin lobes and caudal fin with hyaline white margin on lower fin lobe; no blue iridescence on dorsal and lateral side of whitish pink body; canines somewhat smaller, only upper canines visible when mouth closed; body slender, 40��� 43 % (mean 42 %) of SL; no bony bulge before and above eye even in larger specimens of over 25 cm TL; caudal fin moderately forked. Description. Counts and proportional measurements as percent of SL of the holotype and other specimens (included as synonymized type) of Polysteganus lineopunctatus are given in Table 1. Body depth relatively deeper; head robust, rostro-occipital profile strongly convex; interorbital scaly, scales extending to nearly above anterior nostrils and just above posterior nostrils, anterior nostrils small and rounded, posterior nostrils oval; orbit diameter greater than interorbital width; maxilla naked, its posterior end not reaching vertical at front edge of eye, rear margin of maxilla not concealed by infraorbitals; jaws robust, especially upper jaw; canines slender and very short, 4 in upper and 6 in lower jaw (two outermost canines in upper jaw larger than 2 medial canines, and two outermost canines of lower jaw larger than 4 medial canines; an outer lateral row of conical teeth and a series of villiform teeth on inner side of outer lateral teeth row; mandibular profile slightly convex; rear margin of preopercle rounded and feebly serrated, a narrow naked area (preopercle with keel-like ridge), between preopercle flange scaly area and cheek scales; first to third dorsal-fin spines progressively longer, fourth dorsal-fin spine longest, slightly greater than snout length; no sheath scales on dorsal and pelvic-fin rays; predorsal length subequal to or slightly less than body depth, and clearly less than dorsal-fin base length; lateral line gently curved, almost parallel to curved profile between dorsal outline and lateral line, giving an impression of a more slender body than that of the similar species, P. coeruleopunctus (Figs. 1���2); pectoral-fin base scaly and fin rounded posteriorly, two or three lowest rays slightly extended posteriorly, compared with rays above (Fig. 2 B, of the holotype shows this feature), fin tip reaches vertical at anus, its length clearly less than head length; first pelvic fin ray definitely greater than fourth dorsal-fin spine, fin tip not reaching beyond vertical at pectoral-fin tip and rear margin of anus; soft dorsal and anal fins with no scales; anal fin with moderate scaly sheath, anal-fin spines slender, second and third subequal, and slightly greater than orbit diameter; caudal fin lobes broad and wide, caudal fin moderately forked (Figs. 1 A, 2 A); scales ctenoid; gill-rakers lanceolate; vertebrae 10 + 14. Color when fresh, based on color photographs of specimens (SAIAB 82139, 198 mm SL, Mozambique, and uncatalogued SAIAB photograph, 275 mm TL, Mozambique, photographed by O. Alvheim; Fig. 1 B): head and body uniform reddish orange with silvery reflection at center of each scale, color of upper head and dorsolateral part of body brighter than lower head and abdominal part of body; iris golden and pupil black; all fins uniform reddish orange, but membrane of spinous dorsal-fin hyaline reddish, and more whitish in pelvic and anal fins. Color of preserved specimens: upper sides of head and trunk uniform yellowish-tan. Distribution. Polysteganus lineopunctatus (Boulenger 1903) is currently known only from South Africa and Mozambique, and is endemic to the southwestern Indian Ocean. No confirmed reports of this species from Madagascar. Remarks. Polysteganus lineopunctatus has long been considered a junior synonym of P. coeruleopunctatus (Klunzinger 1873) but is justified as a valid species in this study. The counts and proportional measurements of type material show it to be different (Table 1). Genetic data (Fig. 4) also supports the validity of P. lineopunctatus. Polysteganus lineopunctatus differs from P. coeruleopunctatus in having 5 �� scales rows between 5 th dorsalfin spine base and lateral line (vs. 5 �� or 6 �� scale rows in the latter); 53���55 pored lateral-line scales (vs. 50 or 51 pored lateral-line scales); very weak bony bulge before and above eye, forming a low median ridge (vs. weak bony bulge, becoming prominent in specimens over 25 cm SL); anal-fin rays whitish pink (vs. anal-fin rays pinkish basally); with 7 or 8 distinct irregular blue spotted lines on small to large adults (vs. 7 or 8 faint irregular blue spotted lines). Dentex natalensis Gilchrist & Thompson 1908 also has long been synonymized under Polysteganus coeruleopunctatus (Klunzinger 1870), but one of two syntypes (SAM 9908) clearly has the same counts and morphometric characters as P. lineopunctatus, shown in Table 1. Another syntype was lost in 2009 (M. Bougaardt of SAM, pers. comm.). We accordingly conclude that Polysteganus natalensis (Gilchrist & Thompson 1908) is a junior synonym of P. lineopunctatus (Boulenger 1903)., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2015, Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean, pp. 133-150 in Zootaxa 4059 (1) on pages 140-141, DOI: 10.11646/zootaxa.4059.1.7, http://zenodo.org/record/246088, {"references":["Boulenger, G. A. (1903) Descriptions of six new perciform fishes from the coast of Natal. Annals of the South African Museum, 3 (3), 63 - 67, pls. 2 - 7.","Gilchrist, J. D. F. & Thompson, W. W. (1908) Descriptions of fishes from the coast of Natal. Annals of the South African Museum, 6 (2), 145 - 206.","Smith, J. L. B. & Smith, M. M. (1986) Family No. 183: Sparidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smiths' Sea Fishes. Macmillan, Johannesburg, pp. 580 - 594.","Heemstra, P. C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. NISC and SAIAB, Grahamstown, xxiv + 488 pp.","Klunzinger, C. B. (1870) Synopsis der Fische des Rothen Meeres. I. Theil. Percoiden-Mugiloiden. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 20, 669 - 834."]}

Polysteganus cerasinus new species Proposed English name: Cherry Seabream Figure 4; Table 2 Holotype. SAIAB 84183, 134 mm SL (X), sex not determined, Saya de Malha Bank, Mascarene Plateau (19.10 �� 54.75 'S, 60 �� 58.17 'E), depth 127 m, Trawl No. Mascarene- 19, coll. by D. Tweddle & O. Alvheim, 30 October 2008. Diagnosis. Distinguished from all congeners of Polysteganus by the following combination of characters: scale rows above lateral line 6 ��; scales rows between 5 th dorsal-fin spine base and lateral line 4 ��; lower counts of pored lateral-line scales 49; suborbital depth clearly less than orbit diameter (OD), 60 % OD; no lines of blue spots on body; dorsal-fin rays reddish pink but yellowish membranes of both pinkish dorsal-fin and caudal-fin fin lobes, no whitish margin of lower margin of lower caudal fin lobe; no blue iridescence on body but bright whitish iridescence on each body scale of lateral side of bright pinkish body; body deep, 43 % SL; no bony bulge before and above eye; pinkish upper and lower caudal-fin as well as rear pinkish margin of the caudal fin with yellowish hyaline membrane of most of caudal fin with no whitish upper and lower margins; rear margin of membrane of branchiostegal rays yellow, with yellow hue on some body scales just above upper pectoral-fin base (Fig. 4); no bony bulge before and above eye; caudal fin moderately forked. Description. Counts and proportional measurements as percent of standard length of the holotype of Polysteganus cerasinus n. sp. are given in Table 2. Body depth relatively deeper; head robust, rostro-occipital profile nearly straight; interorbital scaly, scales extend to nearly above anterior nostrils and just above posterior nostrils; anterior nostrils small and rounded, posterior nostrils oval; orbit diameter clearly greater than interorbital width; maxilla naked, its rear end nearly reaching vertical at front edge of eye when mouth closed, rear margin of maxilla not concealed by infraorbitals; jaws robust, especially upper jaw; canines short and small, 4 in upper and 6 in lower jaw (but not visible to naked eye; 2 outermost canines in upper jaw somewhat larger than 2 medial canines, and 2 outermost canines of lower jaw somewhat greater in size than 4 medial canines if seen through binocular microscope); an outer lateral row of conical teeth and a series of villiform teeth, on inner side of outer lateral teeth row; mandible straight; rear margin of preopercle rounded and minutely serrate; narrow naked area between preopercle flange scaly area and cheek scales; first to third dorsal-fin spines progressively longer, fourth dorsal-fin spine longest, and clearly greater than snout length; no sheath scales on dorsal and pelvic-fin rays; predorsal length clearly greater than body depth, and definitely less than dorsal-fin base length; lateral line gently curved, subequally parallel to curved dorsal profile between dorsal outline and curved lateral line; pectoral-fin base scaly and somewhat pointed posteriorly, 2 or 3 lowest rays slightly extended posteriorly, compared with rays above (Fig. 2 B, visible in the holotype of this species if seen through binocular microscope), the fin tip reaching vertical at anal fin origin, its length subequal to head length; first pelvic-fin ray length definitely greater than fourth dorsal-fin spine length, fin tip not beyond vertical at pectoral-fin tip and not beyond rear margin of anus; soft dorsal- and anal-fins with no scales; anal fin with low scaly sheath, anal-fin spines slender, second subequal to third, clearly less than orbit diameter; caudal fin moderately forked; scales ctenoid; gill-rakers lanceolate; vertebrae 10 + 14. Color when fresh, based on color photograph of the holotype (photographed by O. Alvheim; Fig. 4): head and body uniform pink with silvery reflection on each scale, color of upper part of head and dorsolateral part of body brighter pink than lower part of head and abdominal part of body; iris golden and pupil black; all fins uniform pinkish hyaline, especially in hyaline of membrane of pectoral fins; pinkish upper and lower caudal-fin as well as rear pinkish margin of the caudal fin with yellowish membrane of greater part of caudal fin with no whitish margin on lower margin of lower caudal fin lobe (central part of membrane and caudal-fin rays of caudal fin yellow and with pinkish margin); head and body uniform reddish orange with golden yellow reflection at center of each scale, color of dorsal head and dorsolateral part of body brighter than ventral part of head and abdominal part of body; iris golden and pupil black; all fins uniform reddish orange, but membrane of spinous dorsal-fin reddish orange. Color of preserved specimens: upper sides of head and trunk uniform yellowish-tan. Distribution. Polysteganus cerasinus n. sp. is currently known only from Saya de Malha Bank (19.10 �� 54.75 'S, 60 �� 58.17 'E), at a depth of 127 m. This species is presumably more likely to inhabit the somewhat shallower coastal waters around the Mascarene Islands than other Mascarene Polysteganus species: Polysteganus baissaci, P. flavodorsalis n. sp. and P. mascarenensis have all been trawled deeper than 200 m. Etymology. This fish is named cerasinus, ���of the cherry���, alluding to its cherry blossom colour. Remarks. Two other Mascarene species, P. flavodorsalis n. sp., and P. mascarenensis, were collected at depths of over 200 m (Iwatsuki & Heemstra 2011). Further biological information and additional collection data are urgently required for this species., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2015, Redescriptions of Polysteganus coeruleopunctatus (Klunzinger 1870) and P. lineopunctatus (Boulenger 1903), with two new species from Western Indian Ocean, pp. 133-150 in Zootaxa 4059 (1) on pages 144-146, DOI: 10.11646/zootaxa.4059.1.7, http://zenodo.org/record/246088

Ho, Hsuan-Ching, Heemstra, Phillip C., Imamura, Hisashi (2014): A new species of the sandperch genus Parapercis from the western Indian Ocean (Perciformes: Pinguipedidae). Zootaxa 3802 (3): 335-345, DOI: http://dx.doi.org/10.11646/zootaxa.3802.3.3

Parapercis albiventer sp. nov. New English name: Whitebelly sandperch Figures 1A–E, 2A–C, 4A–B. Tables 1 –2 Parapercis xanthozona (non Bleeker): Playfair & Günther, 1867:69, pl. VIII, fig. 3 (Zanzibar). Heemstra, 1986:741, pl. 113, 234.8 (South Africa, with uncertainty). Holotype. SAIAB 15019 (female, 143.2 mm SL), 26°53'S, 32°52'E, off Kosi Bay, KwaZulu-Natal, South Africa, 18.3 m, hook and line, coll. P. C. Heemstra, 1 Nov. 1980. Paratypes. 11 specimens, 63.0– 168.2 mm SL (* indicates specimen with proportional data taken). BMNH 1867.3.9.511* (1 female, 125.1), Tanzania, western Indian Ocean, no other data. MNHN 1992-892 * (1 female 124.2, 1 male 142.3), Vonhira, Madagascar, western Indian Ocean, 15 Mar. 1967. SAIAB 9430 * (1 male, 148.0), 27°31'S, 32°41'E, Sodwana Bay, KwaZulu-Natal, South Africa, coll. P. C. Heemstra et al., 2 Apr. 1979. SAIAB 46245 (1, 70.2), 30°18'S, 30°49'E, Aliwal Shoal southern ledges, KwaZulu-Natal, South Africa, ichthyocide, P. C. Heemstra & C. Buxton, 18 Jun.1994. SAIAB 50377 (1, 63.0), 26°46'S, 32°53'E, Ponta Malongane, Mozambique, 24-26 m, ichthyocide, coll. P. C. Heemstra, 3 Nov. 1995. SAIAB 60347 * (1 female, 140.1), 26°24'S, 32°55'E, inner pinnacle, Ponta Milibangalala, Mozambique, 19 m, ichthyocide, coll. P.C. & E. Heemstra & A. Bentley, 2 May 1999. SAIAB 97481 * (1 female, 154.7), 25°1.7'S, 46°59.7'E, Douane Beach, Fort Dauphin, Anosy, Madagascar, gill net, coll. P. C. Heemstra et al., 5 May 2010. SAIAB 98609 * (1 male, 158.9), 25°1.7'S, 46°59.7'E, Douane Beach, Fort Dauphin, Anosy, Madagascar, gill net, coll. P. C. Heemstra et al., 6 May 2010. SAIAB 188850 (1, 150), 26°40.2'S, 32°54.65'E, Techobanine, S. Mozambique, Africa, 20 m, spear, coll. A. D. Connell, 3 Jan. 2011. SAIAB 191147 * (1 male, 168.2), 30°16.8'S, 30°49.2'E, Aliwal Shoal, South Africa, 31 m, ichthyocide, coll. P. C. Heemstra, 26 Sep. 2002. Diagnosis. A species of Parapercis with a bright white ventral surface, without color markings on lower fourth of body; a row of 10 reddish blotches present at vertical through lower half of pectoral-fin base, the lower part of each blotch with a solid black bar ventrally. It can also be distinguished from its congeners by the following characters: dorsal-fin rays V, 21; anal fin rays I, 17; 3 pairs of enlarged canines at front of lower jaw; pored lateralline scales 55–59 (mainly 58); pectoral-fin rays 17–18 (mainly 17); gill rakers 16–19 in total; and a different coloration: 3 pairs of dark broad bands on upper lip; curved black band on front of underside of chin; medium-sized orange-red irregular spots on entire dorsal surface of head; small irregular spots densely covering nape and dorsal surface of body; row of faint red blotches, on paler background, along body axis; caudal fin with two black longitudinally elongate bars on anterior half, black spots on upper and lower thirds, central third of posterior margin of fin solid black, and a bright white patch on posterior half of middle caudal-fin rays, surrounded by irregular black marks. Description. The following data are provided for the holotype, followed by all measured/counted specimens in parentheses when different, except for other indications. Dorsal-fin rays V, 21 (1 paratype with missing 3rd or 4th spine; 1 with missing 5th spine); anal-fin rays I, 17; all dorsal- and anal-fin rays branched, last to base; pectoral-fin rays 17 (16 or 17, mainly 17), upper ray unbranched, others including lowermost branched; pelvic-fin rays I, 5; branched caudal-fin rays 15; lateral-line scales 59 (55–59, mainly 58), not including 3 (3–5) smaller pored scales on caudal-fin base; scales above lateral line to origin of dorsal fin 6, to base of anterior soft rays of dorsal fin 6.5 (5.5 or 6.5); scales below lateral line in oblique row to origin of anal fin 19 (17–21, mainly 19); circumpeduncular scales 29 (29 or 30); predorsal scales 9 (9 or 10), extending just anterior to a vertical from preopercle margin; gill rakers on 1st arch 6+12=18 (5–7+10–12=16–19); branchiostegal rays 6; vertebrae 10 + 20=30 (5 specimens, including the holotype, examined). Body elongate, relatively depressed anteriorly, becoming rounded and cylindrical medially, compressed posteriorly. Body depth at head 13.6% (12.7–15.7%) SL; body width at head 17.2% (17.1–18.8%) SL; head depressed, particularly anterior portion, elongate; head length 30.8 (29.3–30.8%) SL; snout pointed, its length 10.8% (9.0–10.8%) SL; orbital diameter 6.1% (5.0–6.7%) SL; eyes directed more laterally than dorsally, fleshy interorbital space narrow, 3.0% (2.3–3.1%) SL; caudal-peduncle depth 8.9% (8.0–8.9%) SL; caudal-peduncle length 8.4% (7.2–9.0%) SL. Mouth horizontal to slightly oblique, lower jaw protruding, with curved canine teeth at front of lower jaw slightly projecting, barely visible when mouth is closed; upper jaw extending to vertical at middle of eye (slightly beyond vertical at anterior margin of eye in small specimen), upper-jaw length 12.1% (11.0–12.5%) SL; upper jaw with 26/27 (25–28) outer curved, subequal canines on each side, first 2 slightly larger; broad inner band of villiform teeth anteriorly, narrowing gradually to form two rows at rear of jaw; front of lower jaw on each side with 3 enlarged curved canines in distinctly separate outer row; teeth nearest symphysis smallest, the others gradually increasing in size laterally, last in the series largest; broad inner band of villiform teeth extending posteriorly from symphysis to side of jaw just posterior to largest tooth in outer row; next 9/10 (9–11) teeth in single row, gradually curved and abruptly increasing in size, followed by single row of 12 (10–12) smaller slightly curved conical teeth. Vomer with 3 irregular crescentic rows of subequal teeth, those on 1st row slightly larger. No teeth on palatines. Tongue spatulate with broadly rounded tip. Gill membranes united with broad free fold, not attached to isthmus. Gill rakers short, longest about 2–3 times in length of longest gill filament on 1st gill arch. Anterior nostril small, inconspicuous, anterior to mid-eye, slightly less than half distance from anterior margin of eye to snout tip, with membranous posterior flap, the latter folded around base forming partial tube, flap usually lying flat against snout in preserved specimens. Posterior nostril slightly more than half distance from anterior margin of eye to anterior nostril, dorsoposterior to, and about same width as anterior nostril, its opening simple, oval; internarial distance subequal to diameter of posterior nostril. Opercle with distinctly exposed, robust, pointed spine; angle of subopercle expanded and armed with a cluster of 4 (4 or 5) small poorly defined spinules, remainder of subopercle entire, margins naked and broadly rounded; posterior margin of preopercle free from the gill cover, weakly serrated, with many poorly defined, widely-spaced knobs. Lateral line continuous, ascending smoothly from opercle to below second or third soft dorsal-fin ray, then approximately following contour of back; scales on body ctenoid; on nape cycloid anteriorly and weakly ctenoid posteriorly; on cheeks cycloid, except for a few ctenoid scales near posterior margin of preopercle; ctenoid scales on space between pectoral-fin base and gape of gill chamber; cycloid on breast; cycloid anteriorly on midline of belly, weakly ctenoid posteriorly; scales on cheek extending forward approximately to vertical at anterior margin of orbit; no scales on dorsal, anal or pelvic fins; 4 (3 or 4) rows of small feebly ctenoid scales on base of pectoral-fin rays, except for the very last scales which are cycloid; elongate ctenoid scales densely arranged on proximal threefourths of caudal fin. Origin of dorsal fin just posterior to vertical from tip of opercular flap, predorsal length 32.5% (30.5–33.4%) SL; first four dorsal-fin spines progressively longer, first short, 1.7% (1.5–2.7%) SL; 3rd (3rd or 4th, usually 4th) spine longest, 5.0% (4.8–6.3%) SL; 5th dorsal-fin spine slightly shorter, 3.2% (3.1–4.4%) SL; fifth spine connected by membrane to opposite tip of the spine on 1st soft dorsal-fin ray; 3rd to 15th dorsal-fin soft rays subequal, the longest ray 9.9% (9.2–12.2%) SL; origin of anal fin below base of 5th soft dorsal-fin ray, preanal length 50.0% (48.6–51.3%) SL; anal-fin spine slender, closely attached to 1st soft ray, 3.7% (3.6–4.4%) SL; penultimate anal-fin ray longest, 10.5% (9.7–11.5%) SL; caudal fin truncated to slightly rounded, its length 17.2% (16.5–20.0%) SL; short elongation on upper lobe in most specimens examined, absent in other specimens; pectoral fins rounded, 9th (9th or 10th) ray usually longest, 18.5 (17.3–18.7%) SL, subequal to or slightly longer than pelvic fins; origin of pelvic fins in advance of upper base of pectoral fins and approximately on vertical from upper corner of operculum, prepelvic length 26.7% (24.1–27.7%) SL; pelvic-fin spine closely attached to first soft ray, its termination fleshy and difficult to accurately determine; 4th soft pelvic-fin ray longest, not reaching anus (just anterior to vent in larger specimens, to anal-fin origin in smaller specimens), 20.2 (17.8–23.0%) SL. Pores of cephalic sensory system with row of four large pores above maxilla, one above vertical at anterior nostril, other between verticals at lower half of eye; four pores near nostrils, two above, one between and one below; irregular row of three pores on either side anteriorly in interorbital space; irregular series of small pores medially on posterior half of interorbital space; numerous small pores with subcutaneous canals posteriorly on occiput, pores divided into several irregular double series, from anterior end of lateral line on body, continuing to ventral margin of eye, opercle and subopercle; cheek with 8 (8 or 9) irregular double pore series; irregular double series of small pores (up to 30 in number) along margin of preopercle; series of four large pores on mandible; single large pore at front of chin. Color in life (Figs. 2A–C). Body ground color reddish, with ventral surface bright white; medium orange-red blotches, about pupil size, on dorsal surface of head, eye, cheek and gill cover; smaller irregular reddish spots densely covering dorsal surfaces of nape and body; interspaces of these patches and spots pale red or blue; 3 short deep-red bands on each side of upper lip; pupil black. A series of 10 faint reddish blotches encircled by paler color along a vertical of pectoral fin base; 10 deep-red blotches, subequal to eye, with short solid dark bar on underside of blotches, interspace of blotches bright white with small pale red spot. Small black patch on spinous dorsal fin, between bases of 3rd and 4th spines; 2 rows of deep red spots on soft dorsal fin; pectoral fin reddish, nearly transparent; pelvic fin bright white; anal fin not visible from these photographs; caudal fin with scattered irregular black spots in upper and lower thirds of the fin; 2 longitudinally-elongate black blotches at middle of anterior half of fin; an irregular bright white blotch at middle of posterior half of fin; middle posterior margin of fin black. Color in freshly caught specimens (Figs. 1A, C–E). Upper three-fourths of body reddish and lower fourth of body bright white. Medium red patches, about pupil size, on dorsal surface of head, eye, cheek and gill cover; smaller irregular red spots densely covering nape and dorsal surface of head; three short black bands on upper lip; pupil black. A broad faint red stripe about same height as pectoral fin base along body axis; a row of 10 deep-red blotches, each with a black bar on lower margin, evenly distributed, first below pectoral fin; interspaces white with reddish spots. A small black spot between bases of 3rd and 4th dorsal-fin spine; two rows of small spots on soft dorsal-fin; row of slightly larger black spots on membrane of anal fin; caudal fin with scattered irregular black spots in upper and lower thirds of fin; two longitudinally-elongate black blotches at middle of anterior half of fin, sometimes connected posteriorly; irregular bright white blotch at middle of posterior half of fin; upper and lower thirds of posterior caudal-fin margins reddish; middle third of posterior caudal-fin margin black. Color in preservative (Figs. 1B, 4A–B). Body uniformly yellowish brown; some small brown patches on dorsal surface of head, eye, cheek and gill cover; small faint dots, each about one scale in width, loosely covering dorsal surface of nape and body; three short black bands on each side of upper lip; underside of chin with a curved bar (divided into two spots in holotype and some paratypes). A row of 10 short dark brown bars evenly distributed from vertical at pectoral-fin base, first bar below pectoral fin base. A paler brown spot between short bars; two rows of brown spots on dorsal fin, those on upper row associated with fin membranes, those on lower row associated with fin rays; row of black spots on anal fin associated with fin membranes near the fin base (absent or fewer in number and restricted to posterior portion of the fin in smaller paratypes); caudal fin with scattered irregular black spots in upper and lower thirds of fin; two longitudinally-elongate black blotches at middle of anterior half of fin, sometimes connected posteriorly; no bright white blotch on rear middle half of fin; middle third of posterior caudal-fin margin black. Etymology. The specific name, albiventer, is from the Latin for white ventral surface, in reference to the bright white immaculate color on ventral fourth of the body. Size. Largest specimen examined is 168.2 mm SL. Distribution. The type series was collected from Madagascar, southern Mozambique and South Africa. Literature records for the species (misidentified as P. xanthozona) include Zanzibar (Playfair & Günther, 1867) and South Africa (Heemstra, 1986). Specimens were collected at depths 18.3–31.0 m, mainly at depths less than 30 m. Sexual dimorphism. Most specimens were dissected to determine sex, including four mature males and five mature females, all with well-developed gonads. The range for mature males is slightly larger (142.3–168.2 mm SL) than that of females (124.2–154.7 mm SL). Other specimens are either immature or indeterminate. No distinct differences were observed from external morphology, except that males have two or three faint oblique bands on the cheek whereas the female has fainter and somewhat shorter bands than those of the males. Comparisons. As mentioned above, P. albiventer sp. nov. shares a row of blotches (black bars or spots in preservative) on the lower portion of the body and other similar features of its color pattern with a number of congeners. Of these, the new species is most similar to P. stricticeps (Figs. 3A–B, 4C) from eastern Australia in having a similar body coloration and distribution of spots on the median fins. It differs from P. stricticeps in having 9 or 10 predorsal scales and 55–58 pored lateral-line scales (vs. 10–12 and 57–59, respectively); medium-sized patches on the head, cheek and gill covering (vs. fewer and scattered small black spots on dorsal surface of the head and oblique bars on cheeks and/or gill covering); irregular reddish spots on the nape and dorsal surface densely and evenly distributed (vs. brown spots forming 9 or 10 regular dotted blotches on dorsal surface); no spots on underside of the chin, except for a curved bar at front, and mandible (vs. underside of head including mandible with broad dark bars); a pale chest (vs. usually 3–5 dark spots on chest); and blotches on the lateral body ending as a black bar (vs. blotches extending to ventral surface). The key for species provided by Cantwell (1964) and Randall (2001) did not include P. stricticeps, which was identified as P. xanthozona (Fig. 4B), a species known from the eastern Indian and western Pacific oceans. Our new species differs from P. xanthozona by having blotches on the lower lateral body short, each ending with a black bar ventrally, not extending into lower quarter of the body (vs. no black bar associated with lateral body blotches and blotches long, extending to the ventral surface); black bars on upper lip (vs. absent or very faint when present); row of black spots on basal portion of the anal fin (vs. on middle portion of the fin); medium-sized patches on the head (vs. absent from the head and 1 or 2 oblique stripes on cheek); large irregular black marks on the caudal fin, middle third of the posterior caudal fin margin black (vs. spots on caudal fin, middle third of the posterior caudal fin with dusky margin, but never solid black); spots on the dorsal fin small, not crossing the fin membranes (vs. mid-row of spots extending across fin membranes); pale strip on the body axis about same width as the pectoral-fin base (vs. slightly above the body axis and much narrower); no orange bar on the pectoral-fin base (vs. orange bar present); bright white patch on posterior half of the caudal fin surrounded by black marks or spots (vs. white patch on central portion of the caudal fin, not surrounded by black marks or spots). The five dorsal-fin spines distinguish the new species from P. bimacula, P. clathrata and P. millepunctata, which have modally 4 dorsal-fin spines. The three pairs of canines on front of the lower jaw separate the new species from P. kamoharai and P. multiplicata, which have modally four pairs of canines. The relatively few pored lateral-line scales separate the new species from P. robinsoni (77–84 scales; data from Randall & Stroud, 1985). The new species is sympatric with P. punctulata (Fig. 5A) in the western Indian Ocean and allopatric with P. tetracantha (Fig. 5B) from the eastern Indian and western Pacific oceans. They have a row of black blotches on lower half of body when preserved, but can be easily distinguished from the new species by lacking large black and white marks on the caudal fin. Parapercis albiventer also differs from a recently described species, Parapercis maramara Sparks & Baldwin, 2012, from Madagascar, in having a relatively short pelvic fin (reaching vent in larger specimens and origin of anal fin in smaller specimens vs. reaching 2nd ray of anal fin); a black bar, sometimes breaking into 2 black spots, on the chin (vs. no black bar on chin); a broad, faint red stripe on body axis (vs. a row of pale patches usually with a black central spot on body axis). Comparative material. Parapercis stricticeps: QM I.1708 (153.3), holotype of Percis stricticeps, dry specimen, Southport, Queensland, Australia. QM 23072 (3, 114.2-140.2), 21°18'S, 151°19.8'E, off Swain Reefs, 58-61 m, 14 Sep. 1986. QM 31316 (1, 159.1), 26°39'S, 153°11'E, Gneering Shoals, 10 km E of Mooloolaba, 19 m, 14 Oct. 1998. Parapercis xanthozona: RMNH 5946 (140.4), holotype of Percis xanthozona, Jakarta, Java, Indonesia. ASIZP 72225 (1, 165), Jin-shan, Taiwan, 31 Oct. 2009. HUMZ 122201 (1, 150.6), no data. MNHN 1980-648, 1980-649 (3, 47.4

Acanthopagrus catenula (Lacep��de 1801) New English name: Bridled Seabream Figs. 1 B, 2 B, 3 E���H, 4 C���D & 5 B Labrus catenula Lacep��de 1801: 426, 468, plate 3, figure 3 (type locality: Mauritius). Sparus mylio Lacep��de 1802: 41, 131 (type locality: Mauritius). Holotype. MNHN A- 0961, dried, 171 mm SL, Mauritius. Non-type specimens (19 specimens, 101���364 mm SL). BPBM 27366, 166 mm SL, Natal, South Africa; MNHN A- 4149, 2: 101���104 mm SL, Madagascar; MNHN 1981 -0812, 114 mm SL, Madagascar; MUFS 25806���25808, 3: 266���364 mm SL, off Socotra Island, Somalia, trawls: MUFS 33573���33575, 3: 231���255 mm SL, Madrakah, southern Oman 6 Oct., 2010, hand-line; SAIAB 9672, 139 mm SL, Kosi Bay, KwaZulu-Natal, South Africa; SAIAB 11819, 153 mm SL, Malindi, Kenya; SAIAB 11820, 167 mm SL, Mozambique; SAIAB 11821, 141 mm SL, South Africa; USNM 29538, 4: 280���330 mm SL, north coast of Somalia, southeasternmost part of Gulf of Aden (11 �� 54 '06"N, 51 ��03' 12 "E), trawls. Diagnosis. Distinguished from other species of Acanthopagrus by the following combination of characters: two conspicuous vertical black bars across the head (Figs. 1���3); dorsal fin with wide black margin, especially distinct in distal spinous part of dorsal fin in fish < ca. 30���36 cm SL (Fig. 2 B, 3 E���H), proximal part of dorsal fin yellow; hind margin of caudal fin with narrow black edge in fish to around 30 cm SL, and then disappearing with growth (Fig. 2 B, 3 E���H); scale rows between fifth dorsal-fin spine and lateral line 4 ��; third outer molar teeth row extending to rear end of both jaws; (Fig. 4 B), basal width of 6 enlarged incisor teeth at front of upper jaw larger, 1.0���1.1 mm for 16 cm SL fish, and 5.0���5.5 mm for 34 cm SL fish (Fig. 5 B); dorsal-fin rays XI, 12���14; anal-fin rays III, 10; orbit diameter (Fig. 6), noticeably larger in fish> 15 cm SL. Description. Counts and measurements of the holotype and 20 non-type specimens of Acanthopagrus catenula are given in Table 1. Body deep, compressed; mouth somewhat oblique, lips thick; maxilla reaching to vertical below middle of pupil in young and to vertical at rear edge of pupil in fish over 25 cm SL (Fig. 3 E & F); lower jaw slightly included, especially in fish larger than 35 cm SL; upper jaw usually with 6 distinct incisor-like teeth in juveniles, becoming curved canines in adults; front of lower jaw with 6 canines (Fig. 5 C & D) but 4 middle canines becoming considerably larger than outer teeth in adults (Fig. 6 C & D); upper jaw generally with 6 molar tooth rows anteriorly, 4 rows of larger molar teeth posteriorly, innermost teeth smallest, but outermost teeth at mid-jaw also well-developed; lower jaw with 4 molar tooth rows, teeth small anteriorly, increasing in size and forming 3 rows posteriorly; least infraorbital depth less than eye diameter in juveniles and greater than orbital diameter in adults> 30 cm SL; cheek scales in 4 or 5 transverse rows; occipital profile weakly curved, somewhat gibbous head above eye in specimens> 30 cm SL (Fig. 3 H); dorsal-fin spines strong, heteracanthous, first spine slightly more than half length of second, second clearly shorter than third; fourth spine longest; first dorsal-fin ray shorter than longest spine; first anal-fin spine short, much less than eye diameter; second spine longest, but clearly less than head length; third anal-fin spine less than first anal-fin ray; pectoral fins longer than head, tip reaching beyond vertical at second or third anal-fin spine base in fish Colour of fresh specimens. Based on specimens from South Africa, off Socotra Island, Somalia, and Rodrigues (not kept, ca. 30 cm TL, photographed by T. Hooper and MUFS 25806���25808, 266 ��� 364 mm SL, Fig. 4 H for MUFS 25808). Head and body silvery, sometimes with a black spot at center of each scale (spots more distinct in live specimens or some specimens from off the Horn of Africa, Somalia) forming dotted lines following scale rows parallel to lateral line above lateral line, and also (faintly) below lateral line (Fig. 3 G & H); snout and infraorbital area yellow, especially in fish> 20 cm SL (Fig. 3 H); dorsal, caudal and pectoral fins yellow in young and vivid yellow in specimens over 20 cm SL; dorsal fin with a wide black margin; caudal fin with a narrow black edge but diappearing with growth in specimens> ~ 30 cm SL; pectoral fins often black with a yellow base; juveniles with hyaline yellow pectoral fin, becoming vivid yellow at base on fish> 20 cm SL (Fig. 2 B, 3 G & H); pelvic- and analfin rays dense black; posterior part of anal fin often hyaline in juveniles and presumably first maturity size, 25 cm SL; Fig. 2 B, 3 G), and becoming black in larger adults over 35 cm SL; Fig. 3 H). Colour of preserved specimens. Head and body pale yellowish tan, yellow of dorsal and caudal fins faded, but black margin of dorsal-fin and black rear edge of caudal fin usually remain, caudal fin dark edge disappearing in largest specimen (MUFS 25808, 364 mm SL; Fig. 3 H); faint dotted lines along scale rows often remain but disappears in old specimens; black of pelvic and anal-fin rays and the two conspicuous black bars on head become fainter (Fig. 3 E & F). Distribution. Acanthopagrus catenula is currently known from off the Horn of Africa to off Socotra Island of Somalia, including Kenya, Mozambique, South Africa, Madagascar, Mauritius, and Rodriguez. Recently, A. catenula was collected in Madrakah, southern Oman (MUFS 33573���33575, 3 specimens, 231���255 mm SL, 6 Oct., 2010) during the first author���s visit. Remarks. Lacep��de (1801) described Labrus catenula on the basis of a holotype (MNHN A- 0961, dried, 171 mm SL) from Mauritius. The dark posterior edge of the caudal fin in the dried holotype is still clearly visible as are the counts of 4 �� scale rows between the fifth dorsal-fin spine and lateral line, being diagnostic characters of this species. Unfortunately, the black margin of the dorsal fin was not visible because no dorsal-fin membrane remains on this dried specimen (Fig. 1 B)., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2011, A review of the Acanthopagrus bifasciatus species complex (Pisces: Sparidae) from the Indian Ocean, with redescriptions of A. bifasciatus (Forssk��l 1775) and A. catenula (Lacep��de 1801), pp. 38-50 in Zootaxa 3025 on pages 45-48, DOI: 10.5281/zenodo.205951, {"references":["Lacepede, B. G. E. (1801) Histoire naturelle des poissons. Tome Sixieme. Historie naturelle des poissons, 3, 1 - 558.","Lacepede, B. G. E. (1802) Histoire naturelle des poissons. Histoire naturelle des poissons, 4, 1 - 728."]}

Acanthopagrus bifasciatus (Forssk��l 1775) English name: Doublebar Seabream Figs. 1 A, 2 A, 3 A���D, 4 A��� B & 5 A Chaetodon bifasciatus Forssk��l 1775: 64, xiii (type locality: Jidda, Saudi Arabia, Red Sea). Holocentrus rabaji Lacep��de 1802: 724, 725 (type locality: Red Sea). Chysophrys bifasciatus; Valenciennes 1830: 118 (in part; distribution is that of both Acanthopagrus bifasciatus and A. catenula). Sparus bifasciatus; Fowler 1933: 160 (includes Sparus mylio and Labrus catenula). Holotype. ZMUC-P 50557, dried, 111 mm SL, Jidda, Saudi Arabia, Red Sea. Non-type specimens (37 specimens: 27���362 mm SL). BPBM 18173, 2: 158174 mm SL, Ras Muhammad, Sinai Peninsula, Egypt, Red Sea; BPBM 21294, 207 mm SL, Bahrain; BPBM 29526, 237 mm SL, Bahrain; MNHN 0000-0675, 145 mm SL, Eritrea (formerly Ethiopia), Red Sea; MNHN 0000- 8614, 2: 89���121 mm SL, Red Sea; MNHN 1902 -0161, 161 mm SL, Persian Gulf; MNHN 1966 -0090, 147 mm SL, Red Sea; MNHN 1977 -0953, 6: 28���44 mm SL, Israel; MUFS 25809���25811, 3: 285���362 mm SL, off southern Oman facing Indian Ocean, not Gulf of Oman; SMF 2196, 116 mm SL, Red Sea; SMF 21504, 47 mm SL, Persian Gulf; SMF 21750, 94 mm SL, Red Sea; SMF 24659, 101 mm SL, Red Sea; USNM 19168, 12: 37���75 mm SL, Red Sea; USNM 266730, 93 mm SL, Bahrain, Persian Gulf; USNM 374204, 2: 102���118 mm SL, Eritrea (Ethiopia), Red Sea. Diagnosis. Distinguished from other species of Acanthopagrus by the following combination of characters: two conspicuous vertical black bars across the head, the first bar from occiput through eye and across cheek to rear end of maxilla, the second bar from nape to opercle (Figs. 13); dorsal and caudal fins yellow, without dense black margin of dorsal-fin or narrow black edge along rear margin of caudal fin (Fig. 2 A, 3 A���D); 5 �� or 6 �� scale rows from 5 th dorsal fin spine to lateral line; base of dorsal and anal fins scaly; interorbital area and preopercle flange naked; 2 rows of molars along each side of upper jaw; outer molar teeth row extends to rear end of both jaws (Fig. 4 A); basal width of incisor teeth (usually 6) at front of upper jaw smaller, 0.7���0.8 mm for 15 cm SL fish and 2.2���2.5 mm for fish of 34 cm SL, respectively; Fig. 5 A); dorsal-fin rays XI, 12���15; no spines elongated; anal-fin rays III, 10���11; 2 nd and 3 rd anal-fin spines subequal, not enlarged. Description. Counts and measurements of the holotype and 37 non-type specimens of Acanthopagrus bifasciatus are given in Table 1. Body deep, compressed; mouth somewhat oblique, lips thick; maxilla reaching to vertical at middle of pupil; lower jaw slightly included; front of upper jaw with (usually) 6 incisors (young) that develop into curved canines in adults (Fig. 4 AB) but the 4 middle canines (of 6) generally become larger than the outer ones (Fig. 4 AB, 5 A); molar teeth of upper jaw in 6 rows on each side of jaw anteriorly, 4 rows of larger teeth posteriorly, innermost teeth smallest, but outer teeth at mid-jaw also well-developed; lower jaw with 4 molar tooth rows on each side, teeth small anteriorly, increasing in size posteriorly and forming 3 rows on each side posteriorly (Fig. 4 AB); least infraorbital depth less than eye diameter in young, but greater than eye diameter in fish> 30 cm SL (Fig. 3 A); 5 or 6 transverse rows of scales on cheek; occipital profile weakly curved; dorsal-fin spines strong, heteracanthous, first spine slightly more than half length of second, second clearly shorter than third; fourth spine longest; first dorsal-fin ray shorter than longest spine; first anal-fin spine short, much less than eye diameter; second spine longest, but clearly less than head length; third anal-fin spine shorter than second spine or first anal-fin ray; pectoral-fins longer than head, their tip reaching beyond vertical at first or second anal-fin spine base; pelvic fins clearly shorter than head; pelvic-fin spine longer than snout. 1. Holotypes of A Acanthopagrus bifasciatus and B A. catenula. A, ZMUC-P. 50557, 111 mm SL, Jeddah, Saudi Arabia, Red Sea; B, MNHN A- 0961, 171 mm SL, Mauritius. Colour of fresh specimens. Based on specimens from Red Sea (Egypt), Kuwait, Bahrain, Persian Gulf, and off southern Oman (BPBM 18173, Fig. 2 A, and BPBM 29526, Fig. 3 C, photographed by J. E. Randall; MUFS 25809���25811, Fig. 3 D for MUFS 25811). Head and body silvery, with a black spot at center of each body scale (especially distinct in live adult specimens or in Persian Gulf specimens), forming parallel lines following scale rows above and below lateral line; side of snout and cheek below eye yellow, more prominent in fish> 20 cm SL; dorsal, caudal and pectoral fins yellow in young and vivid yellow in fish> 20 cm SL; dorsal part of dorsal and pectoral fins often denser yellow than lower part, which is more hyaline yellow (Fig. 2 A, 3 C & D); yellow patch before eyes often present (Fig. 3 C); anterior snout region often dark brown or black (Fig. 3 C); pelvic- and anal-fin rays dense black, rear part of anal fin often transparent in fish less than 20 cm SL (Fig. 2 A, 3 A & B) and sometimes yellow in fish of 20���30 cm SL (Fig. 3 C) but dense black in 35 cm SL-specimen (Fig. 3 D). *, Dried right skin; **, Dired left skin; ***, Four specimens (USNM 29538, 4, specimens, 280���330 mm SL, off the Horn of Africa, Somalia) of A. catenula were measured only for standard length and counts of scale rows between the fifth dorsal-fin spine base and lateral line. Hyphen shows no measurement as ���-���. According to Randall (1995), Red Sea individuals of Acanthopagrus bifasciatus (Fig. 2 A) differ from Oman and Persian Gulf fish in lacking the rows of black spots, one per scale, on the body. However, this colour character was not always consistent (Fig. A [Red Sea but exact locality unknown showing visible spots], Fig. 3 B [Sinai Peninsula, northern Red Sea, no clear spots] and Fig. C [Bahrain, Persian Gulf fish showing distinct spots]). Furthermore, large A. bifasciatus from southern Oman���s Arabian Sea (facing the Indian Ocean) lacked distinct dark spots (MUFS 25809���25811, 3 specimens, 285���362 mm SL). Thus it appears that the rows of dark spots become fainter with growth and more distinct if the fish is stressed. Colour of preserved specimens. Head and body pale yellowish tan, yellow or vivid yellow coloration of dorsal and caudal fins faded; dark dotted lines following scale rows often visible; pelvic and anal-fin rays dense black as well as two conspicuous bars on head (Fig. 3 A & B). Distribution. Acanthopagrus bifasciatus is confirmed from the Red Sea (Israel, Egypt, Eritrea [formerly Ethiopia], and Saudi Arabia), Persian Gulf (Bahrain and Kuwait), and off southern Oman (not Gulf of Oman). The species presumably occurs along the whole southern coast of the Arabian Peninsula and is also found from Iran to Pakistan (fig. 5, Day 1875, plate 34, figure 5, based on Sind [= Pakistan]; Blegvad 1944). Remarks. Chaetodon bifasciatus was described from a specimen (Fig. 1 A) from Jeddah, Saudi Arabia, Red Sea by Forssk��l (1775). Lacep��de (1802) described Holocentrus rabaji as a new name for C. bifasciatus, which he apparently considered unacceptable. We consider that the species is not likely to occur in India, but it is known from Sind, Pakistan (Day 1875), which is the easternmost (known) distribution. Furthermore, A. bifasciatus is found in southern Yemen and Djibouti, but the specimens from the northern Somalian coast within the Gulf of Aden may actually be A. catenula. Consequently identification of A. bifasciatus from Somalia should be re-examined. In addition, Acanthopagrus bifasciatus (first females��� maturity size, mean 26.4 cm in fork length) in the southern Persian Gulf (UAE) spawns in January (Grandcourt et al. 2004), which is also a monsoon season when primary production in the Persian Gulf is high (Hirawake et al. 1998)., Published as part of Iwatsuki, Yukio & Heemstra, Phillip C., 2011, A review of the Acanthopagrus bifasciatus species complex (Pisces: Sparidae) from the Indian Ocean, with redescriptions of A. bifasciatus (Forssk��l 1775) and A. catenula (Lacep��de 1801), pp. 38-50 in Zootaxa 3025 on pages 39-42, DOI: 10.5281/zenodo.205951, {"references":["Forsskal, P. (1775) Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit. Post mortem auctoris edidit Carsten Niebuhr. Hauniae. Descr. Animalium Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit ... Post mortem auctoris edidit Carsten Niebuhr. Molleri, Copenhagen, 20 pp., xxxiv + 164 pp. (Pisces on pp. x - xix + 22 - 76).","Lacepede, B. G. E. (1802) Histoire naturelle des poissons. Histoire naturelle des poissons, 4, 1 - 728.","Fowler, H. W. (1933) Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes of the families Banjosidae, Lethrinidae, Sparidae, Girellidae, Kyphosidae, Oplegnathidae, Gerridae, Mullidae, Emmelichthyidae, Sciaenidae, Sillaginidae, Arripidae and Enoplosidae, collected by the United States Bureau of Fisheries steamer \" Albatross, \" chiefly in Philippine seas and adjacent waters. Bulletin of the United States of Natural Museum, No. 100, 12, 1 - 465.","Randall, J. E. (1995) Coastal fishes of Oman. Bathurst, Australia: Craford House Publishing Pty Ltd with University of Hawaii Press, pp. 1 - 439.","Day, F. (1875) The fishes of India; being a natural history of the fishes known to inhabit the seas and fresh waters of India, Burma, and Ceylon (Vol. 1). Member of Asiatic Society of Calcutta and Cotsworld Naturalists. Field Club, India and Burma, 168 pp., 40 pls.","Blegvad, H. (1944) Fishes of the Iranian Gulf. Einar Munksgaard, Copenhagen, 247 pp., 12 pls.","Grandcourt, E. M., Al Abdessalaam, T. Z., Francis, F. & Al Shamsi, A. T. (2004) Biology and stock assessment of the Sparids, Acanthopagrus bifasciatus and Argyrops spinifer (Forsskal, 1775), in the Southern Arabian Gulf. Fisheries Research, 69, 7 - 20.","Hirawake, T., Tobita, K., Ishimaru, T., Satoh, H. & Morinaga, T. (1998) Primary production in the ROPME Sea area. In: A. Otsuki et al. (Eds), Offshore Environment of the ROPME Sea after the War-Related Oil Spill. Terra Scientific Publishing Company, Tokyo, pp. 181 - 191."]}

Iwatsuki, Yukio, Heemstra, Phillip C. (2011): A review of the Acanthopagrus bifasciatus species complex (Pisces: Sparidae) from the Indian Ocean, with redescriptions of A. bifasciatus (Forsskål 1775) and A. catenula (Lacepède 1801). Zootaxa 3025: 38-50, DOI: 10.5281/zenodo.205951

The number of species of the goatfish genus Upeneus from the western Indian Ocean has increased considerably due to recent taxonomic efforts. Here we describe a new species, Upeneus randalli n. sp., based on body and fin coloration and 50 morphometric and meristic characters of eight specimens from the Persian Gulf, and supported by comparative data from a taxonomic review of western Indian Ocean Upeneus species and additional material from collections. Upeneus randalli is distinguished from congeners by the combination of 8 dorsal-fin spines, 14 pectoral-fin rays, 23 to 25 gill rakers, 28 to 30 lateral-line scales, pectoral-fin width 22 to 26 times in SL, and red bars on both caudal-fin lobes. We provide two identification keys: one key is for distinguishing among four species groups and the morphologically unique U. filifer, accounting for all 28 species of the genus, and the other key is for the six closely related species of the “U. tragula group” from the western Indian Ocean, including U. randalli n. sp. Since the colour patterns fade away soon after collection and can be completely lost during preservation, morphometric and meristic characters are important for correctly identifying U. randalli n. sp. and the closely related U. margarethae. The latter species shows considerable variation in body form in different areas of the western Indian Ocean. We discuss the significance of detailed comparative studies of caudal-fin colour patterns and body-form characters for obtaining a deeper understanding of inter- and intraspecific diversity among Upeneus., El número de especies de salmonetes del género Upeneus del Océano Índico Occidental ha aumentado considerablemente debido a los recientes trabajos taxonómicos. En este trabajo se describe una nueva especie, Upeneus randalli n. sp., en base a la coloración del cuerpo y las aletas y a 51 caracteres morfométricos y merísticos de ocho especímenes del golfo Pérsico, apoyados por datos comparativos de una revisión taxonómica de las especies del océano Índico occidental Upeneus y material adicional de colección. Upeneus randalli es differenciada de sus congeners por la combinación de 8 espinas en la aleta dorsal, 14 radios en la aleta pectoral, 23-25 branquiespinas, 28-30 escamas en la línea lateral, anchura de la aleta pectoral 22-26 veces en LS y barras rojas en ambos lóbulos de la aleta caudal. Se proporcionan dos claves de identificación, una para la distinción entre cuatro grupos de especies y la especie U. filifer, morfológicamente única, que responde por el total de las 28 especies del género y la otra clave para las seis especies estrechamente relacionadas del “grupo U. tragula” del océano Índico occidental incluyendo U. randalli n. sp. A causa de que los patrones de color se desvanecen pronto después de la recolección y que pueden perderse completamente durante la conservación, los caracteres morfométricos y merísticos son importantes para la correcta identificación de U. randalli n. sp. y la especie estrechamente relacionada U. margarethae. Esta última muestra una considerable variación en la forma del cuerpo entre diferentes áreas del océano Índico occidental. En este trabajo se discute si la correlación de estudios comparativos detallados de patrones de color de la aleta caudal y forma del cuerpo es significativa para mejorar el conocimiento de la diversidad inter- e intraespecífica entre Upeneus.

Iwatsuki, Yukio, Heemstra, Phillip C. (2011): Polysteganus mascarenensis, a new sparid fish species from Mascarene Islands, Indian Ocean. Zootaxa 3018 (1): 13-20, DOI: 10.11646/zootaxa.3018.1.2, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3018.1.2

Acanthistius joanae new species Figs. 1���5 Acanthistius sp: Heemstra & Randall, 1986: 511, Fig 166.2; Heemstra & Heemstra, 2004. Acanthistius sebastoides: Beckley, 1994: 232, Fig. 109.7; King and Fraser, 2001: 32 & 33. Type material: Holotype: SAIAB 26041, mature female, 113 mm SL; South Africa: off Umdloti, KwaZulu- Natal; depth 80 m; A. R. Thorpe, collector. Paratypes: BPBM 32784, 109 mm SL, South Africa: off KwaZulu- Natal. SAIAB 5766, male, 129 mm SL; South Africa: near Port Alfred, Eastern Cape Province; 22 Feb. 1977; SAIAB 26209, female, 119 mm SL;off Umdloti, SAIAB 28056, 3: 116 ��� 125 mm SL, off Kelso, KwaZulu- Natal, depth 30 m; SAIAB 29406, 66 mm SL; Landers Reef, off Park Rynie, 30 �� 23.2 ' S, 30 �� 50.6 ' E; depth 140 m; 19 August 1991. SAIAB 40259, 90 mm SL; Landers Reef; SAIAB 40459, 2 females: 125 & 130 mm SL; Landers Reef; 30 �� 19 ' S, 30 �� 47 ' E; SAIAB 46440, 3: 116 ���122 mm SL; Aliwal Shoal off Scottburgh, KwaZulu-Natal, 21 June 1994; SAIAB 53810, 118 mm SL, off Umdloti; 74 m, March 1987; SAIAB 59180, 125 mm SL; Aliwal Shoal. SAM 34438, 2: 107 & 130 mm SL, both male; Aliwal Shoal, depth 42 m, 23 June 1994; SAIAB 81415, 113 mm SL; Landers Reef. SAIAB 81416: 126 mm SL; Landers Reef; USNM 283001, 132 mm SL, off Umdloti, USNM 300448, 143 mm SL, off Umdloti, Diagnosis. Interorbital, maxilla, infraorbitals, interopercle, dorsal 4 branchiostegal rays, gular area, isthmus, chest, belly and proximal two-thirds of pectoral fin covered with distinct ctenoid scales. Interorbital area slightly convex. Longest dorsal fin spine equal to or longer than peduncle depth. Caudal, anal, and pelvic fins pale (fin rays mostly reddish orange in life); most prominent dark mark on the head is a large oval, blackish brown blotch covering most of the opercle. Pectoral-fin rays 17���19 Description. (Morphometric data are given in Table 1.) (Data from paratypes are given in parentheses.) Dorsal fin rays XII, 16 (XII, 14 ���16); anal fin rays III, 7 the rays branched; pectoral fin rays 17, 17 (17���19), the upper and lowermost rays unbranched, and the rest branched; pectoral fins reach vertical at anus; pelvic fins also reach anus; branched caudal-fin rays 8 + 7; principal rays 9 + 8; dorsal procurrent caudal rays 4���8, with last 3 segmented; ventral procurrent rays 4���7, with 2 rays segmented; no median fin rays elongated; 4 th to 6 th dorsal-fin spines longest, but slightly shorter than longest dorsal soft-ray or 2 nd anal-fin spine. Body and most of head covered with strongly adherent, distinctly ctenoid scales, extending forward over top of head and interorbital area to a line joining front edge of anterior nostrils; maxilla, infraorbitals, operculum, dorsal branchiostegal rays and gular area with distinct ctenoid scales; snout, preorbitals and lower jaw naked, the skin perforated by numerous minute, close-set, raised pores. Proximal half of all fins are scaly. Lateral-line scales 48 (47���52); scale rows between 6 th dorsal-fin spine and lateral line 16 (13���17). Gill-rakers iii, 3 / 9,v (ii-iv, 2���4 / 8���9, iv-vi; total 19���21); longest gill-raker slightly longer than longest gill-filament. Pseudobranch with 34 (31���35) filaments. Body depth 2.5 (2.4 ���2.6), head length 2.3 (2.2���2.6) in SL. Orbit 4.2 (3.9���4.5) in head length; interorbital flat or slightly convex, width about half orbit diameter, 7.8 (6.8 ���8.0) in head length. Maxilla expanded posteriorly, width including supramaxilla 2.9 (2.4���3.7) times in upper jaw length; posteroventral corner of maxilla with a low knob. Anterior nostrils oval or circular, with a low fleshy rim produced posteriorly as a broad fimbriate flap that can be laid back to cover posterior nostril; posterior nostrils circular, with fleshy fimbriate margin, equidistant from anterior nostril and edge of orbit; infraorbital scales extend forward to a vertical at front edge of eye; about 23 irregular scale series on cheek. Rear edge of opercle with 3 sharp spines, distance from middle spine to lower spine about half distance from middle spine to upper spine; posterior edge of preopercle with 21���23 (14���20) serrae, becoming progressively larger ventrally, 2 (1���3) large, curved, antrorse spines (partly hidden by skin) on lower edge of preopercle. Supratemporal bone rear edge smooth, or with 5 (1���5) small serrae. Premaxilla with a band of small cardiform teeth, the band narrowing posteriorly and widening anteriorly, and an outer series of 17 (16 ���19) short, slightly curved, fixed canines becoming smaller posteriorly. Front half of lower jaw with a band of small, depressible, cardiform teeth, the band narrowing posteriorly, and a few short, stout canines on outer edge of tooth band; 2 or 3 short, stout, curved canines about one-third distance from symphysis to rear end of lower jaw; teeth on rear half of lower jaw are mainly in a single row of small, curved canines directed medially and interspersed with a few much smaller teeth. Two or 3 rows of short, conical teeth in a chevron-shaped band on vomer, and an elongate band of similar teeth on palatines. Second epibranchial with elongate tooth plate. Vertebrae 10 + 16; epipleural ribs on first 9 vertebrae; pleural ribs on vertebrae 3 through 10; last 12���14 dorsal-fin pterygiophores are trisegmental; last 5 anal-fin pterygiophores are trisegmental. Pyloric caeca 4���8. Colour. In life: (125 mm SL, SAIAB 28056) Head and body pale grey, covered with irregular russet spots and blotches; conspicuous, dark, reddish brown blotch covering most of opercle; series of 5 irregular, white blotches along base of dorsal fin and a vertical white streak at base of caudal fin (white markings very noticeable underwater); spinous dorsal fin pale grey, with orange spots on membranes and distal parts of spines; median fin rays orange, inter-radial membranes pale; anal-fin base and rays white, the membranes reddish; pectoral-fin rays orange; first pelvic-fin ray pinkish, remainder white; isthmus, chest and belly white. In alcohol: head and body buff, with faint brown areas and close-set pale spots; dark blotches on head still visible. Comparisons. Acanthistius joanae is most similar to its South African cognate, A. sebastoides. Acanthistius joanae usually has 17 or 18 pectoral fin rays (19-21 in sebastoides). The maxilla of A. sebastoides is apparently naked (scales minute and embedded or absent), but that of A. joanae is covered with distinct, adherent, ctenoid scales that are visible to the naked eye and discernible with the touch of a finger; A. joanae also has conspicuous ctenoid scales on the infraorbitals, interopercles, dorsal 4 branchiostegal rays, gular area, isthmus and chest; but on these areas of A. sebastoides, the scales are absent or minute and embedded. The lateral-line scale tubes of A. joanae have 2 branches, but those of A. sebastoides have 3���5 branches (Figure 5).The apparent rarity of A. joanae may be attributed to its smaller size and its living in deeper water. The specimens from KwaZulu-Natal were caught in depths of 23, 30, 73 and 140 metres; whereas, Acanthistius sebastoides is common from the intertidal region out to depths of 30 m. Etymology. Acanthistius joanae is named to honour Joan Wright, able and conscientious former research assistant of the author. Serranus cuvieri (non Smith): Castelnau, 1861: 2. Serranus sebastoides Castelnau, 1861: 3 (Algoa Bay and Simonstown) Acanthistius sebastoides: Barnard, 1927: Boulenger, 1895: 141; Smith and Smith, 1966: 8; Randall & Heemstra, 1986: 510, Pl. 32, Fig. 166.1; Bianchi, et al. 1999: 165; Heemstra & Heemstra, 2004: 159. Perciform species 1: Brownell, 1979: 24, figs. 162-168. Neotype here designated for 175 mm SL female from Algoa Bay (33 �� 58 ' 54 " S, 25 �� 41 ' 42 " E) SAIAB 85233. Diagnosis. (Characters of the genus given above are not repeated.) Maxilla, infraorbitals, interopercle, branchiostegal rays, gular area, isthmus, chest and proximal two-thirds of pectoral fin naked, or with small, inconspicuous, embedded cycloid scales. Head and body buff, covered with small red spots and a few larger dark brown blotches; most prominent dark mark on the head is a large, oval, blackish brown blotch close behind eye. Caudal and anal fin dark bluish grey; pelvic fins pale yellowish; Pectoral-fin rays 19���21. Description. Morphometric data are given in Table 1; data from the neotype are given first, with data from the other specimens examined in parentheses. Dorsal-fin rays XII, 15 (XI���XIII, 15���17); anal-fin rays III, 7 (2 specimens with 8 anal rays); pectoral-fin rays 19 and 20 (19���21), upper and lowermost rays unbranched and the rest branched; pectoral fins reach vertical at anus; pelvic fins also reach anus; branched caudal-fin rays 8 + 7; principal rays 9 + 8; dorsal procurrent caudal rays 6 (4���8), with last 3 segmented; ventral procurrent rays 5 (4���7), with 2 rays segmented; no median fin rays elongated; 4 th (4 th to 6 th) dorsal-fin spines longest, but slightly shorter than longest dorsal soft-ray or 2 nd anal-fin spine. Body and most of head covered with strongly adherent, distinctly ctenoid scales, extending forward over top of head and interorbital area to a line joining front edge of anterior nostrils; cheek and operculum with ctenoid scales; snout, preorbitals and lower jaw naked, the skin perforated by numerous minute, close-set, raised pores. Proximal two-thirds of all median fins are scaly; pelvic fin rays with scales extending along proximal two-thirds of rays. Lateral-line scales 52 (47���52); lateral-line scale tubes with 3 (3 ��� 5) branches; scale rows between 6 th dorsal-fin spine and lateral line 15 (13���17). Gill-rakers ii, 2 / 8,ii (ii-iv, 2��� 4 / 8���9, ii; total 19 ��� 2); longest gill-raker slightly longer than longest gill-filament. Pseudobranch with 30 (32��� 35) filaments. Body depth 2.5 (2.4���2.6), head length 2.3 (2.2���2.6) in SL. Orbit 3.9 (3.9���4.5) in head length; interorbital flat or slightly convex, width about half orbit diameter, 6.8 ��� 8.0 in head length. Maxilla expanded posteriorly, width 3.1 (2.4���3.7) times in upper jaw length; postero-ventral corner of maxilla with a low knob; supramaxilla well developed, autogenous, hidden by skin and scales. Anterior nostrils oval or circular, with a low fleshy rim produced posteriorly as a broad fimbriate flap reaching to posterior nostril; posterior nostrils circular, with fleshy fimbriate margin, equidistant from anterior nostril and edge of orbit; infraorbital scales extend forward to a vertical at front edge of eye; about 20 irregular scale series on cheek. Rear edge of opercle with 3 sharp spines, distance from middle spine to lower spine equals half distance from middle spine to upper spine; dorsal edge of preopercle with 20 (18���22) serrae, becoming progressively larger ventrally, 2 (1���3) large, exposed, antrorse spines on lower edge of preopercle. Supratemporal bone rear edge smooth, or with 4 (2���4 small serrae. Premaxilla with a band of small, cardiform teeth, the band narrowing posteriorly and widening anteriorly, and an outer series of 4 (3 or 4) larger, slightly curved, fixed canines at symphysis; some inner teeth near symphysis also enlarged. Lower jaw with a band of small, depressible, cardiform teeth, the band narrowing posteriorly, and a few short, stout canines on outer edge of tooth band; 3 (2 or 3) short, stout, curved canines about one-third distance from symphysis to rear end of jaw; teeth on rear half of lower jaw teeth similar but with 3 (3 or 4) stout canines at middle of each side of lower jaw. Vomer with a chevron patch of 3 (3 or 4) rows of short slender teeth and a long band of similar teeth on palatines. Second epibranchial with elongate tooth plate. Vertebrae 10 + 16; 3 supraneural bones: 0/0/0+ 2 / 1 + 1 / 1 / 1 / 1 /; epineural bones on first 9 vertebrae; ribs on vertebrae 3 through 10; last 12 (12���14) dorsal-fin pterygiophores are trisegmental; last 5 anal-fin pterygiophores are trisegmental. Pyloric caeca (7 or 8). Colour. In life: Head, body and dorsal fin buff, covered with small reddish orange spots; caudal fin and rear half of peduncle blackish, the rear edge white; pectoral fins reddish orange distally, the proximal half buff with orange spots; anal fin dusky orange with white edge. In alcohol, head and body buff, covered with small pale spots; and irregular dark brown blotches, the largest and most prominent dark blotch on the head is that immediately behind the eye; nape and top of head blackish, with numerous small pale spots caudal fin and rear part of peduncle dark brownish grey; pelvic and anal fins also dark grey. Attains 28 cm SL. Distribution. Endemic to southern Africa from False Bay, Western Cape Province to Durban. Also reported from Namibia (Bianchi et al. 1999). Discussion: Although the count of 4 anal fin spines for the juvenile reared by Brownell (1979) has not been found in any of the other 60 specimens that have been examined, his specimen is likely to be A. sebastoides. All of the other counts and features (including the colour pattern) agree with the other juveniles of this species that were seen. Acanthistius sebastoides is a sedentary species known from depths of 1��� 30 m. Comparisons. Acanthistius sebastoides was compared with A. joanae in the account above. A. sebastoides differs from both South American species (A. brasilianus and A. patachonicus) in its color pattern, 19���21 pectoral fin rays (15���17 pectoral rays in South American species) and 50���55 lateral-line scales (lateral line scales 56���70 in South American species) The Australian species of Acanthistius (cinctus, ocellatus, pardalotus, paxtoni, and serratus) all differ in color patterns, and various meristic and morphometric characters (See Hutchins, 1981, Hutchins and Kuiter, 1982 and Kuiter, 1994), Remarks. In the original description of Serranus sebastoides, Castelnau (1861, p. 3 mentioned two fish: one from Algoa Bay and one from False Bay [Simons Town] These syntypes are apparently lost. (Bauchot et al. 1984). In view of the considerable similarity between the two South African species of Acanthistius, it will clarify the taxonomic status of these species to designate a neotype for Serranus sebastoides, and this neotype will also provide an objective definition of this species. Acanthistius sebastoides material examined. SAIAB 15280, 9: 55���184 mm SL, Tsitsikamma; SAIAB 78910, 2, 43���188 mm SL, Port Alfred; SAIAB 32150, 3, 122 - 132 mm SL, Port Elizabeth; SAIAB 10953, 163 mm SL, East London; SAIAB 59894, 3,150 ��� 173 mm SL, Pressley���s Bay, Eastern Cape Province; USNM 246689, 173 mm SL, Tsitsikamma; SAM 28499, 24.2 mm SL, specimen reared from egg by Charles Brownell False Bay; Western Cape Province; SAIAB (RUSI) 74-71, 122 mm SL, Algoa Bay; SAIAB (RUSI) 7891, 2, 43��� 188 mm SL, Port Alfred., Published as part of Heemstra, Phillip C., 2010, Taxonomic Review of the perciform fish genus Acanthistius from the east coast of southern Africa, with description of a new species and designation of a neotype for Serranus sebastoides Castelnau, 1861, pp. 59-68 in Zootaxa 2352 on pages 61-67, DOI: 10.5281/zenodo.275626, {"references":["Heemstra, P. C. & Randall, J. E. (1986) Family No. 166: Serranidae, in Smiths' Sea Fishes (M. M. Smith and P. C. Heemstra (editors) Macmillan South Africa, Johannesburg, 1047 pp.","Heemstra, P. C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. South African Institute for Aquatic Biodiversity. Grahamstown, 488 pp.","Beckley, L. E., in Branch, G. M., Griffiths, C. L., Branch, M. L. & Beckley, L. E. (1994) Fish, (pages 211 - 285), in Two Oceans, A Guide to the Marine Life of South Africa. David Philip Publishers, Cape Town 360 pp.","King, D. & Fraser, V. (2001) More Reef Fishes & Nudibranchs. Struik Publishers, Cape Town, 136 pp.","Castelnau, F. L. de. (1861) Memoire sur les Poissons de l'Afrique australe, J. B. Bailliere et Fils. Paris, vii + 78 pp.","Barnard, K. H. (1927) A monograph of the marine fishes of South Africa. Part II (Teleostei - Discocephali to end. Appendix). Annals South African Museum, v. 21, (pt 2), 419 - 1065, pls 18 - 37.","Boulenger, G. A. (1895) Catalogue of the perciform fishes in the British Museum, volume 1, 2 nd ed. London, xix + 394 pp.","Smith, J. L. B. & Smith, M. M. (1966) Fishes of the Tsitsikama Coastal National Park. National Parks Board of Trustees of the Republic of South Africa. 161 pp.","Bianchi, G., Carpenter, K. E., Roux, J. P., Molloy, F. J., Boyer, D. & Boyer, H. J. (1999) FAO Species Identification Guide for Fishery Purposes. Field Guide to the Living Marine Resources of Namibia. 265 pages. Rome, FAO.","Brownell, C. L. (1979) Stages in the early development of 40 marine fish species with pelagic eggs from the Cape of Good Hope. J. L. B. Smith Institute of Ichthyology Bulletin, Rhodes University. 40, 1 - 84.","Kuiter, R. H. (1994) Family Serranidae, in Fishes of Australia's South Coast. (M. F. Gomon, J. M. C. Glover & R. H. Kuiter, (editors) State Print, Adelaide, 992 pp.","Bauchot, M. L., Desoutter, M. & Randall, J. E. (1984) Catalogue critique des trypes de Poissons du Museum national d'Histoire naturelle. (Famille des Serranidae). Bulletin Museum national Histoire naturelle, Paris 4 e serie, 6, section A, no. 3, supplement, 3 - 82."]}

Acanthistius Gill, 1862: 236 Type-species, Plectropoma serratum Cuvier, 1828, by monotypy). Diagnosis. Body robust; scales small, ctenoid, those on the head, chest and belly ctenoid or mostly embedded; lateral line complete, tubed scales 48 ��� 67, not counting several on proximal half of caudal fin, the tubes with 2���5 branches. The scales of Acanthistius ocellatus and A. serratus were described as ���cycloid��� (Kuiter, 1994), but the lateral body scales of the juvenile of each species that I examined are ctenoid, although the ctenii are very small and mostly covered by skin. Dorsal fin single, with 11���13 spines and 13���18 segmented, branched rays; the fin margin slightly notched before the soft-rayed part; anal fin with 3 spines and 7��� 9 segmented, branched rays; caudal fin truncate or convex, with 15 branched, segmented rays; no filamentous or elongated median-fin rays; pectoral fins rounded, with 15���21 rays, most of which are branched and fleshy. Mouth terminal, jaws equal; premaxillae slightly protrusile, maxilla reaches to below rear half of eye; supramaxilla autogenous, well developed, its length about half length of maxilla, hidden by skin on upper rear edge of maxilla. Jaws with inner row or rows of small slender teeth and a few outer teeth enlarged as small canines; vomer and palatines with cardiform teeth; tongue edentate. Opercle with 3 sharp spines posteriorly, but upper and lower spines are mostly hidden by skin and scales; preopercle serrate, with 3 or 4 serrae at angle enlarged and 1���3 large, antrorse spines on ventral edge. Branchiostegal rays 7, the membranes connected to anterior end of isthmus. Lower limb of first gill arch with 13���16 gill-rakers, of which 4���8 are rudiments. Oblong tooth plate on 2 nd epibranchial. Anterior nostrils with a fleshy rim, the rear margin enlarged, forming a large flap reaching to or over posterior nostril. Vertebrae 26 (10 abdominal + 16 caudal or 11 abdominal + 15 caudal); 3 supraneural bones: 0/0/0+ 2 / 1 + 1 / 1 / 1 / 1 or 0+0/0/ 2 / 1 + 1 / 1 / 1 / 1 /. Remarks. Plectropoma aculeatum Valenciennes, 1830, was described from a specimen said to have been collected from the "Banc des Aiguilles" [= Agulhas Bank off the south coast of South Africa]. But, according to Boulenger (1895) who examined the holotypes, P. aculeatum is a synonym of Acanthistius brasilianus (Cuvier, 1828), which is endemic to the coast of Brazil (Irigoyen et al., 2008); and the type locality for P. aculeatum is surely in error (Bauchot et al., 1984: 20). The genus Acanthistius is not clearly defined, and its inclusion within the family Serranidae has recently been questioned (Craig and Hastings, 2007). The genus comprises ten species, occurring in shallow, warmtemperate waters of the Southern Hemisphere: Acanthistius brasilianus (Cuvier), south coast of Brazil; A. cinctus (G��nther), Australia, New Zealand, Lord Howe Id, Norfolk Id, and Kermadec Islands; A. fuscus Regan, Easter Id and Sala y G��mez Id; A. ocellatus ( G��nther), Australia; A. pardalotus Hutchins, Western Australia; A. patachonicus (Jenyns), Argentina and southern Brazil; A. paxtoni Hutchins & Kuiter, Australia; A. pictus Tschudi, Ecuador, Peru and Chile; A. sebastoides (Castelnau), Namibia and southern Africa; and A. serratus (Cuvier), Australia. Species of Acanthistius are superficially similar to those of the genus Hypoplectrodes Gill, 1862, which includes the genus Ellerkeldia Whitley, 1927 (Anderson and Heemstra, 1989); species of both genera have a truncate or convex caudal fin, branched pectoral-fin rays, with the middle rays thickened, and 1���3 strong antrorse spines on the ventral edge of the preopercle. Acanthistius species differ in having 11���13 dorsal-fin spines, versus 10 dorsal-fin spines in all Hypoplectrodes species (except H. wilsoni Allen & Moyer (1980) which has 10 or 11 dorsal-fin spines. Kuiter (2004) added the two South African species of Acanthistius, A. joanae and A. sebastoides to the genus Trachypoma, but the type, T. macracanthum, has a smaller head, with dorsal profile slightly concave, eye diameter 2 times interorbital width, and the interorbital area with a median groove. The first fish to be described from South Africa that may have been a species of Acanthistius was named Serranus cuvierii by Andrew Smith (1831). Smith���s description was published in the South African Quarterly Journal, which even then, must have been an obscure serial. Smith���s description was based on a fish from Algoa Bay and includes the barely legible fin counts of D, 13 / 21 and A, 0/ 14 [dorsal fin with 13 spines, 8 softrays, anal fin with no spinous rays and 14 soft rays]. These fin ray counts do not agree with the dorsal-fin ray count of 12 or 13 spines, plus 15 or 16 rays, and the anal-fin ray count of 3 spines, 7 or 8 rays for A. sebastoides, or the fin counts of any other serranid fish of South Africa. Smith���s description of the colour for his Serranus cuvieri agrees in some respects and disagrees in others with the colour patterns of A. sebastoides. In view of these discrepancies, Serranus cuvierii Smith, 1831, is considered a nomen dubium, as the description does not match any known fish species from South Africa., Published as part of Heemstra, Phillip C., 2010, Taxonomic Review of the perciform fish genus Acanthistius from the east coast of southern Africa, with description of a new species and designation of a neotype for Serranus sebastoides Castelnau, 1861, pp. 59-68 in Zootaxa 2352 on pages 60-61, DOI: 10.5281/zenodo.275626, {"references":["Gill, T. N. (1862) Remarks on the relations of the genera and other groups of Cuban fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 235 - 242.","Cuvier, G. & Valenciennes, A. (1828) Histoire Naturelle des Poissons, 2 (3), [Cuvier author of pp 262 - 386.]","Kuiter, R. H. (1994) Family Serranidae, in Fishes of Australia's South Coast. (M. F. Gomon, J. M. C. Glover & R. H. Kuiter, (editors) State Print, Adelaide, 992 pp.","Boulenger, G. A. (1895) Catalogue of the perciform fishes in the British Museum, volume 1, 2 nd ed. London, xix + 394 pp.","Irigoyen, A. J., Gerhardinger, L. C. & Carvalho-Filho, A. (2008) On the status of the species of Acanthistius (Gill, 1862) (Percoidei) in the South-West Atlantic Ocean. Zootaxa, 1813, 51 - 59.","Bauchot, M. L., Desoutter, M. & Randall, J. E. (1984) Catalogue critique des trypes de Poissons du Museum national d'Histoire naturelle. (Famille des Serranidae). Bulletin Museum national Histoire naturelle, Paris 4 e serie, 6, section A, no. 3, supplement, 3 - 82.","Craig, M. T. & Hastings, P. A. (2007) A molecular phylogeny of the groupers of the subfamily Epinephelinae (Serranidae) with a revised classification of the Epinephelini. Ichthyological Research (2007) 54, 1 - 17.","Whitley, G. P. (1927) Studies in ichthyology. No. 1. Records of the Australian Museum, 15, 289 - 304.","Anderson, W. D. Jr & Heemstra, P. C. (1989) Ellerkeldia, a junior synonym of Hypoplectrodes, with redescriptions of the type species of the genera (Pisces: Serranidae: Anthiinae). Proceedings of the Biological Society of Washington, 102 (4), 1001 - 1017.","Allen, G. R. & Moyer, J. T. (1980) Ellerkeldia wilsoni, a new species of serranid fish from southwestern Australia. Japanese Journal of Ichthyology, 26, 329 - 333.","Kuiter, R. H. (2004) Basses Hamlets and Their Relatives a Comprehensive guide to selected Serranidae and Plesiopidae. TMC Publishing. Chorleywood, UK.","Smith, A. (1831) Dr Smith's contributions to the natural history of South Africa. South African Quarterly Journal ... 17 - 21... [pagination uncertain]."]}