Special Publication J.L.B. Smith Institute of Ichthyology ISSN: 0075-2088 Num. 63, 1999, pp. 1-8

Special Publication, No. 63, pp. 1-13

Code Number: fs99001

ABSTRACT

Three species of the Apogon bandanensis group are found in the Red Sea, the wide-spread Apogon guamensis, the endemic Apogon annularis, and a new endemic species Apogon zebrinus. Color patterns, gill-rakers, body depth, second anal spine length, pectoral fin length, and caudal peduncle depth are important aids for identification of the Red Sea species. Two synonyms of Apogon guamensis were based on juvenile material: Apogon ocellatus from Madagascar and Apogon spongicolus from the Red Sea. Rüppell's, Günther's and Klausewitz's concepts of Apogon annularis are reviewed. Apogon erdmani is a synonym of Apogon annularis. Apogon savayensis and another wide spread new species were not found in any Red Sea collections we examined.

INTRODUCTION

Rüppell described Apogon annularis from the Red Sea in 1824 followed at a long interval by Apogon erdmani Lachner, 1954 and Apogon spongicolus Smith, 1964, both from the Red Sea. One other described species in this complex, Apogon guamensis Valenciennes, 1832, occurs in the Red Sea. Material of two other Indian Ocean species, Apogon savayensis Günther, 1871, and a new species (not known west of the Maldive Islands) have not been collected in the Red Sea despite the attention this area has received by ichthyologists. The last revision of the "bandanensis" group of Apogon was done by Lachner in 1954. He recognized four species: Apogon bandanensis, Apogon savayensis, Apogon nubilus [=Apogon guamensis] and Apogon erdmani [=Apogon annularis]. The senior author and E. A. Lachner,began working on this group in 1974, at the U.S. National Museum after an undescribed species occurring in the Red Sea was identified. J.E. Randall joined the study in 1987 as the result of more material he collected and photographed in the Red Sea. A complete revision of this group is being prepared separately by the first author. The publication of this species is based on the need to make this name available, given other pending regional works for the Red Sea.

METHODS AND MATERIALS

Our methods of taking and recording meristic data and measurements are given in Fraser and Lachner (1985). Data referring to non-type specimens have been abbreviated and include the catalog number, total number of specimens (size range), pertinent locality and only the important published station numbers or specimen numbers. All of the localities for the new species are reported under "Material Examined'' are plotted on a map. The comparative material include western Indian Ocean material from the revision manuscript. The symbols on the maps may represent one or more collections taken at different times and one or more specimens. Oral incubation was checked by examining the mouth cavity of larger specimens with buccal enlargement. Where we found eggs in the mouth, the size of specimens is listed under the section "Remarks."

The following acronyms are used to designate institutions and collections cited: ANSP Academy of Natural Sciences, Philadelphia; BPBM Bernice P. Bishop Museum, Honolulu; FMNH Field Museum of Natural History, Chicago; RMNH National Natuurhistorische Museum, Leiden; RUSI J.L.B. Smith Institute of Ichthyology, Grahamstown, South Africa; USNM collections of the former United States National Museum, deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. UTAI Tel Aviv University, Israel; HUJ Hebrew University, Jerusalem; SMF Natur-Museum und Forshungs-Institut Senckenberg, Frankfurt-am-Main.

Apogon zebrinus new species
Figures 1-5, Tables 1-2

HOLOTYPE; USNM 213422; (59.6); N. Of Râs Burqa, N.W. coast of Aqaba, Egypt; 23 July 1969; to 10 m.; Victor G. Springer; VGS 69-7; x-rayed.

PARATYPES: Red Sea: Egypt: BPBM 31807 was USNM 213424; 4(57-71); Gulf of Aqaba, Râs Muhammad; 0 to 10m.; 26 Sep. 1969; VGS 69-28. USNM 341635; 4(49-73); data same as holotype. TAU 9672; (67); Gulf of Aqaba; UTAI-NS 4181; 8 Oct. 1968. Sudan: BPBM 19741; (38-44); Suakin Harbor, 11 Oct. 1974. BPBM 27417; (41);color; just N. of Port Sudan, 9 Jan. 1980. Eritrea: USNM 213423 (47); N. End of Isola Delemme, off Massawa; 7 Aug. 1969; VGS 69-9. Saudi Arabia: BPBM 30388; 4,(58-74); 29 km. S. of Yanbu, 30 May 1984. Yemen: USNM 213421 15(55-63); Gold Mohur Bay; 21 Dec. 1964; IIOE Cr. 9; F. Talbot Sta. 206.

Other material: Gulf of Aqaba: TAU 9671; (27); UTAI-NS 1805; 13 Sep. 1967. HUJ 12007; 3(17-26); Elat; HUJ 11984; 2(16-25); Elat; HUJ 12859; (20); Dahab.

DIAGNOSIS. An Apogon of the subgenus Ostorhinchus; vertical bars on body, basicaudal mark below lateral-line scales in adults, caudal fin with dark margins, subocular mark broad, triangular, 13 pectoral fin rays, total gillrakers usually 25-28 body depth 40-47%, caudal peduncle depth 18-22%, second anal spine length 19-23%, and pectoral fin length 26-32% of standard length.

Figure 1. Apogon zebrinus, underwater photography by Rudie Kuiter in Egypt.

DESCRIPTION. For general body shape see Figures 1-2. Range of proportions (as percentage of standard length with the holotype in parentheses): greatest body depth 40-47 (44.8); head length 43-52 (45.8); eye diameter 14-18 (16.9); snout length 8-11 (8.5); bony interorbital width 10-14 (12.1); upper jaw length 20-25 (21.8); caudal peduncle depth 18-22 (19.6); caudal peduncle length 18-24 (21.5); first dorsal-fin spine length 2.5-4.1 (2.5); second dorsal-fin spine length 10-14 (11.2); third dorsal-fin spine length 18-23 (21.1); fourth dorsal-fin spine length 19-24 (20.3); second dorsal fin spine 17-21 (17.6); first anal-fin spine length 2.3-7.1 (3.7); second anal-fin spine length 19-23 (19.6); pectoral fin length 26-32 (-); pelvic fin length 24-31 (-).

Dorsal fin VII+I,9; anal fin II,8; pectoral fin usually 13-13, rarely 12-13; pelvic fin I,5; principal caudal rays 9 + 8; pored lateral line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 6; median predorsal scales 3; circumpeduncular scale rows 12 (5+2+5); total gillrakers 26-28 (holotype 27) well developed 24-27 (holotype 25) well developed, (1-3+5-7 upper, 17-20+0-1 lower) . The frequency of certain gill raker counts is given in Table 1.

Figure 2. Holotype of Apogon zebrinus, USMN 213422, 59.6mm SL, N. Ras Burqa, NW Egypt, Coast of Gulf of Aqaba.

Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and vomer; none on ectopterygoid, entopteygoid or basihyal. Vertebrae 10 + 14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural. Three predorsals, one spine on first dorsal pterygiophore. Basisphenoid present. Supramaxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal margins. Infraorbital shelf present on third bone. Scales ctenoid. Pored lateral-line scales complete.

LIFE COLORS. From a 120 mm slide of BPBM 27417: Body brownish gray dorsally, shading to bluish gray on sides and silvery white ventrally, with seven blackish bars two to three times broader than pale interspaces, the first extending ventrally from nape and the last anteriorly on caudal peduncle; a saddle-like black bar posteriorly on caudal peduncle extending to lateral line and tapering below; head grayish brown dorsally, shading to silvery white on side and ventrally (with a lavender-pink cast); an oblique wedge-shaped black mark extending from posteroventral corner of eye nearly to corner of preopercle; an indistinct brown and blackish blotch on opercle; first dorsal fin dusky translucent with a black leading edge from origin of fin to tip of fifth dorsal spine; second dorsal fin dusky translucent, the rays faintly pink, with a blackish leading edge covering spine, first ray, and membrane between; remaining fins translucent with faint pink rays, the caudal with a dusky margin, and the anal with a little dusky pigment distally on the first two soft rays; iris blackish brown. Two transparencies by Rudie H. Kuiter were similar in color except that the caudal margins were brownish to blackish.

Figure 3. Body depth, caudal peduncle depth and caudal peduncle length % SL for Apogon zebrinus (cylinder), Apogon annularis (cube), Apogon guamensis (pyramid).

PRESERVED COLOR PATTERN. Adults: wide, triangular subocular cheek mark behind upper jaw; first dorsal fin with dark membrane between spines 1-4; edges of caudal lobes dark; caudal peduncle with a nearly complete dark wide bar, connected dorsally and almost connected ventrally; pale, slightly wavy bars present on body from below the first dorsal fin and behind the pectoral fin to just behind the ends of the second dorsal fin base and anal fin base, bars extend above the lateral-line scales starting about the second dorsal fin; no dark saddle under the first or second dorsal fin; stomach and intestine black. Juveniles: similar to the adults, except with a complete, dark caudal peduncle band.

DISTRIBUTION. Known only from the Red Sea and the Gulf of Aden.

ETYMOLOGY: from zebra the Abyssinian (Amheric) name for the striped equine of Africa and -inus a Latin suffix pertaining to. The name refers to the somewhat variable broad dark and light bars on the body. A noun in apposition.

REMARKS. This species has been collected at the same stations with Apogon annularis (BPBM 20373, 30386, USNM 212431)) in the Red Sea and Gulf of Aden (USNM 212447) and with Apogon guamensis (USNM 212882). Rather bold pale bars on the body and a caudal peduncle mark is not quite complete along the anterior part of the band will identify the new species from Apogon annularis. A comparison of the preserved color pattern for the three Red Sea species is presented in Table 2.

Apogon zebrinus has a body depth, pectoral fin length and second anal spine length similar to Apogon annularis (Figures 2 and 3). Caudal peduncle depth is slightly greater in Apogon zebrinus than in Apogon annularis. The gill raker counts show, at best, slight modal or average differences, particularly the lower gill arch (Table 1). Apogon guamensis can be distinguished from both of these species by the narrow, oblique cheek mark below the eye, a more slender body, shorter pectoral fin length, well-developed caudal spot in juvenile, diffuse in adults, and no dark margins to the caudal fin edges and slightly lower gillraker counts. Color patterns appear to be one of the important distinguishing characters in this complex, particularly changes in the caudal peduncle and cheek marks, development of pale bars on the side of the body, and development of colored edges to the caudal fin.

Indian Ocean Apogon savayensis may be distinguished from Apogon zebrinus by the lack of well developed bars on the side, an incomplete caudal mark mostly above the lateral line (see Winterbottom et al. 1989, fig. 164) and a shorter second anal spine (14-20% of SL). An undescribed species occuring in the Indian Ocean as far west as the Maldives has numerous, narrow bars (see Jordan and Seale ,1906:240, fig. 33), an incomplete caudal mark mostly above the lateral line and slightly higher gillraker counts.

The largest specimen was 74 mm SL. None of the material indicated oral brooding of the eggs. We believe that this species is an oral brooder based on positive evidence from specimens of Apogon guamensis and Apogon annularis.

Apogon annularis Rüppell, 1828 (Fig 4)
Apogon annularis Rüppell, 1828: 48 (Gulf of Suez)
Apogon erdmani Lachner, 1951: 595, pl.18 (Fig A). ( Red Sea, Saudi Arabia, Jidda)

DIAGNOSIS. An Apogon of the subgenus Ostorhinchus; no bars on body, circumpeduncular basicaudal mark in juveniles and adults, caudal fin with dark margins, subocular mark broad, triangular, 13 pectoral fin rays, total gillrakers usually 26-29, body depth 41-50%, caudal peduncle depth 15-20%, second anal spine length 18-22%, and pectoral fin length 26-31% of standard length.

Figure 4. Apogon annularis, sudan, underwater photograph by John E. Randall.

DESCRIPTION. For general body shape see Figure 4. Range of proportions (as percentages of standard lengths): greatest body depth 41-50; head length 41-46; eye length 16-18; snout length 8-10; bony interorbital width 10-11; upper jaw length 21; caudal peduncle depth 15-19; caudal peduncle length 19-23; first dorsal-fin spine length 2.5-5.3; second dorsal-fin spine length 9-12; third dorsal-fin spine length 17-21; fourth dorsal-fin spine length 18-21; second dorsal spine 17-20; first anal-fin spine length 3.4-5.4; second anal-fin spine length 18-22; pectoral fin length 26-31; pelvic fin length 25-28.

Dorsal fin VII-I,9; anal fin II,8; pectoral fin usually 13-13, rarely 12-13 or 14-13; pelvic fin I,5; principal caudal rays 9 + 8; pored lateral line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 6; median predorsal scales 3; circumpeduncular scale rows 12 (5 +2+5); total gillrakers 25-30, usually 26-29 well developed, (0-3+5-6 upper, 19-21+0-2 lower) . The frequency of certain gill raker counts is given in Table 1.

Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and vomer; none on ectopterygoid, entopteygoid or basihyal. Vertebrae 10 + 14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural. Three predorsals, one spine on first dorsal pterygiophore. Basisphenoid present. Supramaxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal margins. Infraorbital shelf present on third bone. Scales ctenoid. Pored lateral-line scales complete.

LIFE COLOR PATTERNS. From an underwater 35mm Kodachrome transparency of a specimen from Sudan: body light bluish on the body to about the midline, pinkish blue below midline; caudal peduncle with a complete black band within whitish area on each side; broad brownish triangular cheek mark from eye to edge of preopercular ridge; anterior edges of the dorsal fins, caudal fin, anal fin and pelvic fins bluish white, rest of fins pale, pectoral fins pale; no other marks on head, body or fins; iris blackish brown.

PRESERVED COLOR PATTERN. Adults: wide, triangular subocular cheek mark behind upper jaw; first dorsal fin dark membranes between spines 1-4; edges of caudal lobes dark; caudal peduncle with a complete dark wide bar, connected dorsally and ventrally, body uniform without pale bars; no dark saddle under the first or second dorsal fin; stomach and intestine black. Juveniles: similar to the adults.

DISTRIBUTION. Known only from the Red Sea and the Gulf of Aden.

REMARKS. See species comparison discussion under remarks about Apogon zebrinus. Günther (1859) use the combination Apogon annularis var. roseipinnis. Klausewitz (1959) suggested that Apogon annularis is a subspecies of Apogon aureus. These subspecies concepts have not been supported by any subsequent work, but do suggest part of the identification problems with trying to relate appropriate names in these two species groups with caudal peduncle banding.

Apogon annularis was described by Rüppell in 1828 from the Red Sea. Rüppell directly related his specimens to the figure of Ostorhinchus fleurieu in Lacépède, 1801. He noted that the description in Lacépède's text was another kind of fish. In 1835-38, Rüppell commented that Cuvier's identification as Apogon rex mullorum (= Apogon imberbis) was not correct, and again related Apogon annularis with Lacépède's figure. However, a first revisor status of Ostorhinchus fleurieu may be attributed to Gon (1987) by virtue of having created a neotype in 1987 for another species. We recognize Apogon annularis as a valid name and as a Red Sea endemic consistent with the majority of previous studies. We agree with Klausewitz (1959) that Apogon erdmani is a synonym of Apogon annularis.

Two males in USNM 212429 exhibited buccal enlargement but no eggs were found in the mouths. This species has been collected at the same stations with Apogon zebrinus in the Red Sea and Gulf of Aden.

MATERIAL EXAMINED. Types: Apogon annularis Lectotype; SMF 1774 (60.9); Red Sea, Et Tur, Sinai coast, Egypt, Gulf of Suez.; E. Rüppell, 1827. Apogon erdmani Holotype; USNM 147518 (59.5); Saudi Arabia, Jidda, Sams Pier, 2 July 1948, D.S. Erdman Coll; x-rayed.

COMPARATIVE MATERIAL. Apogon annularis: Red Sea: Egypt: Gulf of Aqaba: USNM 212429; 24(39-53); Bay at El Himeira; 16 July 1969; VGS 69-2. BPBM 21513; 3(34-45); color; El Himeira; 12m.; 25 Apr. 1977. USNM 212430; 16(36-59); N. of Râs Burqa; 23 July 1969; VGS 69-7. USNM 212432; 5(40-52); Marsa Muqabila; 17 July 1969; VGS 69-3. USNM 212438; 8(42-50); One mi. N. of Râs Burqa; 21 July 1969; VGS 69-6.USNM 212441; (40); Between Marset Mahashel Ala & Marset Abu Samra; 2 Sept. 1969; VGS 69-18.USNM 212443; 3(14-36; Bay between marsa Mokrakh and El Himeira; 15 July 1969; VGS 69-1. Strait of Gûbâl: USNM 212428 61(45-59); Râs Muhammad; 26 Sept. 1969; VGS 69-28. USNM 212439; (31,50); 2 Jan. 1965; IIOE - H.A. Fehlmann - 30. USNM 212442; 21(25-52); 7 Jan. 1965; IIOE - H.A. Fehhlmann - 36. USNM 212444; (26); 6 Jan. 1965; IIOE - H.A. Fehlmann - 35. Gulf of Suez: USNM 212436; 9(26-30); 4 Jan. 1950; IIOE - H.A. Fehlmann - 32. USNM 212437; 30(32-55); 30 Dec. 1964; IIOE - H.A. Fehlmann - 26.USNM 212440; 3(28-57); El-Tur, Sinai Peninsula; 27 Sept. 1969; VGS 69-29. USNM 212446; 72(24-58); (SOSC Ref. No. 177); 10 Jan. 1965; IIOE - H.A. Fehlmann - 38. Saudi Arabia: ANSP 163226; (52-53); Khor Obhour; SV-RS-77; 15 Apr 1977. RMNH 12892; (61-66); Jedda;1880; J. A. Vragt. BPBM 30386; 4(14-18); 29km. S. of Yanbu; 30 May 1984. Sudan: BPBM 20373; 3(34-45); Suakin Harbor; 11 Oct. 1974. Eritrea: USNM 212431; 29(27-55); North end of Isola Delemme, off Massawa,; 7 Aug. 1969; VGS 69-9. USNM 212433; 47(24-53); Massawa; 12 Aug. 1969; VGS 69-12. USNM 212434; 7(17-29); Difnein I., South shore; 15 Aug., 1969; VGS 69-15. USNM 212435; (31, 35); Melita Bay; 13 Aug. 1969; VGS 69-13. USNM 212445; (28); Ethiopia; Aug. 1969; VGS 69-14. Yemen: Red Sea: BPBM 35701; 4(18-22); Jaz 'ir as Zubayr, 15° 6.8' N., 42° 35.9' E.; 15 May 1993. Gulf of Aden: USNM 212447; 20(20-62); Gold Mohur Bay,; 21 Dec. 1964; IIOE Cr. 9, F. Talbot - 26.

Apogon guamensis Valenciennes, 1832 (Fig 5)
Apogon guamensis Valenciennes, 1832:54 (Guam, Mariana Ids.).
Apogon nubilus Garman, 1903, 229, pl.1, fig 1 (Fiji, Suva reef).
Apogon ocellatus Fourmanoir and Crosnier, 1964: 5, fig 3
( Madagascar I., Nosy-Bè, Ambatoloaka).
Apogon spongicolus Smith, 1964: 529, fig 1. (Red Sea; Ethiopia).

DIAGONSIS. An Apogon of the subgenus Ostorhinchus; no bars on body, basicaudal spot diffuse in adults, caudal fin without dark margins, subocular mark narrow, linear, usually 13 pectoral fin rays, total gillrakers usually 24-28 (25-28 in the Indian Ocean and 24-27 in the Pacific Ocean), body depth 37-42%, caudal peduncle depth 15-18%, second anal spine length 15-17%, and pectoral fin length 24-26% of standard length.

Figure 5. Apogon guamensis, Egypt, underwater photograph by Rudie Kuiter.

DESCRIPTION. For general body shape see Figure 5. Range of proportions (as % of SL): greatest body depth 37-42; head length 40-44; eye length 14-18; snout length 8-9; bony interorbital width 10-12; upper jaw length 20-22; caudal peduncle depth 15-18; caudal peduncle length 22-26; first dorsal-fin spine length 2.1-4.2; second dorsal-fin spine length 8-11; third dorsal-fin spine length 17-22; fourth dorsal-fin spine length 16-19; second dorsal spine 15-19; first anal-fin spine length 2.6-5.1; second anal-fin spine length 15-17; pectoral fin length 24-26; pelvic fin length 21-25.

Dorsal fin VII-I,9; anal fin II,8; pectoral fin usually 13-13, rarely 14-13; pelvic fin I,5; principal caudal rays 9 + 8; pored lateral line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 5-6; median predorsal scales 3-4; circumpeduncular scale rows 12 (5 +2+5); total gillrakers 23-29, usually 25-28 well developed, (1-3+4-6 upper, 17-20+0-1 lower) . The frequency of certain gill raker counts is given in Table 1.

Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and vomer; none on ectopterygoid, entopteygoid or basihyal. Vertebrae 10 + 14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural. Three predorsals, one spine on first dorsal pterygiophore. Basisphenoid present. Supramaxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal margins. Infraorbital shelf present on third bone. Scales ctenoid.

LIFE COLOR PATTERN. From two transparencies of Red Sea specimens by Rudie H. Kuiter: Body more or less uniform bluish brown, darker dorsally under the dorsal fins, may have 3-4 poorly developed lighter vertical bars on side behind pectoral fin, a diffuse darker, offset basicaudal spot; a narrow cheek mark from below eye to edge of preopercular ridge; dark first dorsal fin from first through fourth and upper half of membrane between fourth and fifth, rest of fin pale and translucent; second dorsal fin, anal fin, pelvic and pectoral fins all pale and translucent; caudal fin without any marks on the dorsal and ventral edges, fin pale and translucent; iris blackish brown.

PRESERVED COLOR PATTERN. Adults: narrow, linear subocular cheek mark behind upper jaw; first dorsal fin with dark membrane between spines 1-4; edges of caudal lobes plain; caudal peduncle with a diffuse spot of variable intensity; pale bars, usually four or fewer, occasionally present on body from below the first dorsal fin and behind the pectoral fin to just behind the ends of the second dorsal fin base and anal fin base, bars extend above the lateral-line scales starting about the second dorsal fin; no dark saddle under the first or second dorsal fin; stomach and intestine black. Juveniles: similar to the adults.

DISTRIBUTION. Known from the Red Sea, Indian Ocean and West Pacific to the Tonga Islands, Samoa, Phoenix and Marshall Islands.

REMARKS. See species comparison discussion under remarks about Apogon zebrinus. A number of specimens were found with buccal enlargement, but no eggs were present (USNM 112046 and 147524).

Apogon ocellatus and Apogon spongicolus are juvenile specimens of Apogon guamensis. Figure 5, an adult Red Sea Apogon guamensis, shows the faint basicaudal spot and cheek mark and basicaudal spot. Smith's (1964) material lacks the cheek mark, which is present in Fourmanoir and Crosnier's material. Smith compared Apogon spongicolus with Apogon nubilus [=guamensis] and believed the specimens to be new because of the missing cheek mark. The largest paratype of Apogon spongicolus, RUSI 774, from Kenya is Apogon fleurieu based on the pectoral ray count (14-14) and a low gill total raker count of 23 (2+4-15+2). See Randall et al., (1990) figure 6 for the intense basicaudal spot and lack of a cheek mark in a 33 mm Red Sea specimen. The holotype and other paratypes of Apogon spongicolus have 13 pectoral rays and 17 well developed lower arch gill rakers. Both character states, individually, are rare for Apogon fleurieu. These smaller specimens, 15.7-17.0 mm SL better represent material of Apogon guamensis even though the cheek mark is not present.

Fourmanior and Crosnier (1965) gave a very brief color description of Apogon ocellatus without any other information. In addition to the diagnosis color pattern, the syntypes have pectoral fin counts of 13-13 and total gillraker counts of 24 and 26, all consistent with Apogon guamensis. Their name is a primary homonym of Apogon ocellatus Weber, 1911 and a secondary homonym of Apogon ocellatus (von Bonde, 1923).

MATERIAL EXAMINED. Types: Apogon guamensis Syntypes; MNHN 8767 4(22-56); Guam; Quoy and Gaimard; 1826-1829. Apogon nubilus Holotype; MCZ 28315 (54); Fiji, Suva reef; 12 Dec. 1897. Apogon ocellatus Syntypes; MNHN 1973-41 2(20-20); Madagascar I., Nosy-Bè, Ambatoloaka; x-rayed. Apogon spongicolus Holotype; RUSI 354 (17); Red Sea; Ethiopia; x-rayed. Paratypes: RUSI 441; 2(15.7); Red Sea; Ethiopia; x-rayed. RUSI 774; (22,1);Kenya, Shimoni; Nov.1952; x-rayed.

COMPARATIVE MATERIAL. Apogon guamensis: Red Sea: Israel: Gulf of Aqaba. RUSI 3278 (57); x-rayed; USNM 212876; 5(64-75); USNM 212875; 5(47-63); USNM 212874; 3(52-64); USNM 212878; 7(59-71); USNM 191707; 4(21-28); Elat; USNM 212873; (67). Egypt: USNM 212877; 12(16-66) Muqabila. USNM 212883; 10(39-69); Strait of Jubal, 0-5m. USMN 212884; (31);Strait of Jubal, S. end of Sinai Peninsula. BPBM 18346; (33); Râs Muhammed; 19 Sep. 1974. USNM 213617; 57(32-71); 27° 18' 50"N, 033° 47' 35"E; 0-5m. USNM 212879; 6(23-66). USNM 212880; 46(33-68). USMN 212884; (31);Harghada; USNM 212885; (32); Ghardaqe; USNM 147519; 28(53-68); NW edge Shaib Al Zanadir Reef. Saudi Arabia: Jidda. USNM 191658; 3(48-55); USNM 147523; 15(12-66); USNM 112046; 12(33-65); USNM 212886; 3(40-65); USNM 163808; (11); USNM 147524; 7(37-60). Ethiopia: Sharm Ubhar. USNM 212881; 9(49-60); N. end of Isola Delemme. USNM 212882; (49); naval base, Massawa.

ACKNOWLEDGMENTS

For the loan of material and the use of museum facilities we extend our thanks to many of our colleagues over the past two decades. We especially recognize the following individuals: William G. Saul, Eugenia and James E. Böhlke, William F. Smith-Vaniz (ANSP), Karel F. Liem and M.M. Dick (MCZ), David Catania, William N. Eschmeyer and Tomio Iwamoto (CAS), Robert K. Johnson formerly (FMNH), Arnold Y. Suzumoto (BPBM), M. Boeseman (RMNH), Ofer Gon and M.M. Smith (RUSI), Leslie W. Knapp (SOSC, Smithsonian Institution), and Lev Fishelson (UTAI). Janet C. Gomon, Susan L. Jewett, David G. Smith, and Jeffery T. Williams, all of the Smithsonian Institution, aided in curatorial processes. Rudie Kuiter kindly sent underwater transparencies of the three Red Sea species from his recent trip to Egypt. The first author was supported by a Smithsonian post-doctoral fellowship for an early part of this study. TFH funds were provided by Victor G. Springer for several study trips to the Smithsonian by the first author.

Table 1. Frequency distribution of gill-raker counts (including rudiments) for Apogon species

	Total							Upper Limb			Lower Limb
	24	25	26	27	28	29	30	6	7	8	17	18	19	20	21	22
Apogon annularis		1	2	15	11	2	1	3	23	6			3	17	10	2
Apogon zebrinus	1	5	14	10	9			13	22	4	1	7	16	15
Apogon guamensis	1	5	19	13	4			5	28	11		6	25	11	2
Apogon savayensis		7	9	13	21	9		16	41			8	7	25	9
Apogon sp.			7	29	41	24	12	3	63	40		1	9	30	53	13

Table 2. Preserved pattern characteristics of certain Apogon species.

Characteristic	annularis	guamensis	zebrinus	savayensis	Apogon sp
Pale bars on body	none	0-4	many	0 to few, faint	many
Edges of caudal fin	dark	pale	dark	dark	dark
Basicaudal mark
Juvenile	complete band	diffuse spot	complete band	partial band	partial band
Adult	complete band	diffuse spot or saddle	incomplete, band below LL	incomplete, band above LL	incomplete, band above LL
Cheek mark	wide, triangular	narrow line	wide, trangular	wide, triangular	wide, triangular

REFERENCES

FRASER, T. H. & E. A. LACHNER. 1985. A revision of the cardinalfish subgenera Pristiapogon and Zoramia of the Indo-Pacific region (Teleostei: Apogonidae). Smithsonian Contributions to Zoology. 412:1-47, 20 figs., 4 tabs.

FOURMANOIR, P. & A. CROSNIER. 1964(1963). Deuxième liste complémentaire des poissons du canal de Mozambique. Diagnoses préliminaire de 11 espèces nouvelles. Cah. O.R.S.T.L.M. oceanogr. series Nosy-Be. 2, (6):1-32, Pls. 12-16, 15 figs.

GARMEN, S. 1903. Some fishes from Australasia. Bulletin of the Museum of Comparative Zoology. Harvard, 39(8):229-241, 5 Pls.

GON, O. 1987. Redescription of Apogon (Ostorhinchus) fleurieu (Lacepède, 1802) with notes on its synonymy. Japanese Journal of Ichthyology. 34(2):138-145, 2 figs., 1 tab.

GÜNTHER, A.C.L.G. 1859. Catalogue of the Acanthopterygian fishes in the collection of the British Museum. London, 1:i-xxxiii, 1-524.

GÜNTHER, A.C.L.G. 1871. Report on several collections of fishes recently obtained for the British Museum. Proceedings of the Zoological Society of London. 652-675, Pls. 53-70.

JORDAN, D. S. AND A. SEALE. 1906. The fishes of Samoa. Description of the species found in the Archipelago with a provisional check list of the fishes of Oceania. Bulletin of the Bureau of Fisheries, 1905, 25:175-488, Pls. 23-53, 111 figs.

KLAUSEWITZ, W. 1959. Fische aus dem Roten Meer. II. Knochenfische der familie Apogonidae (Pisces,Percomorphi). Senckenbergiana biologica. 40(5/6):251-262, 11 figs.

LACHNER, E.A. 1951. Studies of certain apogonid fishes from the Indo-Pacific, with descriptions of three new species. Proceedings of the United States National Museum. 101(3290):581-610, Pls. 17-19, Fig. 105, 6 tabs.

RANDALL, J.E., T.H. FRASER AND E.A. LACHNER. 1990. On the validity of the Indo-Pacific cardinalfishes Apogon aureus (Lacepede) and A. fleurieu (Lacepede), with description of a related new species from the Red Sea. Proceedings of the Biolological Society of Washington, 103 (1): 39-62, 9 figs., 4 tabs.

RÜPPELL, W.P.E.S. 1828. Atlas zu der Reise im nordlichen Afrika. Zoologie. Fisches des Rothen Meeres. Frankfurt-a-M., 144pp., 35 Pls.

RÜPPELL, W.P.E.S. 1835-38. Neue wirbelthiere zu der Fauna von Abyssinien gehorig. Fisches des Rothen Meeres. Frankfort-a-M., 1-148, 33 Pls.

SMITH, J.L.B. 1964. A new sponge-dwelling apogonid fish from the Red Sea. Annals & Magazine of Natural History. (13)7:529-531, 1 fig.

VALENCIENNES, A. 1832. Descriptions de plusieurs espèces nouvelles de poisson du genre Apogon. Nouvelles Annales Museum Histoire Naturelle, Paris. 1:51-60, Pl. 4.

VON BONDE, C. 1923. Shallow-water fishes procured by the S.S. Pickle. Rep. Fish. Mar. Biol. Survey, Union S. Africa, 3(1):1-40, Pls. 1-9.

WEBER, M.1911. Die Fische der Aru- und Kei- Iseln. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft. Frankfurt. 34(1):1-49, 2 Pls., 11 figs.

WINTERBOTTOM, R., A. R. EMERY AND E. HOLM. 1989. An annotated checklist of the fishes of the Chagos Archipelago, Central Indian Ocean. Royal Ontario Museum ,Life Sciences Contributions 145: 1-226

The following images related to this document are available:

Photo images

Line drawing images