Four new Indo-Pacific species of cardinalfishes (Apogonidae)

aqua, Journal of Ichthyology and Aquatic Biology Four new Indo-Pacific species of cardinalfishes (Apogonidae) Gerald R. Allen 1 and John E. Randall 2 1) Department of Aquatic Zoology, Western Australian Museum, Francis Street, Perth, Western Australia 6000 and Conservation International, 1919 M. Street N.W. Suite 600, Washington, DC 20036, U.S.A. 2) Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817-2704, U.S.A. Accepted: 24.04.2002 Keywords Taxonomy, marine fishes, Apogon, new species, Apogonidae, Indo-Pacific Abstract The following four Indo-Pacific cardinalfishes of the genus Apogon are described as new: A. lineomaculatus from Bali and Flores, Indonesia from 8-32 m (usually over 18 m) is distinct in having 15 pectoral rays, 4 predorsal scales, 5 + 18-20 developed gill rakers, a narrow mid-lateral black stripe ending in a large black spot at caudal fin base, a black blotch above anal fin base, and numerous faint narrow blackish bars on lower side; A. microspilos, described from two specimens taken in 21.5 m off Lombok, Indonesia, has 13 pectoral rays, 3 median predorsal scales, 3-5 + 14-15 developed gill rakers, and is orange-red in life with a very small dark spot at mid-base of caudal fin; A. cladophilos, described from six specimens taken in 20 m at Lizard Island on the Great Barrier Reef but also photographed in Flores (at both localities in association with soft corals), has 14 pectoral rays, 4 median predorsal scales, 4-5 + 16-17 developed gill rakers, a thin dark mid-lateral stripe and a pupil-size black spot at caudal fin base; and A. posterofasciatus, described from three specimens from the Solomon Islands, Flores, and the Philippines at depths of 18-37 m and distinguished by VI dorsal spines (the above three species with VII), 6 predorsal scales, 2 + 11 developed gill rakers, a membranous flap at the angle of the preopercle, and two dusky bars posteriorly on body, one below rear base of second dorsal fin and one at base of caudal fin. Zusammenfassung Die folgenden vier Kardinalfische der Gattung Apogon aus dem Indo-Pazifik werden neu beschrieben: A. lineomaculatus aus Bali und Flores (Indonesien) aus 8 32 Metern Tiefe (meistens tiefer als 18 m) fällt durch die Anwesenheit von 15 Brustflossenstrahlen, 4 Predorsalschuppen, 5 + 18 20 entwickelten Kiemenreusen, einem schmalen, schwarzen mittleren Seitenstreifen, der in einen schwarzen Punkt auf der Schwanzflossenwurzel ausläuft, einem schwarzen Flecken über der Afterflosse, sowie einer Anzahl von schmalen, leicht Schwärzlichgefärbten Streifen an der Unterseite, auf; A. microspilos, beschrieben anhand von zwei Exemplaren, die in einer Tiefe von 21.5 m vor Lombok (Indonesien) gefangen wurden, hat 13 Brustflossenstrahlen, 3 mittlere Predorsalschuppen, 3-5 + 14-15 entwickelte Kiemenreusen, und lebende Exemplare haben eine orange-rote Färbung mit einem sehr kleinen, dunklen Punkt am mittleren Teil der Schwanzflossenwurzel; A. cladophilos, beschrieben anhand von sechs Exemplaren aus 20 m Tiefe bei Lizard Island am Großen Barrierenriff aber auch vor Flores fotografiert (in beiden Fundorten im Zusammenhang mit Weichkorallen), hat 14 Brustflossenstrahlen, 4 mittlere Predorsalschuppen, 4-5 + 16-17 entwickelte Kiemenreusen und einen schmalen, dunklen mittleren Seitenstreifen und einem pupillen-großen schwarzen Fleck auf der Schwanzflossenwurzel; sowie A. posterofasciatus, beschrieben anhand von drei Exemplaren aus den Salomon Inseln, Flores und den Philippinen aus einer Tiefe von 18 37 m, der sich durch VI Rückenflossenstacheln (die obigen drei Arten jeweils mit VII Stacheln), 6 Predorsalschuppen, 2 + 11 entwickelten Kiemenreusen, einem Hautlappen im Winkel des vorderen Kiemendeckels und zwei dunkle Streifen am Hinterkörper (einer davon an der hinteren Basis der zweiten Rückenflosse und der andere auf der Schwanzflossenwurzel), auszeichnet. Résumé Les quatre espèces d'apogonidés de l'indo-pacifique du genre Apogon décrites ici sont nouvelles: A. lineomaculatus de Bali et Flores (Indonésie), capturé entre 8-32 m (généralement plus de 18 m) se distingue par 15 rayons pectoraux, 4 écailles prédorsales, 5 + 18-20 branchiospines dévéloppées, une étroite bande noire médiane se terminant en grande tache noire à la base de la caudale, une zone noire au-dessus de la base de l'anale et de multiples barres fines, pâles, noirâtres sur le partie inférieure; A. microspilos, décrit sur base de deux spécimens capturés à 21,5 m au large de Lombok, Indonésie, a 13 rayons pectoraux, 3 écailles médianes prédorsales, 3-5 + 14-15 branchiospines développées et, vivant, est rouge orange avec une minuscule 115 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) tache noire au centre de la base de la caudale; A. cladophilos, décrit sur base de six spécimens capturés à 20 m à Lizard Island (Grande Barrière de Corail), mais photographié aussi à Flores (en association avec des coraux mous aux deux endroits), a 14 rayons pectoraux, 4 écailles médianes prédorsales, 4-5 + 16-17 branchiospines développées, une mince bande latérale et une tache noire, de la taille d'une pupille, à la base de la caudale; et A. posteriofasciatus, décrit sur base de trois spécimens des îles Salomon, Flores et les Philippines, à des profondeurs de 18-37 m et se distinguent par VI épines dorsales (les trois espèces ci-dessus avec VII), 6 écailles prédorsales, 2 + 11 branchiospines développées, une excroissance membraneuse à l'angle du préopercule et deux barres grisâtres à l'arrière du corps, l'une sous la base de la seconde dorsale et l'autre à la base de la caudale. Sommario In questo articolo vengono descritte le seguenti quattro nuove specie di pesci cardinale del genere Apogon: A. lineomaculatus, raccolto a Bali e Flores, in Indonesia, a profondità di 8-32 m (di solito maggiori di 18 m), si distingue per avere 15 raggi pettorali, 4 scaglie predorsali, 5 + 18-20 branchiospine sviluppate, una sottile stria nera sui fianchi terminante con una grossa macchia anch essa nera alla base della pinna caudale, una grossa macchia nera diffusa sopra la base della pinna anale e numerose, deboli striature nerastre sul lato inferiore del corpo; A. microspilos, descritto sulla base di due esemplari raccolti a 21.5 m di profondità al largo di Lombok, in Indonesia, possiede 13 raggi pettorali, 3 scaglie predorsali, 3-5 + 14-15 branchiospine sviluppate, e si presenta con un bel color rosso-aranciato e una piccola macchia scura al centro della base della pinna caudale; A. cladophilos, descritto sulla base di 6 esemplari raccolti a 20 m di profondità presso l isola Lizard nella Grande Barriera Corallina Australiana, ma fotografato anche a Flores (in entrambe le località è stato osservato in associazione con coralli molli), ha 14 raggi pettorali, 4 scaglie predorsali, 4-5 + 16-17 branchiospine sviluppate, una banda scura longitudinale mediale molto sottile e una macchia nera grande come la pupilla alla base della pinna caudale; A. posterofasciatus, descritto sulla base di tre esemplari raccolti presso le Isole Solomone, a Flores e nelle Filippine a profondità di 18-37 m, si contraddistingue per avere VI spine dorsali (le altre tre specie sopra descritte ne hanno VII), 6 scaglie predorsali, 2 + 11 branchiospine sviluppate, un lembo membranoso all angolo del preopercolo e due barre fosche posteriormente sul corpo, una al di sotto della base posteriore della seconda pinna dorsale e l altra alla base della pinna caudale. Introduction Cardinalfishes of the family Apogonidae are common inhabitants of coral reefs and adjacent habitats. Although primarily nocturnal, most are readily observed during daylight hours, usually in caves and crevices or around coral formations. The generic classification of the family was reviewed by Fraser (1972), but the documentation of species is far from complete. Fraser recognized 19 genera; with the estimated number of species at about 300, of which approximately 75 percent occur in the Indo-Pacific region. At least half of the species inhabit Australia, New Guinea, Indonesia, and the Philippines regions. This area has been a particularly fertile collecting ground for new species over the past few decades, particularly since the advent of SCUBA diving, which has enabled scientists to reach previously inaccessible habitats and depths. The present paper describes four new species of the genus Apogon, collected mainly by the authors between 1973 and 1992 at the Solomon Islands, Indonesia, Philippines and Australia s Great Barrier Reef. Materials and Methods Lengths given for specimens are standard length (SL), measured from the front of the upper lip to the base of the caudal fin (posterior end of hypural plate); body depth is the maximum depth from the base of the dorsal spines; body width is measured just posterior to the gill opening; head length is taken from the front of the upper lip to the end of the opercular membrane, and snout length from the same anterior point to the fleshy edge of the orbit; orbit diameter is the greatest fleshy diameter, and interorbital width the least bony width; caudal peduncle depth is the least depth, and caudal peduncle length the horizontal distance between verticals at the posterior base of the anal fin and the base of the caudal fin; lengths of fin spines and soft rays are measured to their extreme bases; caudal concavity is the horizontal distance between verticals at the tips of the shortest and longest caudal fin rays; pectoral ray counts include the upper rudimentary ray; lateral line scale counts are made to the base of the caudal fin (hence do not include the pored scales posterior to the hypural plate); gill raker counts are made on the first gill arch and include developed rakers only (those which are higher than the width of their base); the count of the upper-limb rakers is given first, followed by the lower-limb count; the raker at the angle is contained in the lower-limb count. Proportional measurements of type specimens are given in Tables I-III as percentages of the standard length. Data in parentheses in the description refer to paratypes. Type specimens have been deposited at the Bishop Museum, Honolulu (BPBM), Muséum National d Histoire Naturelle, Paris (MNHN); Pusat Penelitian dan Pengembangan Oseanologi, Jakarta, Indonesia (NCIP), U.S. National Museum of Natural History, Washington, D.C. (USNM), and the Western Australian Museum, Perth (WAM). aqua vol. 5 no. 3-2002 116

Gerald R. Allen and John E. Randall Apogon lineomaculatus n. sp. (Figs. 1 and 2; Table I) Apogon thermalis (non Cuvier).- Kuiter, 1992: 62, fig. B (Bali). Holotype: NCIP 6154, 49.0 mm SL, Tulamben, Bali, Indonesia: near rocky point on east side of Tulamben Beach (approximately 8 18 S, 115, 37 E), in 20-25 m, hand net at night, G. R. Allen, 20 May 1992. Paratypes: BPBM 32233, 2 specimens, 44.3-48.5 mm SL, Maumere, Flores, Indonesia: Waipare Reef, off Pertamina fuel storage site (approximately 8 37 46 S, 122 16 35 E), in 32 m, hand net, R. H. Kuiter, 20 September 1987; BPBM 34081, 43.2 mm SL, Japanese shipwreck east of Maumere, Flores, Indonesia, in 23 m, spear, J. E. Randall, 17 September 1988; BPBM 34159, 2 specimens, 38.7-46.4 mm SL, Maumere Bay, Flores, Indonesia: off Sao Wisata Resort, in 8 m, rotenone, J. E. Randall, 19 September 1988; MNHN 1998-1380, 2 specimens, 37.6-42.6 mm SL, same data as BPBM 34159; NCIP 6155, 3 specimens, 23.4-36.0 mm SL, collected with holotype; USNM 351559, 2 specimens, 37.7-38.7 mm SL, same data as BPBM 34159; WAM 30514-002, 2 specimens, 45.0-47.2 mm SL, collected with holotype. Diagnosis Dorsal rays VII-I,9; anal rays II,8; pectoral rays 15; lateral line scales 25; predorsal scales 4; developed gill rakers 5 + 18-20; preopercular margin serrate; preopercular ridge finely serrate; body depth usually 3.1- Fig. 1. Apogon lineomaculatus, paratype, 48.5 mm SL, Maumere, Flores, Indonesia. Photo by J. E. Randall. Fig. 2. Underwater photograph, taken at night, of Apogon lineomaculatus, about 50 mm SL, Tulamben, Bali, Indonesia. Photo by G. R. Allen. 117 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) 3.2 in SL; fourth dorsal spine longest; overall colour pale with a narrow black stripe along middle of side and a large black spot at base of caudal fin; additional colour pattern features include a series of narrow dark bars on the lower side, a black patch above the anal fin base, and black outer edge on the first dorsal fin; peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Description Dorsal rays VII-I,9; anal rays II,8 ; all dorsal and anal soft rays branched, the last to base; pectoral rays 15, the lowermost and uppermost pair unbranched; pelvic rays I, 5, all rays branched; principal caudal rays 17, the upper and lower rays unbranched; upper and lower procurrent caudal rays 8-10, the last (and occasionally the one preceding it) segmented; lateral line scales 25, plus 3 pored scales posterior to hypural plate, the last long, narrow, and somewhat triangular; scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 6; median predorsal scales 4; circumpeduncular scales 12; developed gill rakers 5 + 20 (5 + 18 or 19), usually 2-3 tiny r u d i m e n t s a t b e g i n n i n g o f u p p e r l i m b ; branchiostegal rays 7; supraneural (predorsal) bones 3; vertebrae 10 + 14. Body depth 3.1 (3.1-3.2, except one specimen 3.6) in SL; body width 2.0 (2.0-2.5) in depth; head length 2.6 (2.5-2.7) in SL; dorsal profile of head straight, snout pointed; snout length 3.8 (3.8-5.3) in head; orbit diameter 3.0 (2.9-3.3) in head; interorbital width 5.2 (5.3-5.5) in head; caudal peduncle depth 3.1 (3.0-3.6) in head; caudal peduncle length 1.5 (1.4-1.6) in head. Mouth large, the maxilla reaching a vertical through middle of eye, the upper jaw length 2.3 (2.4-2.5) in head; mouth oblique, the gape forming an angle of about 33 (34-40) degrees to horizontal axis of head and body; posterior edge of maxilla slightly concave; no supramaxilla present; lower jaw slightly protruding; jaws with dense patches of small villiform teeth arranged in irregular rows, the patches decreasing in width posteriorly; an irregular row of small teeth forming a V on vomer and a straight, narrow row on each palatine. Tongue broad-based with rounded tip. Anterior nostril a small, low-rimmed, membranous tube directly in front of centre of eye about half distance from edge of orbit to front of upper lip; posterior nostril ovate without a rim, on a line between anterior nostril to top of orbit, more than twice as large as anterior nostril. Cephalic lateralis pores numerous, especially on interorbital, around eye, and on ventral surface of lower jaw. Preopercular margin finely serrate (40 serrae on left side of holotype and 17-49 on paratypes, becoming more numerous with increased size); preopercular ridge finely serrate, particularly around angle and on lower edge. Table I. Proportional measurements of type specimens of Apogon lineomaculatus expressed as percentages of the standard length. * = fin damaged. Character Holotype Paratype Paratype Paratype Paratype Paratype NCIP BPBM BPBM BPBM MNHN USNM 6154 32233 34159 32233 1998-1380 351559 Standard length (mm) 49.0 48.5 46.4 44.3 42.6 38.7 Body depth 32.7 31.1 32.5 27.8 31.2 31.0 Body width 16.7 15.5 12.9 11.3 15.7 14.7 Head length 39.0 38.8 37.5 37.0 40.6 37.7 Snout length 10.2 9.7 9.1 7.9 7.7 7.8 Orbit diameter 12.9 13.8 12.3 12.4 12.4 12.9 Interorbital width 7.6 7.2 6.5 7.0 7.5 7.0 Upper jaw length 16.7 16.7 15.7 15.3 16.7 15.8 Caudal peduncle depth 12.7 13.2 11.4 11.7 11.3 12.4 Caudal peduncle length 25.9 25.6 26.7 26.0 24.9 28.4 Predorsal length 39.0 40.0 37.7 37.9 41.1 39.8 Pre-anal length 61.2 61.9 61.0 58.7 62.9 59.9 Prepelvic length 37.3 37.5 36.2 34.3 38.5 35.4 Length 1st dorsal spine 1.6 2.3 1.9 2.7 2.6 1.8 Length 2nd dorsal spine 9.6 10.9 11.0 9.9 13.1 10.9 Longest dorsal spine 16.3 15.1 14.7 13.1* 17.1 14.7 Spine of 2nd dorsal fin 11.6 12.8 10.8 12.6 12.7 12.1 Longest soft dorsal ray 21.0 18.4 19.2 19.2 21.1 21.4 Length 1st anal spine 1.6 2.1 1.5 1.8 1.9 1.8 Length 2nd anal spine 10.8 11.3 10.1 12.0 12.0 11.1 Longest anal soft ray 18.4 19.6 16.8 19.9 18.3 17.6 Caudal fin length 25.7 26.4 26.1 23.0* 27.0 23.0* Caudal concavity 6.1 5.8 8.6 * 7.0 * Pectoral fin length 21.4 23.5 21.6 23.3 23.5 23.3 Pelvic spine length 13.1 14.6 12.9 14.9 16.0 13.7 Pelvic fin length 19.2 20.6 19.0 20.3 21.1 18.9 aqua vol. 5 no. 3-2002 118

Gerald R. Allen and John E. Randall Scales weakly ctenoid. Lateral line conspicuous, nearly paralleling dorsal contour of body, and ending at caudal fin base (three pored scales posterior to hypural, the last pointed). No scales on dorsal and anal fins except a narrow sheath at base of second dorsal and anal fins; small scales on basal quarter of caudal fin; no scales on paired fins except a pair of large mid-ventral scales at base of pelvic fins. Origin of first dorsal fin above third lateral line scale; first dorsal spine very short, 23.9 (13.7-20.9) in head; fourth dorsal spine longest, 2.4 (2.4-2.6) in head; anterior soft dorsal rays longest, 1.9 (1.8-2.1) in head; origin of anal fin below base of third or fourth dorsal soft ray; first anal spine very short, 23.9 (19.1-24.9) in head; second anal spine 3.6 (3.1-3.7) in head; anterior anal soft rays longest, 2.1 (1.9-2.3) in head; caudal fin forked, its length 1.5 (1.4-1.6) in head; caudal concavity 6.4 (4.4-6.8) in head; upper pectoral rays longest, 1.8 (1.6-1.8) in head; origin of pelvic fins below pectoral fin base; first pelvic soft ray longest, reaching about midway between anus and origin of anal fin, its length 2.0 (1.8-2.0) in head. Colour in life: ground colour mauve (darkest on back) to nearly whitish with golden reflections; a dark brown stripe from snout to front of eye, continuing behind eye to middle of caudal peduncle, tapering in width posteriorly and ending in front of pupil-sized black spot mid-laterally at caudal fin base, anterior half of spot usually ringed with golden yellow; narrow, faint brown bars on lower half of side, about one per transverse scale row, originating from mid-lateral body stripe; pair of faint brown stripes extending from snout, over each eye and ending above opercle; a faint middorsal str ipe from snout tip to dorsal fin origin; a faint brown stripe sometimes evident on narrow scaly sheath of second dorsal fin; small, but brilliant white spot on tip of snout; tip of lower jaw noticeably dusky; elongate blackish patch immediately above anal fin base; fins translucent except for blackish outer part of first dorsal fin. Colour in alcohol: preserved specimens overall tan or yellowish with a dark mid-lateral stripe and prominent dark spot in the middle of the caudal fin base; elongate blackish patch immediately above anal fin base; tip of lower jaw dusky brown; fins yellowish or whitish except outer part of first dorsal fin broadly blackish. Peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Remarks Apogon lineomaculatus is currently known only from the islands of Bali and Flores, but probably occurs at least throughout the Nusa Tenggara chain, which extends from Lombok to Timor. It is found solitary or in small groups near the bottom, either over open sand at night or in caves and crevices during the day. It is often seen at the base of steep slopes on sand or rubble bottoms. Depth of collection ranged between 8 and 32 m, but the species is usually encountered below 18 m. A fish described as Apogon semilineatus (non Temminck & Schlegel) and reported from north-western Australia by Sainsbury et al. (1985: colour illustration on p.149) is possibly undescribed. It is superficially similar to A. lineomaculatus, but differs in having 14 pectoral rays and only 3 + 14-15 developed gill rakers. Moreover, it has a more distinct stripe above the eye that extends to below the second dorsal fin rather than terminating above the opercle as in A. lineomaculatus. It also lacks the blackish patch immediately above the anal fin base. Apogon lineomaculatus belongs to the subgenus Nectamia Jordan, as defined by Fraser (1972). However, this name has subsequently been replaced by Ostorhinchus Lacépède (Randall, et al. 1990). This subgenus is by far the largest within Apogon, containing at least 70 species. The new species appears to be allied to A. franssedai and A. ocellicaudus, both described by Allen, Kuiter, and Randall (1994) from Indonesian seas. The three species share similar meristics and proportions, as well as a pale ground colour, and large dark spot on the caudal fin base. However, they are easily separated on the basis of colour pattern: A. ocellicaudus lacks a dark mid-lateral stripe, the stripe above the eye on A. franssedai extends to the middle of the back, and both species lack the dark supra-anal patch and series of faint bars on the lower sides, which are typical of A. lineomaculatus. In addition, both A. franssedai and A. ocellicaudus have 14 pectoral rays and 14-16 rakers on the lower limb of the first gill arch. Kuiter (1992) identified this species as Apogon thermalis Cuvier, presumably going by similarities to the illustration of this species in Smith and Heemstra (1986: Plate 49, fig. 175.26). However, this species, a senior synonym of A. sangiensis Bleeker from the Indo- Malay Archipelago, does not have a stripe along the middle of the body, lacks a dark patch above the anal fin base, and possesses six spines in the first dorsal fin. Etymology The species is named lineomaculatus: (line-spotted) with reference to the most distinctive features of its colour pattern. Apogon microspilos n. sp. (Fig 3; Table II) Holotype: NCIP 6156, 54.0 mm SL, Gili Ayer, Lombok, Indonesia, in 21.5 m, spear, J. E. Randall, 22 February 1984. Paratype: BPBM 30149, 40.2 mm SL, collected with holotype. Diagnosis Dorsal rays VII-I,9; anal rays II,8; pectoral rays 13; 119 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) Fig. 3. Underwater photograph of Apogon microspilos, holotype, 54.0 mm SL, Lombok, Indonesia. Photo by J. E. Randall. lateral line scales 24; predorsal scales 3; developed gill rakers 3-5 + 14-15; preopercular margin serrate; preopercular ridge smooth ; body depth 2.5-2.6 in SL; third dorsal spine longest; overall colour pale (orangered in life) with small, dark spot at middle of caudal fin base and pair of silvery stripes through eye; peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Description Dorsal rays VII-I,9; anal rays II,8 ; all dorsal and anal soft rays branched, the last to base; pectoral rays 13, the lowermost and uppermost pair unbranched; pelvic rays I, 5, all rays branched; principal caudal rays 17, the upper and lower rays unbranched; upper and lower procurrent caudal rays 7-8, the most posterior (and occasionally the one preceding it) segmented; lateral line scales 24, plus 3 (4) pored scales posterior to hypural plate, the last rounded on holotype, but triangular on paratype; scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 5; median predorsal scales 3; circumpeduncular scales 12; developed gill rakers 5 + 14 (3 + 15), usually 1-3 tiny rudiments at beginning of upper limb; branchiostegal rays 7; supraneural (predorsal) bones 3; vertebrae 10 + 14. Body depth 2.5 (2.6) in SL; body width 2.4 (2.2) in depth; head length 2.5 in SL; dorsal profile of head straight, snout relatively blunt; snout length 4.7 (5.1) in head; orbit diameter 2.3 (2.5) in head; interorbital width 5.3 (5.2) in head; caudal peduncle depth 2.2 (2.3) in head; caudal peduncle length 1.7 in head. Mouth large, the maxilla reaching a vertical past middle of eye, the upper jaw length 2.0 (2.1) in head; mouth oblique, the gape forming an angle of about 37 degrees to horizontal axis of head and body; posterior edge of maxilla slightly concave; no supramaxilla present; lower jaw slightly protruding; upper and lower jaw with dense band of small villiform teeth, narrowing towards rear of mouth, with a toothless gap at front of Table II. Proportional measurements of type specimens of Apogon microspilos expressed as percentages of the standard length. * = fin damaged Character Holotype Paratype NCIP BPBM 6156 30149 Standard length (mm) 54.0 40.2 Body depth 40.6 38.3 Body width 16.7 17.2 Head length 39.3 40.3 Snout length 8.3 8.0 Orbit diameter 17.0 16.4 Interorbital width 7.4 7.7 Upper jaw length 19.6 19.4 Caudal peduncle depth 18.0 17.9 Caudal peduncle length 22.8 24.4 Predorsal length 60.2 42.8 Pre-anal length 66.5 64.9 Prepelvic length 39.6 39.1 Length 1st dorsal spine 1.5 1.7 Length 2nd dorsal spine 6.1 7.5 Longest dorsal spine 19.4 19.9 Spine of 2nd dorsal fin 14.1 17.4 Longest soft dorsal ray 23.7 26.1 Length 1st anal spine 1.3 1.7 Length 2nd anal spine 10.4 13.2 Longest anal soft ray 18.5 19.7 Caudal fin length 25.4* 32.8 Caudal concavity * * Pectoral fin length 24.8 26.6 Pelvic spine length 14.4 15.9 Pelvic fin length 21.1 23.6 aqua vol. 5 no. 3-2002 120

Gerald R. Allen and John E. Randall jaw; an irregular row of small teeth forming a V on vomer and a straight, narrow row on each palatine. Tongue broad-based with rounded tip. Anterior nostril a small, low-rimmed, membranous tube directly in front of centre of eye, but closer to upper lip than to eye; posterior nostril ovate without a rim, on a line connecting anterior nostril to top of orbit, more than twice as large as anterior nostril. Cephalic lateralis pores numerous, especially on interorbital, around eye, and on ventral surface of lower jaw. Preopercular margin finely serrate (67 serrae on left side of holotype and 41on paratype), the number apparently increasing with growth); preopercular ridge smooth. Scales weakly ctenoid. Lateral line conspicuous, nearly paralleling dorsal contour of body, and ending at caudal fin base (3-4 pored scales posterior to hypural). No scales on dorsal and anal fins except a narrow sheath at base of second dorsal and anal fins; small scales on basal third of caudal fin; no scales on paired fins except a pair of large mid-ventral scales at base of pelvic fins (missing from damaged holotype). Origin of first dorsal fin above third lateral line scale; first dorsal spine very short, 26.5 (23.1) in head; third dorsal spine longest, 2.0 in head; anterior soft dorsal rays longest, 1.7 (1.5) in head; origin of anal fin below base of third or fourth dorsal soft ray; first anal spine very short, 30.3 (23.1) in head; second anal spine 3.8 (3.1) in head; anterior anal soft rays longest, 2.1 in head; caudal fin forked, but badly damaged in both types; upper pectoral rays longest, 1.6 (1.5) in head; origin of pelvic fins slightly anterior to pectoral fin base; first pelvic soft ray longest, reaching to about anus, its length 1.9 (1.7) in head. Colour in life: mainly orange-red grading to dull mauve on back and caudal peduncle; a tiny, inconspicuous dark spot at middle of caudal fin base; a pair of thin silver stripes extending from snout through eye, bordering upper and lower edge of pupil; fins translucent to slightly reddish except blackish membrane between third and fourth dorsal spines. Colour in alcohol: uniform tan with faint dark spot at middle of caudal fin base; fins translucent to tan. Peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Remarks Apogon microspilos is known only from the type specimens collected at a depth of 21.5 m at Lombok, Indonesia. This species belongs to the subgenus Ostorhinchus Lacépède (see comments in Remarks section for A. Table III. Proportional measurements of type specimens of Apogon cladophilos expressed as percentages of the standard length. * = fin damaged. Character Holotype Paratype Paratype Paratype Paratype Paratype WAM BPBM AMS AMS USNM BPBM P.31480 37873 I.38996 I.38996 351560 37873-001 -001-001 Standard length (mm) 30.0 29.3 28.5 27.9 25.9 24.3 Body depth 31.7 30.7 30.9 30.5 30.9 29.2 Body width 13.3 12.6 12.6 14.3 11.6 9.9 Head length 37.0 40.3 40.7 39.1 40.5 39.1 Snout length 7.3 7.5 7.0 7.2 7.7 7.0 Orbit diameter 13.7 12.6 13.3 13.3 14.3 13.6 Interorbital width 7.7 7.8 7.4 8.2 7.7 6.2 Upper jaw length 18.0 19.1 17.5 17.9 19.3 17.7 Caudal peduncle depth 12.0 12.6 12.6 12.9 13.5 9.9 Caudal peduncle length 24.0 24.2 22.8 23.3 23.9 23.5 Predorsal length 38.0 37.5 40.7 39.1 41.3 38.7 Pre-anal length 63.7 62.1 62.5 59.9 63.7 61.7 Prepelvic length 37.0 39.2 37.2 37.6 37.5 35.8 Length 1st dorsal spine 2.0 2.0 2.1 1.8 1.9 2.1 Length 2nd dorsal spine 7.7 8.2 9.1 8.2 8.5 9.1 Longest dorsal spine 14.0 15.7 15.8 14.0 15.1 16.5 Spine of 2nd dorsal fin 13.0 13.3 13.0 11.8 11.6 12.3 Longest soft dorsal ray 23.7 25.6 24.6 22.2 23.2 18.5* Length 1st anal spine 1.7 2.4 1.8 1.8 1.9 1.6 Length 2nd anal spine 13.3 11.9 10.9 10.8 11.6 11.5 Longest anal soft ray 19.3 22.5 16.8* 19.7 19.3 19.8 Caudal fin length 11.3* 33.1 21.1* 19.4* 29.0 27.2 Caudal concavity * 9.2 * * 8.5 6.2 Pectoral fin length 26.0 24.9 25.3 25.4 27.4 24.7 Pelvic spine length 13.3 13.7 13.7 12.9 13.5 12.3 Pelvic fin length 20.7 20.8 20.7 19.4 20.1 20.6 121 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) lineomaculatus). It does not appear to have any obvious close relatives, but is similar in some respects to A. griffini Seale from Sabah and the Philippines. Both species possess a similar shape, although A. griffini has distinctly elongate rays at the beginning of the second dorsal fin which form a filamentous extension. Both species also have a pair of silvery stripes through the eye and a faint spot at the middle of the caudal fin base. However, they differ in general coloration of the body and fins. A. griffini lacks the orange colour on the sides and has orange to yellow dorsal and anal fins. It also has an additional pectoral fin ray (14 versus 13). Moreover, it is a shallow-dwelling species usually encountered in areas of mixed weed and coral at depths of less than 8 m. A. sabahensis Allen & Kuiter 1994 is a junior synonym of A. griffini. Etymology The species is named microspilos (small-spot) in reference to the unusually small dark spot at the midbase of the caudal fin. The Greek word spilos is used as a noun in apposition. Apogon cladophilos n. sp. (Fig. 4 and 5, Table III) Holotype: WAM P.31480-001, 30.0 mm SL, Lizard Island (approximately 14 40 S, 145 28 E), Great Barrier Reef, Australia: off Granite Bluff, in 20 m, rotenone, J. E. Randall and H. W. Kendall, 10 December 1982. Paratypes: (all collected with holotype) AMS I.38996-001, 2 specimens, 27.9-28.5 mm SL; BPBM 37873, 2 specimens, 24.3-29.3 mm SL; USNM 351560, 25.9 mm SL. Diagnosis Dorsal rays VII-I,9; anal rays II,9; pectoral rays 14; lateral line scales 24; predorsal scales 4; developed gill rakers 4-5 + 16-17; preopercular margin serrate; preopercular ridge smooth; body depth 3.2-3.4 in SL; fourth dorsal spine longest; overall colour pale (semitransparent or whitish in life) with a thin, dark midlateral stripe on sides and prominent pupil-sized black spot at caudal fin base; peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Description Dorsal rays VII-I,9; anal rays II,9; all dorsal and anal soft rays branched, the last to base; pectoral rays 14, the lowest and uppermost pair unbranched; pelvic rays I,5, all rays branched; principal caudal rays 17, the upper and lower rays unbranched; upper and lower procurrent caudal rays 8-10, the most posterior (and occasionally the one preceding it) segmented; lateral line scales 24, plus 3 pored scales posterior to hypural plate; scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 5 or 6; median predorsal scales 4; circumpeduncular scales 12; developed gill rakers 4 + 16 (4-5 + 16-17), usually 1-2 tiny rudiments at beginning of upper limb; branchiostegal rays 7; supraneural (predorsal) bones 3; vertebrae 10 + 14. Body depth 3.2 (3.2-3.4) in SL; body width 2.4 (2.1-3.0) in depth; head length 2.7 (2.5-2.6) in SL; dorsal profile of head straight, snout relatively pointed; snout length 5.0 (5.3-5.8) in head; orbit diameter 2.7 (2.8-3.1) in head; interorbital width 4.8 (4.7-6.3) in head; caudal peduncle depth 3.1 (3.0-4.0) in head; caudal peduncle length 1.5 (1.7-1.8) in head. Fig. 4. Underwater photograph of Apogon cladophilos, about 40 mm SL, Flores, Indonesia. Photo by R. H. Kuiter. aqua vol. 5 no. 3-2002 122

Gerald R. Allen and John E. Randall Mouth large, the maxilla reaching a vertical well past middle of eye (to about rear edge of pupil), the upper jaw length 2.1 (2.1-2.3) in head; mouth oblique, the gape forming an angle of about 37 (34-39) degrees to horizontal axis of head and body; posterior edge of maxilla slightly concave; no supramaxilla present; lower jaw slightly protruding; upper and lower jaw with dense band of small villiform teeth, narrowing towards rear of mouth, with a toothless gap at front of jaw; an irregular row of small teeth forming a V on vomer and a straight, narrow row of inconspicuous teeth on each palatine. Tongue broad-based with rounded tip. Anterior nostril tubular, directly in front of centre of eye, about halfway between front of eye and outer edge of upper lip; posterior nostril ovate without a rim, on a line connecting anterior nostril to top of orbit, more than twice as large as anterior nostril. Cephalic lateralis pores numerous, especially on interorbital, around eye, and on ventral surface of lower jaw. Rear edge of preopercular margin finely serrate (35 serrae on left side of holotype and 17-31 on paratypes); preopercular ridge smooth. Scales weakly ctenoid. Lateral line conspicuous, nearly paralleling dorsal contour of body, and ending at caudal fin base (3 pored scales posterior to hypural). No scales on dorsal and anal fins except a narrow sheath at base of second dorsal and anal fins; small scales on basal fourth of caudal fin; no scales on paired fins except a pair of large mid-ventral scales at base of pelvic fins (missing from damaged holotype and three paratypes. Origin of first dorsal fin above third lateral line scale; first dorsal spine very short, 18.5 (19.0-21.8) in head; fourth dorsal spine longest, 2.6 (2.4-2.8) in head; anterior soft dorsal rays longest, 1.6 (1.6-2.1) in head; origin of anal fin below base of first or second dorsal soft ray; first anal spine very short, 22.2 (16.9-23.8) in head; second anal spine 2.8 (3.4-3.7) in head; anterior anal soft rays longest, 1.9 (1.8-2.4, but most damaged) in head; caudal fin forked, but badly damaged in holotype and three paratypes, its length in intact paratypes 1.2-1.4 in head; upper pectoral rays longest, 1.4 (1.4-1.6) in head; origin of pelvic fins slightly anterior to pectoral fin base; first pelvic soft ray longest, reaching to about anus, its length 1.8 (1.9-2.0) in head. Colour in life: overall translucent to whitish with silvery reflections on head and belly; a diffuse to distinct brown stripe extending from snout tip through eye to middle of base of caudal fin, tapering posteriorly and ending in prominent black spot (about pupil size or slightly larger); a pair of silvery stripes on snout passing through eye at level of upper and lower margin of pupil; fins translucent whitish. Fig. 5. Underwater photograph of Apogon cladophilos, about 40 mm SL, Flores, Indonesia. Photo by R. H. Kuiter. 123 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) Colour in alcohol: yellowish-tan with an inconspicuous, faint mid-lateral stripe and prominent black spot at base of caudal fin; fins translucent whitish. Peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Remarks Apogon cladophilos is known only from the northern Great Barrier Reef, New Caledonia, and Indonesia, where it has been photographed at Maumere on the island of Flores (R. H. Kuiter, pers. comm.). The Australian specimens were associated with black coral (Antipathes) and the Indonesian fish were sheltering in an unidentified soft coral. The depth was about 20 m at both localities. A large aggregation sheltering among the spines of a sea urchin (probably Diadema setosum) at New Caledonia was illustrated as A. fraenatus by Fourmanoir and Laboute (1976: 289). This species belongs to the subgenus Ostorhinchus Lacépède (see comments in remarks section for A. lineomaculatus). It superficially resembles Apogon fraenatus, a member of the subgenus Pristiapogon (see Fraser and Lachner, 1985 for a review). Both species are characterized by a pale ground colour and tapering dark mid-lateral stripe that terminates in a centrally located black peduncular spot. However, A. fraenatus, like other members of its subgenus, is characterized by serrations on the upper edge of the infraorbitals and preopercular ridge. It is further distinguished from A. cladophilos by 8 soft anal rays, 7-11 well-developed rakers on the lower limb of the first gill arch, and a more vivid mid-lateral stripe that is solid rather than diffuse. Etymology The species is named cladophilos from the Greek (klados + philo = branch-loving), in reference to its habit of sheltering among branches of antipatharian and alcyonarian corals as well as sea urchins. Apogon posterofasciatus n. sp. (Fig. 6, Table IV) Holotype: BPBM 15669, 42.8 mm SL, Florida Island, Solomon Islands: south side of Tanavula Point (9 02 44 S, 160 04 07 E), in 18-37 m, rotenone, G. R. Allen and J. E. Randall, 30 July 1973. Paratypes: USNM 351561, 34.3 mm SL, Maumere, Flores, Indonesia: fringing reef about 1.5 km north of Maumere town, in 30 m, rotenone, J. E. Randall, 13 October 1987; USNM 349197, 49.0 mm SL, Balicasag Island, Philippine Islands: drop-off on west side (9 31 14 N, 123 40 00 E), 0-24 m, rotenone, V. G. Springer et al., 10 June 1978. Diagnosis Dorsal rays VI-I,9; anal rays II,8; pectoral rays 13; lateral line scales 24; predorsal scales 6; developed gill rakers 2 + 11; preoperculum serrate on rear margin with well-developed membranous flap at the lower corner and along lower margin; preopercular ridge smooth; body depth 2.6-2.7 in SL; second dorsal spine longest; overall colour pale (light yellowish in life) with a pair of dusky bars on posterior part of body; peritoneum pale with scattered melanophores; outer surface of intestinal tract black. Fig. 6. Apogon posterofasciatus, holotype, 42.8 mm SL, Florida Island, Solomon Islands. Photo by J. E. Randall. aqua vol. 5 no. 3-2002 124

Gerald R. Allen and John E. Randall Table IV. Proportional measurements of type specimens of Apogon posterofasciatus expressed as percentages of the standard length. * = fin damaged Character Holotype Paratype Paratype BPBM USNM USNM 15669 349197 351561 Standard length (mm) 42.8 49.0 34.3 Body depth 37.6 37.8 36.4 Body width 17.8 20.4 16.0 Head length 41.4 41.6 39.9 Snout length 10.0 9.0 8.2 Orbit diameter 14.7 14.1 13.7 Interorbital width 6.5 6.5 6.7 Upper jaw length 21.3 21.0 19.8 Caudal peduncle depth 14.5 15.1 14.6 Caudal peduncle length 24.5 25.9 25.1 Predorsal length 45.8 44.9 45.8 Pre-anal length 64.5 61.2 62.7 Prepelvic length 39.7 39.0 35.9 Length 1st dorsal spine 7.7 6.5 4.4 Length 2nd dorsal spine 24.3 23.9 23.3 Longest dorsal spine 24.3 23.9 23.3 Spine of 2nd dorsal fin 15.4 17.3 14.6 Longest soft dorsal ray 21.0 20.8* 24.5 Length 1st anal spine 2.1 2.7 1.2 Length 2nd anal spine 15.4 16.7 15.2 Longest anal soft ray 14.7* 18.8* 21.9 Caudal fin length 23.8* 33.3* 36.4 Caudal concavity * * 10.8 Pectoral fin length 30.1 17.1* 29.2 Pelvic spine length 17.1 17.3 18.1 Pelvic fin length 18.0* 25.5 25.7 Description Dorsal rays VI-I,9; anal rays II,8 ; all dorsal and anal soft rays branched, the last to base; pectoral rays 13, the lowermost and uppermost pair unbranched; pelvic rays I,5, all rays branched; principal caudal rays 17, the upper and lower rays unbranched; upper and lower procurrent caudal rays 7-8, the last (and occasionally the one preceding it) segmented; lateral line scales 24, plus 3 pored scales posterior to hypural plate; scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 6; median predorsal scales 6; circumpeduncular scales 12; developed gill rakers 2 + 11, usually 3 smaller rudiments at anterior end of both upper and lower limb; branchiostegal rays 7; supraneural (predorsal) bones 3; vertebrae 10 + 14. Body depth 2.7 (2.6-2.7) in SL; body width 2.1 (1.9-2.3) in depth; head length 2.4 (2.4-2.5) in SL; dorsal profile of head straight, snout relatively blunt; snout length 4.4 (4.6-5.1) in head; orbit diameter 2.8 (2.9-3.0) in head; interorbital width 6.3 (6.0-6.4) in head; caudal peduncle depth 2.9 (2.7-2.8) in head; caudal peduncle length 1.7 (1.6) in head. Mouth large, the maxilla reaching a vertical well past middle of eye (to about rear edge of pupil), the upper jaw length 1.9 (2.0) in head; mouth oblique, the gape forming an angle of about 35 (33) degrees to horizontal axis of head and body; posterior edge of maxilla slightly concave; no supramaxilla present; lower jaw slightly protruding; upper and lower jaw with dense band of small villiform teeth, narrowing towards rear of mouth, with a toothless gap at front of jaw; an irregular row of small teeth forming a V on vomer and a straight, narrow row of inconspicuous teeth on each palatine. Tongue broad-based with rounded tip. Anterior nostril tubular, directly in front of centre of eye, but much closer to upper lip than to eye; posterior nostril ovate without a rim, on a line connecting anterior nostril to top of orbit, more than twice as large as anterior nostril. Cephalic lateralis pores numerous, especially on interorbital, around eye, and on ventral surface of lower jaw. Rear edge of preopercular margin finely serrate (12 serrae on left side of holotype and 18 on paratypes) with well-developed membranous flap at the lower corner and along lower margin; preopercular ridge smooth. Scales weakly ctenoid. Lateral line conspicuous, nearly paralleling dorsal contour of body, and ending at caudal fin base (3-4 pored scales posterior to hypural). No scales on dorsal and anal fins except a narrow sheath at base of second dorsal and anal fins; small scales on basal third of caudal fin; no scales on paired fins except a pair of large midventral scales at base of pelvic fins (missing from damaged holotype). Origin of first dorsal fin above third lateral line scale; first dorsal spine slender, but moderately long, 5.4 (6.4-9.1) in head; second dorsal spine longest, 1.7 in head; anterior soft dorsal rays longest, 2.0 (1.6) in head; origin of anal fin below base of first or second dorsal soft ray; first anal spine very short, 19.7 (15.7-34.3) in head; second anal spine 2.7 (2.5-2.6) in head; anterior anal soft rays longest, 2.8 (1.8-2.2, but damaged) in head; caudal fin forked, (badly damaged in holotype), its length in paratypes 1.1-1.3 in head; upper pectoral rays longest, 1.4 in head; origin of pelvic fins slightly anterior to pectoral fin base; first pelvic soft ray longest, reaching to about origin of anal fin, its length 1.6 in head in paratypes (damaged in holotype). Colour in life: head light pinkish, grading to light yellowish on body; a broad dusky bar at caudal fin base and a narrower one between posterior rays of anal and second dorsal fins; first dorsal fin pink anteriorly shading to whitish posteriorly, the rays light red; remaining fins with light yellow-orange rays and clear membranes. Shortly after death the entire body assumed a pinkish coloration (Fig. 6 ) Colour in alcohol: light yellowish-tan with a diffuse dusky brown bar at caudal fin base and a narrower, more distinct, bar below rear part of second dorsal fin; fins translucent whitish. Peritoneum pale with scattered melanophores; outer surface of intestinal tract black. 125 aqua vol. 5 no. 3-2002

Four new Indo-Pacific species of cardinalfishes (Apogonidae) Remarks Apogon posterofasciatus is known only from the type specimens collected in the Solomon Islands, Indonesia (Flores), and the Philippines (Balicasag Island). It is probably more widespread in the Indo- Austalian Archipelago, but has escaped detection due to its small size, cryptic habits, and relatively deep distribution (18-37 m). The holotype from the Solomon Islands was collected from a steep-walled gorge with abundant caves in the reef, whereas the Indonesian paratype was taken from an isolated rock on a sand bottom at the base of a fringing reef. This species belongs to the subgenus Apogon Lacépède as defined by Fraser (1972). It is most closely related to A. deetsie Randall 1998, from the Hawaiian Islands and Tuamotu Archipelago, and A. caudicinctus Randall & Smith 1988, which ranges from Réunion in the western Indian Ocean to Rapa Iti in French Polynesia. This trio is characterised by a membranous flap at the lower corner and along the ventral margin of the preopercle. Apogon deetsie has the same general pattern of broad blackish bars posteriorly on the body, but the background colour in life is red, and the outer quarter of the caudal lobes is blackish and the tips of the second dorsal and anal fins are dusky. Apogon caudicinctus is transparent red overall with blackish scale edges and lacks the anterior dark bar, although the posterior bar is evident. Both A. caudicinctus and A. deetsie differ from A. posterofasciatus in having 12 (rarely 11) rather than 13 pectoral rays. Fraser,T. H. 1972. Comparative osteology of the shallow water cardinal fishes (Perciformes: Apogonidae) with reference to the systematics and evolution of the family. Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 34: 1-105. Fraser, T. H. & E. A. Lachner. 1985. A revision of the cardinalfish subgenera Pristiapogon and Zoramia (genus Apogon) of the Indo-Pacific region (Teleostei: Apogonidae). Smithsonian Contributions to Zoology, 412: 1-47. Kuiter, R. H. 1992. Tropical reef fishes of the western Pacific and Indonesia and adjacent waters. PT Gramedia Pustaka Utama, Jakarta, 314 pp. Randall, J. E. 1996. Caribbean reef fishes (2nd edition). T.F.H. Publications, Neptune City, NJ, 369 pp. Randall, J. E. 1998. Review of the cardinalfishes (Apogonidae) of the Hawaiian Islands, with the descriptions of two new species. aqua, Journal of Ichthyology and Aquatic Biology, 3 (1): 25-38. Randall, J. E., Fraser, T. H., & E. A. Lachner. 1990. On the validity of the Indo-Pacific cardinalfishes Apogon aureus (Lacépède) and A. fleurieu (Lacépède), with description of a related new species from the Red Sea. Proceedings of the Biological Society of Washington, 103 (1): 39-62. Sainsbury, K. J., Kailola, P. J., & G. G. Leyland. 1984. Continental shelf fishes of northern and northwestern Australia. CSIRO Division of Fisheries Research, Canberra, Australia, 374 pp. Etymology The species is named posterofasciatus (Latin: with posterior bands ) with reference to its most conspicuous colour feature. Acknowledgements We thank Rudie H. Kuiter (Associate of Museum of Victoria, Melbourne, Australia) for providing photographs and information concerning the Indonesian distribution of Apogon cladophilos. Thomas H. Fraser critically reviewed the manuscript. References Allen, G. R. & R. H. Kuiter. 1994. Descriptions of two new species of cardinalfishes (Apogonidae) from Malaysia and Indonesia. Revue française d Aquariologie, 21 (1-2): 19-23. Allen, G. R., Kuiter, R. H., & J. E. Randall. 1994. Descriptions of five new species of cardinalfishes (Apogon: Apogonidae) from Maumere Bay, Flores, Indonesia and surrounding regions. Revue française d Aquariologie, 21 (1-2): 27-38. Fourmanoir, P. & P. Laboute. 1976. Poissons de Nouvelle Calédonie et des Nouvelles Hébrides Les éditions du Pacifique, Papeete, Tahiti, 376 pp. aqua vol. 5 no. 3-2002 126