Review of the Cardinalfishes (Apogonidae) of the Hawaiian Islands, with Descriptions of Two New Species

John E. Randall aqua, Journal of Ichthyology and Aquatic Biology Review of the Cardinalfishes (Apogonidae) of the Hawaiian Islands, with Descriptions of Two New Species by John E. Randall Bishop Museum, 1525 Bernice St., Honolulu, Hawaii 96817, USA Accepted: 01.12.1997 Keywords Apogonidae, Pseudamiops, Apogon, Apogonichthys, Foa, Lachneratus, new species, Hawaiian Islands. Abstract The 10 species of apogonid fishes of the Hawaiian Islands are differentiated in a key. Pseudamiops diaphanes, sp. n., endemic to Hawaii, is distinct in having VI-I,8 dorsal rays, II,8 anal rays, 16-18 pectoral rays, a spine-like ventral projection posteriorly on the maxilla, deciduous cycloid scales, no lateral line, a rhomboid caudal fin, and in being transparent in life. Apogon deetsie, sp. n., from the Hawaiian Islands and Tuamotu Archipelago, has VI-I,9 dorsal rays, II,8 anal rays; 11-12 (usually 12) pectoral rays, 24 lateral-line scales, 6 predorsal scales, 3-4 + 12-13 gill rakers, a protruding membranous flap on corner and lower edge of preopercle, a forked caudal fin with rounded lobes, and is light red with two blackish bars posteriorly on the body. The remaining species are diagnosed, and their distributions given. Apogon erythrinus Snyder appears to be endemic to Hawaii and Johnston Island; A. maculiferus Garrett is also a Hawaiian endemic; A. evermanni Jordan and Snyder is widespread in the Indo-Pacific and is also known from the western Atlantic; A. kallopterus Bleeker, A. taeniopterus Bennett (A. menesemus Jenkins is a synonym), Apogonichthys perdix Bleeker (A. waikiki Jordan and Evermann is a synonym), Foa brachygramma (Jenkins), and Lachneratus phasmaticus Fraser and Struhsaker are wide-ranging Indo-Pacific species. Zusammenfassung Die 10 Spezies der Apogonidae Fische der hawaiianischen Inseln werden mit einem Schlüssel versehen; Pseudamiops diaphanes, sp. n. ist endemisch um Hawaii, hat VI-1,8 hintere Strahlen, 11,8 anale Strahlen, 16-18 Bruststrahlen, eine wirbelartige, vorgewölbte Maxilla, abfallende, ringlförmige Schuppen, keine Seitenlinie, eine rhombenförmige Caudale, und ist insgesamt transparent. Apogon deetsie, sp. n. kommt bei Hawaii und im Luamotu Archipel vor. Hat VI-1,9 hintere Strahlen, 11,8 anale Strahlen 11-12 (für 25 gewöhnlich 12) Bruststrahlen, 24 seitliche Linien- Schuppen, 6 predorsale Schuppen, 3-4 + 12-13 Kiemenlamellen, einen hervortretenden, hautartigen Lappen an der Ecke und am unteren Ende des Preopercles einen gabelartigen Schwanz mit abgerundeten Enden. Die Art ist hellrot gefärbt mit zwei schwarzen Balken am hinteren Körperteil. Für die verbleibenden Spezies ist eine Diagnose erstellt worden und ihre Verbreitung wird erwähnt. Apogon erythrinus Snyder scheint bei Hawaii endemisch zu sein. A. evermanni Jordan und Snyder ist weitverbreitet im Indo Pazifik und lebt auch im westlichem Atlantik. A. kallopterus Bleeker, A. taeniopterus Bennett (A. menesemus Jenkins ist ein Synonym), Apogonichthus perdix Bleeker (A. waikiki Jordan und Evermann ist ein Synonym), Foa brachygramma (Jenkis) und Lachneratus phasmaticus Fraser und Struhsaker sind weitverbreitete Indo-Pazifik Spezies. Résumé Une clé est proposée qui permet de déterminer les 10 espèces d Apogonidés des îles Hawaii. Pseudamiops diaphanes sp. n., endémique des Îles Hawaii, se distingue par la présence de VI-I,8 rayons dorsaux, II,8 rayons anaux, 16-18 rayons pectoraux, une projection épineuse ventrale insérée postérieurement sur le maxillaire, des écailles cycloïdes décidues, l absence de ligne latérale, une nageoire caudale rhomboïde et la transparence du corps de l animal en vie. Apogon deetsie, sp. n., des Îles Hawaii et de l archipel des Tuamotu, présente VI-I,9 rayons dorsaux, II,8 rayons anaux, 11-12 (habituellement 12) rayons pectoraux, 24 écailles sur la ligne latérale, 6 écailles prédorsales, 3-4 + 12-13 branchiospines, une extension membraneuse à l angle inférieur du préoperculaire, une nageoire caudale fourchue avec des lobes arrondis, une coloration rouge clair avec deux barres noirâtres à l arrière du corps. Pour les autres espèces, la diagnose et la distribution sont indiquées. Apogon erythrinus Snyder semble endémique d Hawaii et de Johnston Island; A. maculiferus Garrett est également endémique d Hawaii; A. evermanni Jordan et Snyder est très répandu dans l Indo-Pacifique et également connu de l Atlantique occidental; A. kallopterus Bleeker, A. taeniopterus Bennett (synonyme: A. menesemus aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids Jenkins), Apogonichthys perdix Bleeker (synonyme: A. waikiki Jordan et Evermann), Foa brachygramma (Jenkins) et Lachneratus phasmaticus Fraser et Struhsaker sont des espèces indo-pacifiques à large rèpartition. Sommario Le 10 specie di pesci apogonidi delle isole Hawaii sono differenti in una cosa. I Pseudamiops diaphanes, nuova specie endemica delle Hawaii, si distingue per il possesso di VI-l,8 raggi dorsali, II, 8 raggi anali, 16-18 raggi pettorali, una proiezione addominale tipo spina posteriore sulla scala cicloide posteriore della maxilla, senza linee lateriali, una spina caudale romboide e nell essere trasparente durante la sua vita. L Apogon deetsie, nuova specie, delle isola Hawaii e l Arcipelago Tuamotu ha Vi-I,9 raggi dorsali, II,8 raggi anali, 11-12 (solitamente 12) raggi pettorali, 24; squame sulla linea laterale, 6 squame predorsali, 3-4 + 12-13 spine branchiali, un lembo di membrana sporgente sull angolo e l estremità inferiore del preopercle, una spina caudale biforcuta con lobi rotondi e è rosso chiaro con due strisce nerastre sulla parte posteriore del corpo. Le specie restanti vengono analizzate e ne viene segnalata la diffusione. L Apogon erythrinus Snyder sembra essere endemico delle Hawaii e delle isole Johnston; l A. maculiferus Garret è anche endemica delle Hawaii; l A. evermanni Jordan e Snyder è diffuso nell indo-pacifico e e viene conosciuto anche nell Atlantico occidentale; l A. kallopterus Bleeker, l A. taeniopterus Bennett (A. menesemus Jenkins è un sinonimo), l Apogonichthus perdix Bleeker (A. waikiki Jordan e Evermann è un sinonimo), il Foa brachygramma (Jenkins), e Lachneratus phasmaticus Fraser e Struhsaker sono specie molto diffuse in Indo-pacifico. Introduction The Apogonidae, popularly known as cardinalfishes, are a large family of small fishes well-represented in tropical to warm temperate seas from a variety of habitats. The species are nocturnal, in general, and among the few marine fishes which are mouthbrooders. The male retains the fertilized eggs within the mouth until hatching (however, there are also reports of female apogonids as mouthbrooders). Fishes of this family are distinct in having two dorsal fins, the first with VI to VIII spines, the second with a single spine and 8-14 soft rays; an anal fin with II spines and 8-18 soft rays; large eyes; a large oblique mouth; a single opercular spine, and the edge of preopercle preceded by a parallel ridge. Fraser (1972) reviewed the classification of the family from the subfamily to the subgeneric level. He recognized three subfamilies: Apogoninae, Pseudaminae, and Epigoninae; the last-mentioned taxon is now given family status. The Hawaiian Islands have relatively few apogonid fishes compared to most other areas of the Indo- Pacific region. Gosline and Brock (1960) recognized only eight species (not counting Epigonus atherinoides and Synagrops argyrea, the latter placed in the Pericichthyidae by Fraser, 1972). Fraser and Struhsaker (1991) described Lachneratus phasmaticus as a new genus and species of apogonid fish from the Hawaiian islands (type locality), Fiji, and the Comoro Islands. The small number of apogonid species for the islands is undoubtedly related to the relatively short larval life in the plankton due to the early development taking place within the mouth of the brooding adult fish. Chave (1979) provided a detailed analysis of the general ecology of the six most common apogonid fishes of the Hawaiian Islands. In the present paper two of the species names used by Gosline and Brock (1960) are changed, the species identified by them as Pseudamiops gracilicauda (Lachner) is described as new, a new species of Apogon is described, and a revised key to the Hawaiian species of the family is provided. Materials and Methods Type specimens of the new species have been deposited in the Australian Museum, Sydney (AMS); Natural History Museum, London (BMNH); Bernice P. Bishop Museum, Honolulu (BPBM); California Academy of Sciences, San Francisco (CAS); and the U.S. National Museum of Natural History, Washington, D.C. (USNM). Lengths recorded for specimens are standard length (SL), the straight-line distance from the tip of the snout in the median plane to the base of the caudal fin (end of hypural plate). Body depth is the greatest depth; body width is the greatest width just posterior to the head. Head length is measured from the front of the upper lip in the median plane (with mouth fully closed) to the most posterior point of the opercular membrane; snout length is taken from the same anterior point to the fleshy edge of the orbit. Eye diameter is the greatest diameter of the eye ball (this can be more accurately measured than the fleshy orbit diameter); interorbital width is the least fleshy width. Caudal-peduncle depth is the least depth, and caudalpeduncle length the horizontal distance between verticals at the rear base of the anal fin and the base of the caudal fin; lengths of spines and rays of fins are measured from their extreme bases in a straight line to their tips. Caudal concavity is the horizontal distance between verticals at the tips of the longest and shortest caudal rays. Pectoral-fin length is the length of the longest ray; pelvic-fin length is measured from the base of the spine to the tip of the longest ray. Counts of pectoral rays include the splint-like uppermost ray. Counts of lateral-line scales are made to the base of the caudal fin (hence do not include one aqua vol. 3 no. 1-1998 26

John E. Randall or two pored scales that extend onto the base of the fin). Often a pored scale lies directly over the end of the hypural plate, hence partly onto the base of the caudal fin; this scale is then included in the count of lateral-line scales. Gill-raker counts include all rudiments; the count of lower-limb rakers contains the long raker at the angle. Developed gill rakers are those which are higher than the diameter of their base. Meristic and morphometric data given in parentheses refer to paratypes. Tables of measurements of type specimens are presented as percentages of the standard length. Proportional measurements in the text are rounded to the nearest 0.05. Key to the Hawaiian Species of Apogonidae 1a. Dorsal rays VI-I,8; pectoral rays 16-18; a sharp, downward-projecting, bony process posteriorly on ventral edge of maxilla; body very elongate, the depth 4.25-4.8 in SL; caudal fin rhomboid (Hawaiian Islands)......Pseudamiops diaphanes, new species 1b. Dorsal rays VI or VII, 9-13; pectoral rays 10-14; no sharp bony process posteriorly on ventral edge of maxilla; body not very elongate, the depth 2.3-3.9 in SL; caudal fin rounded or forked...2 2a. Dorsal soft rays 12-13; anal soft rays 13-16; pectoral rays 10; edge of preopercle smooth with 2 or 3 acute spines at corner; scales cycloid; no lateral line (Hawaiian Islands, Fiji, and Comoro Islands)...Lachneratus phasmaticus 2b. Dorsal soft rays 9; anal soft rays 8; pectoral rays 11-15; edge of preopercle smooth or serrate, without acute spines at corner; scales ctenoid; lateral line present...3 3a. Caudal fin rounded; edge of preopercle smooth; supramaxilla present; predorsal scales 3-4...4 3b. Caudal fin forked; edge of preopercle serrate; supramaxilla absent; predorsal scales 5-12...5 4a. Lateral line complete; palatine teeth absent; pectoral rays 14; body not deep, the depth 2.5-2.9 in SL (Indo-Pacific)...Apogonichthys perdix 4b. Lateral line interrupted, the anterodorsal part ending beneath second dorsal fin; palatine teeth present; pectoral rays 12; body deep, the depth 2.2-2.5 in SL (Indo-Pacific)...Foa brachygramma 5a. Scales of lateral line about twice as large as scales of rest of body; predorsal scales 10-12; red with a blackish stripe on head from front of snout through eye to end of opercle, and a large blackish spot at rear base of second dorsal fin with a small white spot adjacent and posterior to it (Indo- Pacific and western Atlantic)...Apogon evermanni 5b. Scales of lateral line not larger than scales of rest of body; predorsal scales 5-6; color not as in 5a...6 6a. Spines of first dorsal fin VI; color in life red...7 6b. Spines of first dorsal fin VII; color in life not red (may be orangish)...8 7a. Pectoral rays 14-15; scales above lateral line to origin of dorsal fin 3; circumpeduncular scales 16; color in life transparent red with no distinct dark markings (Hawaiian Islands)......Apogon erythrinus 7b. Pectoral rays 11-12; scales above lateral line to origin of dorsal fin 1; circumpeduncular scales 12; color in life light red with 2 broad b l a c k i s h bars, one between posterior halves of second dorsal and anal fins and the other posteriorly on caudal peduncle (Hawaiian Islands and Tuamotu Archipelago)...Apogon deetsie, new species 8a. Pectoral rays usually 14; preopercular ridge smooth or with a few small serrae; third dorsal spine clearly longest; orangish with a small distinct dark brown spot on scales of body forming longitudinal rows; no black stripe in second dorsal and anal fins, and no black anteriorly on first dorsal in (Hawaiian Islands)......Apogon maculiferus 8b. Pectoral rays usually 13; preopercular ridge strongly serrate; third and fourth dorsal spines subequal; no rows of distinct small dark brown spots on body (diffuse dark blotches may be present on body of A. taeniopterus); a black stripe in second dorsal and anal fins near base; first dorsal fin with a black anterior border... 9 9a. Gill rakers 17-20; circumpeduncular scales 12-14; longest dorsal spine 1.7-1.95 in head length; a midlateral dark brown stripe on head and body (may be diffuse or absent on body of large specimens); a dark brown spot at caudal-fin base; no black bar in caudal fin (Indo-Pacific)......Apogon kallopterus 9b. Gill rakers 21-23; circumpeduncular scales 16-18; longest dorsal spine 1.95-2.3 in head length; no dark brown stripe on body; no dark brown spot at base of caudal fin; a black bar in caudal fin about one-third distance from base, linked to a black submarginal band in each lobe of fin (Indo- Pacific)...Apogon taeniopterus Pseudaminae Pseudamiops diaphanes, sp. n. Transparent Cardinalfish (Figs. 1 & 2) Pseudamiops gracilicauda (Fig. 3) (non Lachner) Gosline, 1955: 450 (Johnston Island and Hawaiian Islands); Gosline and Brock, 1960: 162, fig. 179; Randall et al., 1985: 54 (Johnston Island). Holotype: BPBM 37247, female, 30.2 mm, Hawaiian Islands, Oahu, Makaha, Kepuhi Point, cave at shore, 3 m, hand net at night, J. L. Earle, 6 June 1991. Paratypes: BPBM 37412, 3: 24.4-26.0 mm, Hawaiian 27 aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids Fig 1. Holotype of Pseudamiops diaphanes, BPBM, 30.2 mm SL, Makaha, Oahu. Fig 2. Underwater photograph at night of Pseudamiops diaphanes, about 40 mm TL, Makaha, Oahu. Dorsal rays VI-I,8; anal rays II,8; pectoral rays 16-18; scales thin, cycloid, highly deciduous, about 23-24 in longitudinal series, lateral line absent; a sharp, downward-projecting, bony process posteriorly on ventral margin of maxilla; body very elongate, the depth 4.25-4.8 in SL; preopercular margin serrate; preopercular ridge smooth; caudal fin rhomboid; transparent in life. Description Fig 3. Underwater photograph at night of Pseudamiops gracilicauda, about 45 mm TL, Kwajalein Atoll, Marshall Island. Islands, Oahu, Waimea coast, rotenone, V.E. Brock, W. A. Gosline et al., 25 July 1955 (formerly University of Hawaii collection); BPBM 7913, 2: 16.9-22.6 mm, Oahu, Moku Manu, cave in 23 m, rotenone, J.E. Randall, W. J. Baldwin, and G. S. Losey, 3 October 1969; CAS 91613, 25.2 mm, Oahu, Waimea Bay, SW end, cave in 10.5 m, rotenone, J. E. Randall, L. A. Randall, and P. M. Allen, 27 July 1970; BPBM 37221, 30.3 mm, Oahu, Makaha, cave, 18 m, hand net at night, J. L. Earle, 10 October 1990; BPBM 37401, 17.8 mm, same data as holotype; BPBM 37252, 32.8 mm, Oahu, Makaha, hand net at night, J. L. Earle, July, 1991; USNM 343981, 28.8 mm, Oahu, Makaha, Kepuhi Point, cave at shore, hand net at night, 3 m, J. L. Earle, 5 July 1991; AMS I.37827-001, 31.0 mm, Oahu, off main channel to Kaneohe Bay, base of drop-off, 25 m, D.W. Greenfield, J. E. Randall, G. Cockrell, and J. L. Mahon, 30 August 1991; BMNH 1997.2.4.1, 24.6 mm, Northwestern Hawaiian Islands, Midway Atoll, Sand Island, SE side, 200 m SW of bulk garbage pier, outside reef, cave, 8 m, rotenone, J. E. Randall, J. L. Earle, T. Hayes, and R. L. Pyle, 16 September l991; BPBM 37413, 6: 18.5-25.3 mm, Oahu, off Kaneohe Bay, Sampan Channel, spur and groove zone, 6-7 m, rotenone, D.W. Greenfield and K. R. Longenecker, 1 November 1993. Dorsal rays VI-I,8; anal rays II,8; all dorsal and anal soft rays branched, the last to base; pectoral rays 17 (two paratypes with 16, four with 18, 11 with 17), the upper and lower 3-5 unbranched; pelvic rays I,5; principal caudal rays 17, the upper and lower unbranched; upper procurrent caudal rays 7 (6-8), the posterior 2 segmented; lower procurrent caudal rays 6 (6-7), the posterior 2 segmented; gill rakers 16, with 1 upper and 7 lower developed (15-17, with 1 upper and 6-7, usually 7, developed); pseudobranchial filaments rudimentary; branchiostegal rays 7; vertebrae 10 +14; no supraneural (predorsal) bones. Body very elongate, the depth 4.65 (4.25-4.8) in SL, the greatest depth just posterior to head (except in fully ripe females); body moderately compressed, the width 2.0 (1.9-2.35) in depth; head length 2.95 (2.75-2.95) in SL; snout length 3.65 (3.45-3.85) in head; eye diameter 4.3 (4.35-5.0) in head; interorbital width 3.4 (3.1-3.65) in head; caudal peduncle slender and long, the depth 3.15 (2.9-3.5) in head, the length 3.4 (3.4-3.7) in SL. Mouth slightly inferior and large, the maxilla extending slightly posterior to orbit, the upper-jaw length 1.7 (1.75-1.85) in head; mouth slightly oblique, the gape forming an angle of about 20 to horizontal axis of body; a sharp, spine-like, downward-projecting, bony process posteriorly on ventral margin of maxilla; no supramaxilla; upper jaw with 1-3 irregular rows of villiform teeth; 2 (1-2) large, nearly sessile, inward-projecting canine teeth; medial to villiform teeth on each aqua vol. 3 no. 1-1998 28

John E. Randall side of symphysis of upper jaw; lower jaw with an irregular outer row of small conical teeth and an inner row of small, slender, well-separated, incurved canine teeth, those in middle of side of jaw largest (largest about half pupil diameter); a single row of small conical teeth on palatines, and a double row forming a V on vomer. Anterior part of tongue very slender. Longest gill raker at angle, its length in holotype 1.75 in eye diameter. A single, flat, flexible, right-angle opercular spine; edge of preopercle serrate, with 14 (11-13) irregular serrae, the largest at angle; preopercular ridge smooth; rows of small papillae on head forming a crosshatch pattern, with 11 transverse rows across the top of the head and continuing downward on side of head (those posterior to preopercle indistinct); five short rows extending ventrally from eye (four of the five oblique), and six across mandible; two longitudinal rows dorsally on snout and extending through interorbital space; five longitudinal rows across preopercle, and one along mandible. Anterior nostril a short flaccid tube in front of center of eye at edge of groove separating snout from upper lip; posterior nostril a large opening just in front of eye and only slightly dorsal to anterior nostril; nasal chamber cavernous, the skin over the roof thin and sagging inward. Scales cycloid, thin, and very easily shed (few scales now present on any type specimens, hence the count of 23 or 24 scales in longitudinal series is only approximate); large scales present on opercle, but none now apparent on preopercle or elsewhere on head. No lateral line evident. Origin of first dorsal fin slightly posterior to base of middle pectoral rays, the predorsal distance 2.6 (2.6-2.65) in SL; fin spines slender and flexible; first dorsal spine 2.7 (2.9-3.5) in head; third dorsal spine longest, slightly longer than second spine, 2.45 (2.45-2.7) in head; second dorsal fin well-separated from first dorsal fin, its origin slightly anterior to anal-fin origin; spine of second dorsal fin 3.05 (3.05-3.85) in head; second dorsal soft ray longest, 1.8 (1.65-2.0) in head; first anal spine short, 9.0 (6.5-10.1) in head; second anal spine 2.9 (3.0-3.5) in head; second anal soft ray longest, 1.75 (1.6-1.9) in head; caudal fin rhomboid, 3.35 (3.05-3.4) in SL; pectoral fins long and pointed, the eighth or ninth ray longest, 3.9 (3.25-4.0) in SL; pelvic fins just reaching anus, 2.05 (2.0-2.15) in head. Color of holotype in preservative: pale with dusky spots on lips (from individual melanophores), a single melanophore ventroposteriorly at edge of eye, a double row of melanophores midventrally on abdomen, and a dusky bar basally on caudal fin (this marking absent on some paratypes, prominent and broader on another). Color of holotype when fresh: transparent with dark markings as noted in preserved specimen; body cavity opaquely covered with blackish red; vertebral column reddish black; gill filaments showing through operculum as bright red. Life color from an underwater photograph, shown in Fig. 2. Etymology This species of Pseudamiops is named diaphanes from the Greek for transparent. Pseudamiops diaphanes is known only from the Hawaiian Islands and Johnston Atoll (Gosline, 1955 reported a 23-mm specimen from the atoll as Pseudamiops gracilicauda). An uncatalogued lot of six specimens, 17-26 mm SL, collected by V. E. Brock et al. at Johnston in 1963, was found in the reference fish collection of the University of Hawaii. This lot is now BPBM 37413. The specimens are in poor condition, therefore not designated as paratypes. All Hawaiian specimens of P. diaphanes were collected at the island of Oahu except for one from Midway. The species may be expected at the other islands of the archipelago. This cardinalfish has been observed alive only at night and only in caves where it is generally seen well above the substratum. Rotenone collections during the day have resulted in specimens from the depth range of 3-25 m, all from caves, except one in a spur and groove zone. The holotype, the 24.6-mm paratype, the 31-mm paratype, and a 32.8-mm specimen collected in July, 1991 (BPBM 37252, not recorded as a paratype due to bad condition) are fully mature females. Pseudamiops diaphanes is most closely related to P. gracilicauda, described from the Marshall Islands by Lachner in Schultz and collaborators (1953: 497, fig. 84) from six specimens, 13.5-33.5 mm SL. P. gracilicauda has since been reported from the Samoa Islands by Wass (1984: 14) and from Rapa by Randall et al. (1990: 19, fig. 24). It differs from P. diaphanes in having the edge of the preopercle smooth with two stout serrae at the angle (edge fully serrate in diaphanes), in having 15-16 pectoral rays (16-18 in diaphanes; only two of 18 specimens with 16 rays), and in color. P. gracilicauda has a dusky blotch on the preopercle behind the eye and a sprinkling of melanophores along the base of the dorsal and anal fins; these markings are not apparent on diaphanes. The edges of the scales of gracilicauda are faintly red in life, but not in diaphanes. Lachner gave the gillraker count of gracilicauda, including rudiments, as 3 + 8 which would seem to be another meristic difference. However, the gill-raker count of Lachner s holotype was checked by Jeffrey T. Williams who corrected it to 4 + 11 (Randall et al., 1990). Williams also determined that the holotype has 1 weak opercular spine, not 2 as recorded by Lachner, and 2 stout serrae at the angle with an additional small one. If the Rapa and Samoa specimens are correctly identified as 29 aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids gracilicauda, this species appears to attain a larger size than diaphanes, the largest specimen of which measures 32.8 mm SL. Randall et al. (1990) reported five Bishop Museum specimens from Rapa, 35-50 mm SL. One specimen from Tutuila, American Samoa (BPBM 17501) measures 41 mm. Figure 3 is an underwater photograph of P. gracilicauda taken in 7 m at night at Kwajalein Atoll, Marshall Islands by the author; the fish was estimated to be 45 mm in total length. Lachner described gracilicauda in the genus Gymnapogon, but he did so tentatively because of the poor condition of the type specimens. One character favoring the placement by him in Gymnapogon was the apparent lack of scales. Although Smith (1954: 784) doubted that gracilicauda lacked scales, he wrote, If it should, however, prove that gracilicauda Lachner is truly scaleless, it unquestionably merits generic distinction from Gymnapogon, and the new genus Lachneria is provisionally proposed, its type gracilicauda Lachner, 1953. Gosline (1955) correctly concluded that Lachner s specimens of gracilicauda were scaled but lost them due to their deciduous nature; therefore there was no need to recognize Lachneria. Smith (1954: 783) described the new genus Pseudamiops, with P. pellucidus Smith (1954: 785, fig. 2, Pl. XXIII,A-C) from Bazaruto, Mozambique as the type species; he included gracilicauda in this genus. P. pellucidus is clearly distinct from gracilicauda and diaphanes in having 9 anal soft rays, 14 pectoral rays, about 33 scales in longitudinal series, and prominent canines on the side of the upper jaw. Apogoninae Apogon deetsie, sp. n. Deetsie s Cardinalfish (Fig. 4) Apogon sp. Madden, 1973: 145 (off Makapuu Point, Oahu). Holotype: BPBM 37400, male, 40.4 mm, Hawaiian Islands, Oahu, off entrance to channel at NE end of Kaneohe Bay, small cave, 29 m, rotenone, J. E. Randall, E. H. Chave, and W. Hashimoto, 10 October 1969. Paratypes: BPBM 10217, 32.8 mm, Hawaiian Islands, Oahu, off Makapuu Point, one-fourth mile E of Manana Island (Rabbit Island), from bilge of a boat raised from an initial depth of 160 m (boat sank on June 4, 1970; on June 20 it was dragged to a depth of 68 m; on October 14 it was dragged farther inshore and raised to the surface), W.D. Madden et al.; BPBM 14004, 26.6 mm, Tuamotu Archipelago, Rangiroa Atoll, outside reef 1 mile E of Avatoru Pass, 60 dropoff, sand and coral, 46-53 m, rotenone, J. E. Randall, D. B. Cannoy, R.M. McNair, and D. Bryant, 9 April 1971 (fins damaged); BPBM 13983, 59.7 mm, Oahu, off Kaneohe Bay, under ledge in 26 m, rotenone, W. D. Madden and R. J. Shallenberger, 2 July 1971; USNM 343982, 37.0 mm, Oahu, Makaha, under ledge, 44 m, rotenone, J. E. Randall, L. R. Taylor, and P. M. Allen, 16 June 1973; BMNH 1997.2.4.2, 38.7 mm, Oahu, off Kaneohe Bay, near entrance to ship channel at NW end of bay, cave at base of drop-off, 25 m, rotenone, D. W. Greenfield, K.R. Longenecker, and J. Mendez, 7 September 1995; CAS 91614, 40.7 mm and AMS I.37828-001, 50.9 mm, same data as preceding. Fig 4. Holotype of Apogon deetsie, BPBM 37400, 40.4 mm SL, off Kaneohe Bay, Oahu. Dorsal rays VI-I,9; anal rays II,8; pectoral rays 12 (rarely 11); lateral-line scales 24; predorsal scales 6; gill rakers 3-4 + 12-13; body depth 2.9-3.15 in SL; upper preopercular edge serrate, the corner and lower edge with a protruding membranous lobe; preopercular ridge smooth; caudal fin forked with rounded lobes; color light red with two dusky bars posteriorly on body. Description Dorsal rays VI-I,9; anal rays II,8; all dorsal and anal soft rays branched, the last to base; pectoral rays 12 (seven paratypes with 12, one with 11), the upper and lower 2 unbranched; pelvic rays I,5; principal caudal rays 17, the upper and lower unbranched; procurrent caudal rays 6 (6-7, usually 6), the most posterior segmented; lateral line complete, the pored scales 24 (plus 2 smaller pored scales on caudal-fin base); predorsal scales 6; scales above lateral line to origin of first dorsal fin 1; scales below lateral line to origin of anal fin 7; circumpeduncular scales 12; gill rakers 3 + 12, the upper 2 and lower 11 developed (4 + 12-13, the upper 3 and lower 11 or 12 developed); pseudobranchial filaments 8 (8-11); branchiostegal rays 7; vertebrae 10 + 14; supraneural (predorsal) bones 2. Body depth 3.1 (2.9-3.15) in SL; body moderately compressed, the width 1.9 (1.75-2.1) in depth; head length 2.5 (2.45-2.55) in SL; dorsal profile of head aqua vol. 3 no. 1-1998 30

John E. Randall slightly convex to straight (except for rounded front of snout); snout length 4.1 (3.85-4.05) in head; eye diameter 3.0 (2.8-3.15) in head; interorbital width 4.15 (4.05-4.66) in head; caudal-peduncle depth 2.7 (2.55-2.7) in head; caudal-peduncle length 3.5 (3.3-3.5) in SL. Mouth slightly inferior and large, the maxilla extending nearly to a vertical at posterior edge of orbit, the upper-jaw length 1.85 (1.75-1.9) in head; mouth slightly oblique, the gape forming an angle of about 20 to horizontal axis of body; supramaxilla not present; a band of villiform teeth in jaws (maximum of about 10 rows in upper jaw and 5 in lower of holotype); a single row of very small conical teeth on palatines (15 teeth in row of holotype); small conical teeth forming a V-shaped patch on vomer, in 1 to 2 rows anteriorly and 3 rows posteriorly (vomerine teeth larger than those in jaws). Anterior part of tongue slender and spatulate with papillae on the upper surface. Longest gill rakers at angle and adjacent to angle on lower limb, its length half eye diameter in holotype. A single, acute, sharp, opercular spine at level of center of eye; upper edge of preopercle serrate, with 18 (11-21) serrae; corner and ventral part of preopercular edge with a large membranous flap which extends slightly posterior to serrate edge; lateralis system of head with numerous close-set pores having slightly elevated rims; 2 longitudinal rows of pores from front of snout to occiput, crossed by 27 (26-30) transverse rows; two suborbital rows of pores with the same transverse pore system. Anterior nostril a short membranous tube slightly below center of eye about a nostril diameter from groove at base of upper lip; posterior nostril a large vertically ovate aperture directly in front of center of eye, the internarial distance about half pupil diameter; nasal chamber cavernous, the skin over the roof thin. Scales weakly ctenoid and thin; scales present on nape, opercle, and preopercle, those on opercle larger than scales of body; no scales on occiput, interorbital, snout, or ventrally on head; no scales on fins except for small scales on about basal fourth of caudal fin and a scaly process of 2 large scales extending midventrally from base of pelvic fins to slightly more than half length of pelvic spines. Origin of first dorsal fin over third lateral-line scale, the predorsal distance 2.35 (2.25-2.4) in SL; fin spines strong and sharp; first dorsal spine 4.25 (4.1-4.35) in head; second dorsal spine longest, 2.45 (2.45-2.7) in head; second dorsal fin separated by a median scale (indented posteriorly) from first dorsal fin, its origin above ninth lateral-line scale; spine of second dorsal fin abnormally short in holotype (2.5-2.8 in head of paratypes); second dorsal soft ray longest, 1.45 (1.5-1.55) in head; origin of anal fin below or slightly posterior to origin of second dorsal fin; first anal spine short, 8.6 (7.9-9.35) in head; second anal spine 2.5 (2.3-2.55) in head; second anal soft ray longest, 2.5 (2.3-2.55) in head; caudal fin forked with broadly rounded lobes, 3.1 (2.8-3.15) in SL; caudal concavity 3.45 (2.75-3.25) in head; pectoral fins long and pointed, the seventh ray longest, 3.55 (3.25-3.6) in SL; pelvic fins nearly or just reaching origin of anal fin, 1.55 (1.5-1.65) in head. Color of holotype in alcohol: pale yellowish with a faint dusky bar below basal half of second dorsal fin and a broader bar posteriorly on caudal peduncle and extending slightly onto base of caudal fin; fins pale except about outer fourth of caudal lobes which is dusky. Color of holotype when first collected: body translucent, the edges of scales broadly light red; two blackish bars posteriorly on body as described above; head light red with an iridescent area of light yellow and light blue over operculum; iris pale yellowish; fins transparent with pale pink rays except about outer fourth of caudal lobes which is blackish and tips of second dorsal and anal fins which are dusky. Etymology This species of Apogon is named deetsie in honor of E. H. Deetsie Chave, one of the collectors of the holotype, in recognition of her research on the ecology of apogonid fishes of the Hawaiian Islands. The specific name is treated as a noun in apposition. Apogon deetsie is known only from the Hawaiian Islands and Rangiroa Atoll in the Tuamotu Archipelago. All specimens from the Hawaiian Islands were collected off Oahu except for one of 62 mm which was taken by trawl by Paul J. Struhsaker from the R/V. Townsend Cromwell off the north shore of Maui (20 57.5 N, 156 28.6 W) in 66-84 m; the trawl was towed between 11 and 12 p.m on 7 June 1970. The specimen was deposited at the Honolulu Laboratory of the National Marine Fisheries Service but later discarded due to its very poor condition. Meristic data were taken by E. H. Chave. This species occurs in moderately deep water, sometimes deeper than normal SCUBA-diving depths, hence the probable reason for the few specimens that have been collected. Apogon deetsie is most closely related to an undescribed species represented by a single specimen collected by the author and Gerald R. Allen in 18-37 m at Florida Island, Solomon Islands on 30 July 1973. This species has the same two broad blackish bars posteriorly on the body, but the ground color in life was light yellowish, and there is no dark pigment on the median fins. It differs further in having 13 pectoral rays, 5 + 14 gill rakers, and in lacking a broad membranous flap at the corner and lower margin of the preopercle. 31 aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids Six species of Apogon with two blackish bars posteriorly on the body occur in the tropical western Atlantic. Böhlke and Randall (1968: fig. 2) illustrated the barred pattern of five of these, and Bö hlke and Chaplin (1968: 240) the sixth. In view of the occurrence of Apogon evermanni in the western Atlantic and the Indo-Pacific, A. deetsie was compared with these six Atlantic species. A. phenax Böhlke and Randall (1968: fig. 5; illustrated in color by Randall, 1996a: 124) appears to be the closest relative among the Atlantic species. It differs from A. deetsie in the more posterior position and usual wedge-shape of the first blackish bar on the body, 2 scales above the lateral line to the origin of the dorsal fin, usually 7 predorsal scales, 5 + 13-14 gill rakers, and no protruding membranous lobe on the corner and ventral margin of the preopercle. A. planifrons Longley and Hildebrand (illustrated by Böhlke and Chaplin, 1968: 245) is also similar; it differs in having more pointed caudal lobes, 2 scales above the lateral line to the origin of the dorsal fin, 6 + 16-17 gill rakers, and no lower membranous preopercular lobe. Apogon erythrinus Snyder, 1904 Hawaiian Ruby Cardinalfish (Fig. 5) Apogon erythrinus Snyder, 1904: 526, pl. 9, fig. 17 (type locality, Puako Bay, Hawaii). Amia erythrina Jordan and Evermann, 1905: 217; pl. 34 (as Apogon erythrinus). Fig. 5. Underwater photograph at night of Apogon erythrinus, about 45 mm TL, Black Rock, West Maui. Dorsal rays VI-I,9; anal rays II,8; pectoral rays 13-15 (one of 20 with 13 and one with 15); lateral-line scales 24; scales above lateral line to base of first two dorsal spines 3; predorsal scales 5 or 6; circumpeduncular scales 16; gill rakers 3-4 + 11-13 (one of 20 with 4 and one with 13); upper margin of preopercle serrate; corner and lower edge of preopercle membranous and crenate but not protruding posteriorly; preopercular ridge smooth; body depth 2.5-2.8 in SL; head length 2.35-2.6 in SL; caudal-peduncle length 3.65-3.95 in SL; second dorsal spine longest, 1.45-1.75 in head; caudal fin forked with rounded lobes; transparent red; blackish pigment on occiput, opercle, and as a small blotch below anterior part of first dorsal fin. Attains about 45 mm SL. Apogon erythrinus appears to be endemic to the Hawaiian Islands and Johnston Island. Hayashi and Kishimoto (l983) have recorded it from the Ryukyu Islands and the Philippines; however, these records need confirmation. Apogon erythrinus is a close relative of A. crassiceps Garman, 1903 (type locality, Fiji), widely distributed in the Indo-Pacific except the Arabian region (Red Sea to Persian Gulf) where another close relative, A. coccineus Rüppell,1838, is found. Randall et al., 1990 reported on the examination of the holotype of Apogon crassiceps (MCZ 28214, 27 mm SL). They concluded that it is the small, wideranging, transparent red species that has been confused in the past with A. erythrinus and A. coccineus. The species that Jordan and Seale (1906: 239, fig. 32), Masuda et al. (1984: 149, pl. 132-L), and Winterbottom et al. (1989: 29, fig. 159) identified as A. crassiceps is the one Smith (1961: 387, pl. 47 A) described as A. talboti. Apogon crassiceps differs from A. erythrinus in usually having 13 pectoral rays, 2 scales above the lateral line to the base of first two dorsal spines, and a more elongate body on average (body depth 2.65-3.2 in SL). The Bishop Museum has numerous specimens of Apogon crassiceps from Rapa (as reported by Randall et al., 1990), Pitcairn Group, Society Islands, Tuamotu Archipelago, Marquesas Islands, Line Islands, Phoenix Islands, Caroline Islands, Kiribati, Marshall Islands, Minami-tori-shima (Marcus Island), Fiji, New Caledonia, Lord Howe Island, Great Barrier Reef, Solomon Islands, Indonesia, Philippines, Ryukyu Islands, Ogasawara Islands, Sri Lanka, Seychelles, and Mauritius. A. coccineus differs in having 3-5 + 13-15 gill rakers and a prominent, broad, longitudinal, blackish streak on the side of the caudal peduncle. Apogon erythrinus is a shallow-water, coral-reef species that has been observed alive only at night, close to the bottom, and never far from shelter. Chave (1979) reported not seeing it more than 3 cm from the substratum. It is a difficult fish to photograph in the wild because it quickly retreats to a hole in the reef when a light is shone in its direction. Chave noted that only Apogonichthys perdix among the six common Hawaiian apogonids is more sensitive to light. She found only crustaceans in the stomachs of 68 specimens examined from Oahu. Hobson (1974) found food in eight specimens from Kona, Hawaii, all crustaceans, of which xanthid crabs were the major prey. aqua vol. 3 no. 1-1998 32

John E. Randall Apogon evermanni Jordan and Snyder, 1904 Oddscale Cardinalfish (Fig. 6) Apogon evermanni Jordan and Snyder, 1904: 123 (type locality, Honolulu). Amia evermanni Jordan and Evermann, 1905: 213, fig. 84. Apogon anisolepis Böhlke and Randall, 1968: 182, fig. 1 (type locality, Green Cay, Bahamas). Fig 6. Underwater photograph of Apogon evermanni, about 110 mm TL, cave in Kona, Hawaii. Dorsal rays VI-I,9; anal rays II,8; pectoral rays 12; lateral-line scales 24; remaining scales of body small; scales above lateral line to origin of dorsal fin 3-4; scales below lateral line to origin of anal fin 19-21; predorsal scales 10-12; circumpeduncular scales 33-35; gill rakers 4-6 + 15-17; upper edge of preopercle finely serrate, the corner and lower edge smooth, membranous, and not extending posterior to serrate margin; preopercular ridge smooth; body moderately elongate, the depth 2.7-3.7 in SL; head length 2.3-2.7 in SL; dorsal profile of head straight except for rounded front of snout; caudal peduncle long, 3.2-4.2 in SL; second and third dorsal spines subequal, 2.2-3.2 in head; caudal fin forked with rounded lobes; red in life with a blackish stripe from front of snout, through eye, to end of opercle, a black spot at rear base of second dorsal fin and an adjacent small white spot at base of last dorsal ray and onto caudal peduncle. Smaller individuals tend to be paler red; still smaller ones translucent. Largest specimen examined, 121.5 mm. Böhlke and Randall (1968) described Apogon anisolepis from one specimen from the Bahamas and one from Curação, never suspecting that the species had been described from the Hawaiian Islands as A. evermanni in 1904. That realization came in 1969 when the author, S. N. Swerdloff, and E. H. Chave collected a specimen of A. evermanni (BPBM 7826, 102.3 mm) off Pokai Bay, Oahu. Randall and Böhlke (1981) placed A. anisolepis in the synonymy of A. evermanni and recorded the latter from the following localities in the Indo-Pacific, in addition to Hawaii: Marquesas Islands, Marshall Islands, Solomon Islands, Molucca Islands, Cocos-Keeling Islands, Maldive Islands, Chagos Archipelago, and Mauritius. They added Cozumel Island, Mexico as an additional record for the western Atlantic. A. evermanni is unknown from the Red Sea and Gulf of Aden; it appears to be replaced there by A. isus Randall and Böhlke. This species is very similar in color and morphology to A. evermanni but differs notably in having the scales of the body equal in size to the lateral-line scales. Randall and Böhlke (1981: Table l) noted that the number of scales below the lateral line and the number of circumpeduncular scales of Apogon evermanni is highest in the Hawaiian Islands and Marquesas, intermediate in the western Pacific and Indian Ocean, and lowest in the western Atlantic. Apogon evermanni occurs in clear-water reef areas of full salinity. It is well hidden in caves or beneath ledges by day, and even at night is rarely seen outside of caves. Like A. erythrinus, it is very difficult to approach with a light at night. Specimens have been collected in the depth range of 3-69 m, usually in more than 25 m. Apogon kallopterus Bleeker, 1856 Iridescent Cardinalfish (Fig. 7) Apogon kallopterus Bleeker, 1856: 33 (type locality, Manado, Sulawesi). Amia kallopterus Bleeker, 1873-1876: 89; Bleeker, 1876-1877: pl. 59, fig. 2. Apogon snyderi Jordan and Evermann, 1903: 180 (type locality, Honolulu); Jordan and Evermann, 1905: 214, fig. 85, pl. 36. Dorsal rays VII-I,9; anal rays II,8; pectoral rays 13-14 (rarely 14); lateral-line scales 25; scales above lateral Fig 7. Underwater photograph of Apogon kallopterus, about 80 mm TL, Oahu. 33 aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids line to origin of dorsal fin 2; median predorsal scales 5-6 (usually 6); circumpeduncular scales 12-14 (rarely 12, usually 14); gill rakers 4-5 + 13-15; edge of preopercle and preopercular ridge serrate; body depth 2.5-3.3 in SL; head length 2.4-2.55 in SL; dorsal profile of head convex; caudal-peduncle length 3.65-4.35 in SL; third and fourth dorsal spines subequal, 1.7-1.95 in head; caudal fin forked, with slightly rounded lobes; color in life light brown, the scale edges darker, with blue-green and yellow iridescence (especially at night); a dark brown stripe from front of snout to eye, continuing more broadly and diffusely on head behind eye and along side of body (stripe on body absent or nearly so on large individuals); a dark brown spot about size of pupil posteriorly on caudal peduncle, its lower edge on lateral line; leading edge of first dorsal fin broadly dusky to dark brown; a dark brown to dusky band basally in second dorsal and anal fins, on second pelvic ray, and submarginally in caudal-fin lobes. Largest specimen reported by Fraser and Lachner (1985), 122 mm SL. Apogon kallopterus is a wide-ranging Indo-Pacific species, occurring from the Red Sea to South Africa, east to the Hawaiian Islands and Pitcairn Group; in the western Pacific it is distributed from Wakayama Prefecture, Honshu to the southern Great Barrier Reef. Bruce C. Mundy (pers. comm.) has offered an explanation of the broad distribution of this cardinalfish. He has caught small juveniles in tows in the open ocean far from shallow water, thus indicating that this species can transform from the late postlarval (prejuvenile) stage and still survive in the pelagic realm. Apogon kallopterus is often the most common cardinalfish at many of the localities within its range, including the Hawaiian Islands. Fraser and Lachner (1985) examined about 4000 specimens in museums during the course of their study of the species of Apogon of the subgenus Pristiapogon. They reported it as occurring in a wide array of habitats from tidepools to reef environments as deep as 60 m. Chave and Mundy (1994) reported submarine observations to depths of 158 m in the Hawaiian Islands. This species is more inclined than other Hawaiian apogonids to move over sand and rubble substrata away from the shelter of the reef when feeding at night; it is seen closely oriented to the bottom. Hobson (1974) examined the stomach contents of 24 specimens from Kona, Hawaii; he found decapod shrimps in 12 fish, xanthid crabs in six, crab megalops in four, fishes in two, and mysids in two. Chave (1979) reported the food items eaten by 162 individuals collected off Oahu as 74.9% crustaceans, 12.3% polychaetes, 6.0% fishes, 4.1% ophiuroids, and 2.7% gastropods. Apogon maculiferus Garrett, 1863 Spotted Cardinalfish (Fig. 8) Apogon maculiferus Garrett, 1863: 105 (type locality, Hawaiian Islands). Amia maculifera Jordan and Evermann, 1905: 212, fig. 83. Fig 8. Underwater photograph of Apogon maculiferus, about 90 mm TL, Kaha Bay, Oahu. Dorsal rays VII-I,9; anal rays II,8; pectoral rays 13-14 (usually 14); lateral-line scales 25; scales above lateral line to origin of dorsal fin 2; predorsal scales 5; circumpeduncular scales 14; gill rakers 5-6 + 15-16; edge of preopercle serrate; preopercular ridge smooth in small specimens, finely and irregularly serrate in large ones; body depth 2.6-3.3 in SL; head length 2.35-2.55 in SL; dorsal profile of head slightly convex to straight (except for rounded front of snout); caudal-peduncle length 3.75-4.05 in SL; third dorsal spine clearly longest, 1.75-2.1 in head; caudal fin forked with slightly rounded lobes; color in life pinkish orange with iridescence; longitudinal rows of small yellowish brown spots on body, one per scale, the row just above lateral line nearly joined to form a narrow stripe (spots faint ventrally on body); a blackish spot at base of caudal fin centered on lateral line. Reaches 110 mm SL. This species is known only from the Hawaiian Islands; it appears to be more common in shallow water at the northwestern end of the chain than in the main islands. Bishop Museum collections have been obtained from depths of 1-119 m. Chave and Mundy (1994) reported submarine observations to depths of 153 m. Chave (1979) observed that Apogon maculiferus forages mainly above the substratum, tends to form aggregations while feeding, and feeds more heavily toward dawn. The food eaten by 132 specimens collected off Oahu consisted of crustaceans (94%), fishes (4.3%), polychaetes (1.2%), and gastropods (0.5%). aqua vol. 3 no. 1-1998 34

John E. Randall Apogon taeniopterus Bennett, 1835 Bandfin Cardinalfish (Fig. 9) Apogon taeniopterus Bennett, 1835: 206 (type locality, Mauritius). Apogon menesemus Jenkins, 1903: 448, fig. 19 (type locality, Honolulu). Amia menesema Jordan and Evermann, 1905: 215, fig. 86, pl. 13. Apogon menesemops Lachner in Schultz and collaborators, 1953: 455, fig. 78 (type locality, Bikini Atoll, Marshall Islands). Fig 9. Underwater photograph of Apogon taeniopterus, about 130 mm TL, cave at Moku Manu, Oahu. Dorsal rays VII-I,9; anal rays II,8; pectoral rays 13 (rarely 12 or 14); lateral-line scales 25; scales above lateral line to origin of dorsal fin 2; median predorsal scales 5-6 (usually 6); circumpeduncular scales 16-18; gill rakers 5-6 + 16-18; preopercular edge and ridge serrate; body depth 2.65-3.1 in SL; head length 2.4-2.65 in SL; caudal-peduncle length 3.55-4.4 in SL; third and fourth dorsal spines subequal, 1.95-2.35 in head; caudal fin forked with slightly rounded lobes; color in life pale purplish to reddish brown with iridescence, the edges of scales darker; longitudinal series of indistinct dark blotches often present below lateral line; a black band, edged in pale blue, basally in second dorsal and anal fins, anteriorly on first dorsal and pelvic fins, and at upper and lower margin of caudal fin; black marginal bands in caudal fin joined by a curved black bar, edged in pale blue, about one-third distance from base of fin; a narrow blackish stripe on side of snout. The dark bands are more intense at night and there is much more iridescence, particularly blue-green. Subadults have a large oval blackish spot across the entire caudal peduncle at the base of the caudal fin (see Fraser and Lachner, 1985: fig. 12 of a 60.8-mm specimen from Pohnpei). Largest specimen reported by Fraser and Lachner (1985) 150 mm SL. 35 Randall (1985) placed Apogon menesemus Jenkins from the Hawaiian Islands, and A. menesemops Lachner from the Marshall Islands, in the synonymy of A. taeniopterus. Fraser and Lachner (1985) recognized menesemus as distinct on the basis of the black bar in the caudal fin being complete in menesemus vs. interrupted in taeniopterus, and as shown in their Table 3, 36 specimens of menesemus had 14 pectoral rays (only one with 13), compared to 49 specimens of taeniopterus with 13 pectoral rays (only one with 14). Randall (1996b) followed Fraser and Lachner in recognizing menesemus as a valid endemic Hawaiian species. Further study, however, has shown that specimens of A. taeniopterus from Pitcairn Island (BPBM 16686, 5: 96-122 mm; BPBM 16898, 101 mm) have identical coloration, including a complete bar across the caudal fin, as in menesemus in Hawaii. Furthermore, there is an error in Table 3 of Fraser and Lachner; the count of pectoral rays of menesemus should be 13 for 36 specimens and 12 for one. Accordingly, A. menesemus is again placed in the synonymy of A. taeniopterus. Apogon taeniopterus has an interesting disjunct distribution. It occurs at islands of Oceania from the Pitcairn Group, Marquesas, and Hawaiian Islands west to the Marianas and New Caledonia. There are as yet no records from the Indo-Malayan region, Australia, Taiwan, or Japan. There are four records from the Indian Ocean, all south of the Equator: Christmas Island, Cocos-Keeling Islands, Mauritius, and St. Brandon s Shoals (Fraser and Lachner, 1985: Fig. 10). Bishop Museum specimens of this species have been collected from coral reefs in the depth range of less than 1 to 42 m; it is most common in less than 10 m. At night it is generally found about a metre above the substratum, feeding on zooplankton, but it also feeds at times on benthic prey. Hobson (1974) reported the diet of 31 specimens collected in Kona, Hawaii. He found that crustaceans dominate the prey animals, especially crab megalops and decapod shrimps. Chave (1979) analyzed the food of 132 specimens from Oahu; she found crustaceans (73%), fishes (24.2%), polychaetes (2.8%), and ophiuroids (0.3%). Apogonichthys perdix Bleeker, 1854 Waikiki Cardinalfish (Fig. 10) Apogonichthys perdix Bleeker, 1854: 321 (type locality, Larantuka, Flores). Amia perdix Bleeker, 1873-1876: 100. Apogonichthys perdix Bleeker, 1876-1877: pl. 322, fig. 2. Apogonichthys waikiki Jordan and Evermann, 1903: 179 (type locality, Waikiki, Honolulu). Mionurus waikiki Jordan and Evermann, 1905: 210, pl. 35. aqua vol. 3 no. 1-1998

Review of the Hawaiian apogonids Fig 10. Aquarium photograph of Apogonichthys perdix, about 45 mm TL, Kaneohe Bay, Oahu. Dorsal rays VII-I,9; anal rays II,8; pectoral rays 14; lateral-line scales 23; scales above lateral line to origin of dorsal fin 2; median predorsal scales 4; circumpeduncular scales 12; gill rakers 2-3 + 10-13; preopercular edge membranous; preopercular ridge smooth; supramaxilla present; no palatine teeth; body depth 2.5-2.9 in SL; head length 2.2-2.45 in SL; third dorsal spine longest, 1.9-2.3 in head length; caudal fin rounded; color in life orangish brown with brown blotches, dark brown spots, and whitish flecks, often with white rims on scales of body; indistinct broad dark brown bands radiating posteriorly from eye; a small white spot usually present at rear base of second dorsal fin; caudal fin with a translucent whitish posterior margin. Individual fish may alter their life color greatly, sometimes with red dominating. Largest Hawaiian specimen, 45 mm SL. This species occurs from the Red Sea south to Natal and east to the Hawaiian Islands and Rapa; in the western Pacific from Honshu to New Caledonia. However, there are many intermediate localities for which there is no record. It is generally found in shallow, well-protected coral reefs, often where there is considerable benthic algae. On coasts exposed to heavy surf, it is not apt to occur in less than 20 m. Chave (1979) noted that it hovers a short distance above its place of refuge in the reef while feeding at night. She found crustaceans (91.2%), fishes (7.8%), polychaetes (0.9%), and gastropods (0.1%) in the stomachs of 68 fish from Oahu. Foa brachygramma (Jenkins, 1903) Bay Cardinalfish (Fig. 11) Fowleria brachygrammus Jenkins, 1903: 447, fig. 20 (type locality, Honolulu). Foa brachygramma Jordan and Evermann, 1905: 211, fig. 82. Fig 11. Underwater photograph at night of Foa brachygramma, about 45 mm TL, Kaneohe Bay, Oahu. Dorsal rays VII-I,9; anal rays II,8; pectoral rays 12; lateral line incomplete, ending beneath front of second dorsal fin, the tubed scales 9-11, followed by two or three scales with a small pore; a midlateral row of 7 or 8 scales on caudal peduncle with a small pore; longitudinal scale series 22; scales above lateral line to origin of first dorsal fin 1; median predorsal scales 3, with a pair of overlapping median scales anterior to first predorsal scale; circumpeduncular scales 12; gill rakers 4 + 10-12; preopercular edge and ridge smooth; supramaxilla present; palatine teeth present; body deep, the depth 2.2-2.5 in SL; head length 2.1-2.4 in SL; dorsal profile of head straight to slightly convex; third dorsal spine longest, 1.65-2.05 in head; caudal fin rounded; pale gray to light yellowish brown with golden and pale blue-green iridescence; edges of scales narrowly dark; faint dark bars sometimes evident on body, and indistinct brown bands may radiate from eye; outer part of anterior membranes of first dorsal fin white. Attains 65 mm SL. The Bishop Museum has specimens of Foa brachygramma only from the Hawaiian Islands. It is otherwise reported from the Philippines (Fowler and Bean, 1930), Japan (Hayashi and Kishimoto, 1983, who reviewed earlier records of the species from Japanese waters), Natal (Gon in Smith and Heemstra, 1986), Maldive Islands (Randall and Anderson, 1993), and Queensland (Paxton et al., 1989). Not all of the above records may be valid; a revision of the genus Foa is needed. Even more than the preceding species, Foa brachygramma is a species of protected habitats, particularly bays and harbors. However, it is known from relatively deep water; one collection came from a trawl haul off Molokai in 78-134 m. It is common around dead coral, sponges, and heavy algal growth in bays such as Kaneohe Bay, Oahu. It readily enters brackish environments and has been found in fresh water. aqua vol. 3 no. 1-1998 36

John E. Randall Chave (1979) observed that this species is usually near the bottom after dusk but rises during calm nights if there is half to full moon. Her analysis of the diet of 96 fish from Oahu consisted of crustaceans (90.7%), polychaetes (5.1%), fishes (3.2%), ophiuroids (0.6%), and gastropods (0.4%). Lachneratus phasmaticus Fraser and Struhsaker, 1991 Phantom Cardinalfish (Fig. 12) Lachneratus phasmaticus Fraser and Struhsaker, 1991: 718, figs. 1, 2 (type locality, Ka u Loa Point, Kona, Hawaii). Fig 12. Underwater photograph of Lachneratus phasmaticus, about 80 mm TL, cave in Kona, Hawaii. Dorsal rays VI-I,12-13; anal rays II,13-16; pectoral rays 10; scales cycloid and highly deciduous (no scale counts possible, and none given in original description); no lateral line; gill rakers 2-3 + 14-16; preopercular edge smooth with 2 or 3 distinct spines at corner; preopercular ridge smooth; lower jaw slightly projecting; slender incurved canine teeth in jaws and on vomer, in addition to villiform teeth; a row of slender conical teeth on palatines; body elongate, the depth 3.2-3.5 in SL; head length 2.35-2.50 in SL; caudalpeducle length 4.1-4.6 in SL; fin spines slender; second dorsal spine longest, 2.35-2.7 in head; caudal fin deeply forked, the lobes pointed; transparent light red in life. Largest specimen, 73.6 mm SL. This species is known from three widely separated localities in the Indo-Pacific region: Hawaii, Fiji, and the Comoro Islands. Thirteen of the type specimens, including the holotype, were collected by the author and Bruce A. Carlson from only 3-5 m at the rear part of a cave at the island of Hawaii that was totally dark during daylight. The fish were hovering in midwater. The largest paratype was taken by trawl off Haleiwa, Oahu in 95-104 m. The Fiji specimen was collected in 33.5 m on a vertical drop-off. The Comoro Island paratype, 59.2 mm, was netted at a night light. Acknowledgements Thanks are due the many collectors, in particular John L. Earle and David W. Greenfield, whose specimens provided us with the above descriptions and diagnoses. I am grateful to E. H. Chave and Gerald R. Allen for their critical review of the manuscript. I also thank Jeffrey T. Williams for information on the type specimens of Gymnapogon gracilicauda and Arnold Y. Suzumoto and Loreen R. O Hara for taking radiographs. References Bennett, E. T. 1835. Characters of several fishes from Isle de France. Proc. Zool. Soc. London 3: 206-208. Bleeker, P. 1854. Bijdrage tot de kennis der ichthyologische fauna van het eiland Flores. Nat Tijdschr. Nederlandaises. Indi 6: 311-338. Bleeker, P. 1856. Beschrijvingen van nieuwe of weinig bekende vischsoorten van Manado en Makassar, grootendeels verzameld op eene reis naar den Molukschen Archipel in het gevolg van den Gouverneur-Generaal Duymaer van Twist. Act. Soc. Sci. Indo-Nederl. 1: 1-76. Bleeker, P. 1873-1876. Atlas Ichthyologique des Indes Orientales Nederlandaises, vol. 7: 126 pp., Fréderic Muller, Amsterdam. Bleeker, P. 1876-1877. Atlas Ichththologique des Indes Orientales Nederlandaises, vol. 8: 156 pp., Fréderic Muller, Amsterdam. Böhlke, J. E. & C. C. G. Chaplin. 1968. Fishes of the Bahamas and Adjacent Tropical Waters, xxiii + 771 pp., Livingston Publishing Co., Wynnwood, Pennsylvania. Böhlke, J. E. & J. E. Randall. 1968. A key to the shallow-water west Atlantic cardinalfishes (Apogonidae), with descriptions of five new species. Proc. Philad. Acad. Nat. Sci. 120: 175-206. Chave, E. H. 1979. General ecology of six species of Hawaiian cardinalfishes. Pac. Sci. 32(3): 245-270. Chave, E. H. & B. C. Mundy. 1994. Deep-sea benthic fish of the Hawaiian Archipelago, Cross Seamount, and Johnston Atoll. Pac. Sci. 48(4): 367-409. Fowler, H. W. & B. A. Bean. 1930. Contribution to the biology of the Philippine Archipelago and adjacent regions. The fishes of the families Amiidae, Chandidae, Duleidae, and Serranidae, obtained by the United States Bureau of Fisheries steamer Albatross in 1907-1910, chiefly in the Philippine Islands and adjacent seas. Bull. U.S. Natl. Mus. 100, vol. 10: ix + 334 pp. Fraser, T. H. 1972. Comparative osteology of the shallow water cardinal fishes (Perciformes: Apogonidae) with reference to the systematics and evolution of the family. Ichth. Bull. 34: 1-105. 37 aqua vol. 3 no. 1-1998