KeywordsApogonidae, Pseudamiops, Apogon, Apogon-

ichthys, Foa, Lachneratus, new species, HawaiianIslands.

AbstractThe 10 species of apogonid fishes of the Hawaiian

Islands are differentiated in a key. Pseudamiopsdiaphanes, sp. n., endemic to Hawaii, is distinct inhaving VI-I,8 dorsal rays, II,8 anal rays, 16-18 pectoralrays, a spine-like ventral projection posteriorly on themaxilla, deciduous cycloid scales, no lateral line, arhomboid caudal fin, and in being transparent in life.Apogon deetsie, sp. n., from the Hawaiian Islands andTuamotu Archipelago, has VI-I,9 dorsal rays, II,8 analrays; 11-12 (usually 12) pectoral rays, 24 lateral-linescales, 6 predorsal scales, 3-4 + 12-13 gill rakers, aprotruding membranous flap on corner and loweredge of preopercle, a forked caudal fin with roundedlobes, and is light red with two blackish bars posteri-orly on the body. The remaining species are diag-nosed, and their distributions given. Apogon erythri-nus Snyder appears to be endemic to Hawaii andJohnston Island; A. maculiferus Garrett is also aHawaiian endemic; A. evermanni Jordan and Snyderis widespread in the Indo-Pacific and is also knownfrom the western Atlantic; A. kallopterus Bleeker, A.taeniopterus Bennett (A. menesemus Jenkins is asynonym), Apogonichthys perdix Bleeker (A. waikikiJordan and Evermann is a synonym), Foa brachy-gramma (Jenkins), and Lachneratus phasmaticusFraser and Struhsaker are wide-ranging Indo-Pacificspecies.

ZusammenfassungDie 10 Spezies der Apogonidae Fische der hawai-

ianischen Inseln werden mit einem Schlüssel verse-hen; Pseudamiops diaphanes, sp. n. ist endemischum Hawaii, hat VI-1,8 hintere Strahlen, 11,8 analeStrahlen, 16-18 Bruststrahlen, eine wirbelartige,vorgewölbte Maxilla, abfallende, ringlförmige Schup-pen, keine Seitenlinie, eine rhombenförmige Caudale,und ist insgesamt transparent. Apogon deetsie, sp. n.kommt bei Hawaii und im Luamotu Archipel vor. HatVI-1,9 hintere Strahlen, 11,8 anale Strahlen 11-12 (für

gewöhnlich 12) Bruststrahlen, 24 seitliche Linien-Schuppen, 6 predorsale Schuppen, 3-4 + 12-13Kiemenlamellen, einen hervortretenden, hautartigenLappen an der Ecke und am unteren Ende desPreopercles einen gabelartigen Schwanz mitabgerundeten Enden. Die Art ist hellrot gefärbt mitzwei schwarzen Balken am hinteren Körperteil. Fürdie verbleibenden Spezies ist eine Diagnose erstelltworden und ihre Verbreitung wird erwähnt. Apogonerythrinus Snyder scheint bei Hawaii endemisch zusein. A. evermanni Jordan und Snyder ist weitverbrei-tet im Indo Pazifik und lebt auch im westlichemAtlantik. A. kallopterus Bleeker, A. taeniopterus Ben-nett (A. menesemus Jenkins ist ein Synonym),Apogonichthus perdix Bleeker (A. waikiki Jordan undEvermann ist ein Synonym), Foa brachygramma(Jenkis) und Lachneratus phasmaticus Fraser undStruhsaker sind weitverbreitete Indo-Pazifik Spezies.

RésuméUne clé est proposée qui permet de déterminer les 10espèces d’Apogonidés des îles Hawaii. Pseudamiopsdiaphanes sp. n., endémique des Îles Hawaii, se dis-tingue par la présence de VI-I,8 rayons dorsaux, II,8rayons anaux, 16-18 rayons pectoraux, une projectionépineuse ventrale insérée postérieurement sur lemaxillaire, des écailles cycloïdes décidues, l’absencede ligne latérale, une nageoire caudale rhomboïde etla transparence du corps de l’animal en vie. Apogondeetsie, sp. n., des Îles Hawaii et de l’archipel desTuamotu, présente VI-I,9 rayons dorsaux, II,8 rayonsanaux, 11-12 (habituellement 12) rayons pectoraux,24 écailles sur la ligne latérale, 6 écailles prédorsales,3-4 + 12-13 branchiospines, une extension membra-neuse à l’angle inférieur du préoperculaire, unenageoire caudale fourchue avec des lobes arrondis,une coloration rouge clair avec deux barres noirâtres àl’arrière du corps. Pour les autres espèces, la diagnoseet la distribution sont indiquées. Apogon erythrinusSnyder semble endémique d’Hawaii et de JohnstonIsland; A. maculiferus Garrett est également en-démique d’Hawaii; A. evermanni Jordan et Snyder esttrès répandu dans l’Indo-Pacifique et également connude l’Atlantique occidental; A. kallopterus Bleeker,A. taeniopterus Bennett (synonyme: A. menesemus

aqua vol. 3 no. 1 - 199825

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

Jenkins), Apogonichthys perdix Bleeker (synonyme:A. waikiki Jordan et Evermann), Foa brachygramma(Jenkins) et Lachneratus phasmaticus Fraser etStruhsaker sont des espèces indo-pacifiques à largerèpartition.

SommarioLe 10 specie di pesci apogonidi delle isole Hawaii

sono differenti in una cosa. I Pseudamiops diaphanes,nuova specie endemica delle Hawaii, si distingue peril possesso di VI-l,8 raggi dorsali, II, 8 raggi anali, 16-18 raggi pettorali, una proiezione addominale tipospina posteriore sulla scala cicloide posteriore dellamaxilla, senza linee lateriali, una spina caudale rom-boide e nell’essere trasparente durante la sua vita. L’A-pogon deetsie, nuova specie, delle isola Hawaii el’Arcipelago Tuamotu ha Vi-I,9 raggi dorsali, II,8 raggianali, 11-12 (solitamente 12) raggi pettorali, 24;squame sulla linea laterale, 6 squame predorsali, 3-4+ 12-13 spine branchiali, un lembo di membranasporgente sull’angolo e l’estremità inferiore delpreopercle, una spina caudale biforcuta con lobi roton-di e è rosso chiaro con due strisce nerastre sulla parteposteriore del corpo. Le specie restanti vengono anal-izzate e ne viene segnalata la diffusione.L’Apogon ery-thrinus Snyder sembra essere endemico delle Hawaiie delle isole Johnston; l’A. maculiferus Garret è ancheendemica delle Hawaii; l’A. evermanni Jordan e Sny-der è diffuso nell’indo-pacifico e e viene conosciutoanche nell’Atlantico occidentale; l’A. kallopterusBleeker, l’A. taeniopterus Bennett (A. menesemusJenkins è un sinonimo), l’Apogonichthus perdix Bleek-er (A. waikiki Jordan e Evermann è un sinonimo), ilFoa brachygramma (Jenkins), e Lachneratus phas-maticus Fraser e Struhsaker sono specie molto diffusein Indo-pacifico.

IntroductionThe Apogonidae, popularly known as cardinalfishes,

are a large family of small fishes well-represented intropical to warm temperate seas from a variety ofhabitats. The species are nocturnal, in general, andamong the few marine fishes which are mouthbrood-ers. The male retains the fertilized eggs within themouth until hatching (however, there are also reportsof female apogonids as mouthbrooders). Fishes ofthis family are distinct in having two dorsal fins, thefirst with VI to VIII spines, the second with a singlespine and 8-14 soft rays; an anal fin with II spines and8-18 soft rays; large eyes; a large oblique mouth; asingle opercular spine, and the edge of preoperclepreceded by a parallel ridge.

Fraser (1972) reviewed the classification of the fam-ily from the subfamily to the subgeneric level. He rec-ognized three subfamilies: Apogoninae, Pseudami-nae, and Epigoninae; the last-mentioned taxon is nowgiven family status.

The Hawaiian Islands have relatively few apogonidfishes compared to most other areas of the Indo-Pacific region. Gosline and Brock (1960) recognizedonly eight species (not counting Epigonus atheri-noides and Synagrops argyrea, the latter placed in thePericichthyidae by Fraser, 1972). Fraser and Struh-saker (1991) described Lachneratus phasmaticus asa new genus and species of apogonid fish from theHawaiian islands (type locality), Fiji, and the ComoroIslands. The small number of apogonid species forthe islands is undoubtedly related to the relativelyshort larval life in the plankton due to the early devel-opment taking place within the mouth of the broodingadult fish.

Chave (1979) provided a detailed analysis of thegeneral ecology of the six most common apogonidfishes of the Hawaiian Islands. In the presentpaper two of the species names used by Gosline andBrock (1960) are changed, the species identified bythem as Pseudamiops gracilicauda (Lachner) isdescribed as new, a new species of Apogon isdescribed, and a revised key to the Hawaiian speciesof the family is provided.

Materials and MethodsType 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 Acade-my 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 thesnout in the median plane to the base of the caudal fin(end of hypural plate). Body depth is the greatestdepth; body width is the greatest width just posteriorto the head. Head length is measured from the frontof the upper lip in the median plane (with mouth fullyclosed) to the most posterior point of the opercularmembrane; snout length is taken from the same ante-rior point to the fleshy edge of the orbit. Eye diameteris the greatest diameter of the eye ball (this can bemore accurately measured than the fleshy orbit diam-eter); interorbital width is the least fleshy width. Caud-al-peduncle depth is the least depth, and caudal-peduncle length the horizontal distance between ver-ticals at the rear base of the anal fin and the base ofthe caudal fin; lengths of spines and rays of fins aremeasured from their extreme bases in a straight lineto their tips. Caudal concavity is the horizontal dis-tance between verticals at the tips of the longest andshortest caudal rays. Pectoral-fin length is the lengthof the longest ray; pelvic-fin length is measured fromthe base of the spine to the tip of the longest ray.

Counts of pectoral rays include the splint-likeuppermost ray. Counts of lateral-line scales are madeto the base of the caudal fin (hence do not include one

aqua vol. 3 no. 1 - 1998 26

Review of the Hawaiian apogonids

or two pored scales that extend onto the base of thefin). Often a pored scale lies directly over the end of thehypural plate, hence partly onto the base of the caudalfin; this scale is then included in the count of lateral-linescales. Gill-raker counts include all rudiments; the countof lower-limb rakers contains the long raker at theangle. Developed gill rakers are those which are higherthan the diameter of their base.

Meristic and morphometric data given in paren-theses refer to paratypes. Tables of measurements oftype specimens are presented as percentages of thestandard length. Proportional measurements in thetext 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 onventral edge of maxilla; body very elongate, thedepth 4.25-4.8 in SL; caudal fin rhomboid (Hawai-ian Islands)...........................................................................Pseudamiops diaphanes, new species

1b. Dorsal rays VI or VII, 9-13; pectoral rays 10-14; nosharp bony process posteriorly on ventral edge ofmaxilla; body not very elongate, the depth 2.3-3.9in SL; caudal fin rounded or forked ..............2

2a. Dorsal soft rays 12-13; anal soft rays 13-16; pec-toral rays 10; edge of preopercle smooth with 2 or3 acute spines at corner; scales cycloid; no later-al line (Hawaiian Islands, Fiji, and ComoroIslands).........................Lachneratus phasmaticus

2b. Dorsal soft rays 9; anal soft rays 8; pectoral rays11-15; edge of preopercle smooth or serrate, with-out acute spines at corner; scales ctenoid; lateralline 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; pec-toral rays 14; body not deep, the depth 2.5-2.9 inSL (Indo-Pacific)..................Apogonichthys perdix

4b. Lateral line interrupted, the anterodorsal part end-ing beneath second dorsal fin; palatine teethpresent; pectoral rays 12; body deep, the depth2.2-2.5 in SL (Indo-Pacific).....Foa brachygramma

5a. Scales of lateral line about twice as large asscales of rest of body; predorsal scales 10-12; redwith a blackish stripe on head from front of snoutthrough eye to end of opercle, and a large black-ish spot at rear base of second dorsal fin with asmall white spot adjacent and posterior to it (Indo-Pacific and western Atlantic)...Apogon evermanni

5b. Scales of lateral line not larger than scales of restof body; predorsal scales 5-6; color not as in5a........................................................................6

6a. Spines of first dorsal fin VI; color in lifered.......................................................................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 toorigin of dorsal fin 3; circumpeduncular scales16; color in life transparent red with no distinctdark markings (Hawaiian Islands)...........................................................................Apogon erythrinus

7b. Pectoral rays 11-12; scales above lateral line toorigin of dorsal fin 1; circumpeduncular scales12; color in life light red with 2 broad b l a c k i s hbars, one between posterior halves of seconddorsal and anal fins and the other posteriorly oncaudal peduncle (Hawaiian Islands and TuamotuArchipelago)............Apogon deetsie, new species

8a. Pectoral rays usually 14; preopercular ridgesmooth or with a few small serrae; third dorsalspine clearly longest; orangish with a small dis-tinct dark brown spot on scales of body forminglongitudinal rows; no black stripe in second dor-sal and anal fins, and no black anteriorly on firstdorsal in (Hawaiian Islands).................................................................................Apogon maculiferus

8b. Pectoral rays usually 13; preopercular ridgestrongly serrate; third and fourth dorsal spinessubequal; no rows of distinct small dark brownspots on body (diffuse dark blotches may be pre-sent on body of A. taeniopterus); a black stripe insecond dorsal and anal fins near base; first dorsalfin 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; amidlateral dark brown stripe on head and body(may be diffuse or absent on body of large speci-mens); a dark brown spot at caudal-fin base; noblack 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; nodark brown stripe on body; no dark brown spot atbase of caudal fin; a black bar in caudal fin aboutone-third distance from base, linked to a blacksubmarginal 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 HawaiianIslands); Gosline and Brock, 1960: 162, fig. 179; Ran-dall et al., 1985: 54 (Johnston Island).Holotype: BPBM 37247, female, 30.2 mm, HawaiianIslands, 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

aqua vol. 3 no. 1 - 199827

John E. Randall

Islands, Oahu, Waimea coast, rotenone, V .E. Brock, W.A. Gosline et al., 25 July 1955 (formerly University ofHawaii collection); BPBM 7913, 2: 16.9-22.6 mm,Oahu, Moku Manu, cave in 23 m, rotenone, J.E. Ran-dall, 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, samedata as holotype; BPBM 37252, 32.8 mm, Oahu, Maka-ha, hand net at night, J. L. Earle, July, 1991; USNM343981, 28.8 mm, Oahu, Makaha, Kepuhi Point, cave atshore, hand net at night, 3 m, J. L. Earle, 5 July 1991;AMS I.37827-001, 31.0 mm, Oahu, off main channel toKaneohe Bay, base of drop-off, 25 m, D.W. Greenfield,J. E. Randall, G. Cockrell, and J. L. Mahon, 30 August1991; BMNH 1997.2.4.1, 24.6 mm, NorthwesternHawaiian 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: 18.5-25.3mm, Oahu, off Kaneohe Bay, Sampan Channel, spurand groove zone, 6-7 m, rotenone, D.W. Greenfield andK. R. Longenecker, 1 November 1993.

DiagnosisDorsal rays VI-I,8; anal rays II,8; pectoral rays 16-18;

scales thin, cycloid, highly deciduous, about 23-24 inlongitudinal series, lateral line absent; a sharp, down-ward-projecting, bony process posteriorly on ventralmargin of maxilla; body very elongate, the depth 4.25-4.8 in SL; preopercular margin serrate; preopercularridge smooth; caudal fin rhomboid; transparent in life.

Description

Dorsal rays VI-I,8; anal rays II,8; all dorsal and analsoft rays branched, the last to base; pectoral rays 17(two paratypes with 16, four with 18, 11 with 17), theupper and lower 3-5 unbranched; pelvic rays I,5; prin-cipal caudal rays 17, the upper and lower unbranched;upper procurrent caudal rays 7 (6-8), the posterior 2segmented; lower procurrent caudal rays 6 (6-7), theposterior 2 segmented; gill rakers 16, with 1 upperand 7 lower developed (15-17, with 1 upper and 6-7,usually 7, developed); pseudobranchial filaments rudi-mentary; branchiostegal rays 7; vertebrae 10 +14; nosupraneural (predorsal) bones.

Body very elongate, the depth 4.65 (4.25-4.8) in SL,the greatest depth just posterior to head (except infully ripe females); body moderately compressed, thewidth 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; eyediameter 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 extend-ing slightly posterior to orbit, the upper-jaw length 1.7(1.75-1.85) in head; mouth slightly oblique, the gapeforming an angle of about 20° to horizontal axis ofbody; a sharp, spine-like, downward-projecting, bonyprocess posteriorly on ventral margin of maxilla; nosupramaxilla; upper jaw with 1-3 irregular rows of villi-form teeth; 2 (1-2) large, nearly sessile, inward-pro-jecting canine teeth; medial to villiform teeth on each

aqua vol. 3 no. 1 - 1998 28

Review of the Hawaiian apogonids

Fig 1. Holotype of Pseudamiops diaphanes, BPBM, 30.2 mmSL, Makaha, Oahu.

Fig 2. Underwater photograph at night of Pseudamiopsdiaphanes, about 40 mm TL, Makaha, Oahu.

Fig 3. Underwater photograph at night of Pseudamiops gracili-cauda, about 45 mm TL, Kwajalein Atoll, Marshall Island.

side of symphysis of upper jaw; lower jaw with anirregular outer row of small conical teeth and an innerrow of small, slender, well-separated, incurved canineteeth, those in middle of side of jaw largest (largestabout half pupil diameter); a single row of small coni-cal teeth on palatines, and a double row forming a Von vomer. Anterior part of tongue very slender.Longest gill raker at angle, its length in holotype 1.75in eye diameter.

A single, flat, flexible, right-angle opercular spine;edge of preopercle serrate, with 14 (11-13) irregularserrae, the largest at angle; preopercular ridgesmooth; rows of small papillae on head forming acrosshatch pattern, with 11 transverse rows acrossthe top of the head and continuing downward on sideof head (those posterior to preopercle indistinct); fiveshort rows extending ventrally from eye (four of thefive oblique), and six across mandible; two longitudi-nal rows dorsally on snout and extending throughinterorbital space; five longitudinal rows across preop-ercle, and one along mandible.

Anterior nostril a short flaccid tube in front of centerof eye at edge of groove separating snout from upperlip; posterior nostril a large opening just in front of eyeand only slightly dorsal to anterior nostril; nasal cham-ber cavernous, the skin over the roof thin and sagginginward.

Scales cycloid, thin, and very easily shed (few scalesnow present on any type specimens, hence the countof 23 or 24 scales in longitudinal series is only approx-imate); large scales present on opercle, but none nowapparent on preopercle or elsewhere on head. No lat-eral line evident.

Origin of first dorsal fin slightly posterior to base ofmiddle pectoral rays, the predorsal distance 2.6 (2.6-2.65) in SL; fin spines slender and flexible; first dorsalspine 2.7 (2.9-3.5) in head; third dorsal spine longest,slightly longer than second spine, 2.45 (2.45-2.7) inhead; second dorsal fin well-separated from first dor-sal 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; secondanal spine 2.9 (3.0-3.5) in head; second anal soft raylongest, 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 duskyspots on lips (from individual melanophores), a singlemelanophore ventroposteriorly at edge of eye, a doublerow of melanophores midventrally on abdomen, and adusky bar basally on caudal fin (this marking absent onsome paratypes, prominent and broader on another).

Color of holotype when fresh: transparent with darkmarkings as noted in preserved specimen; body cavi-ty opaquely covered with blackish red; vertebral col-umn reddish black; gill filaments showing through

operculum as bright red. Life color from an under-water photograph, shown in Fig. 2.

EtymologyThis species of Pseudamiops is named diaphanes

from the Greek for transparent.

RemarksPseudamiops diaphanes is known only from the

Hawaiian Islands and Johnston Atoll (Gosline, 1955reported a 23-mm specimen from the atoll asPseudamiops gracilicauda). An uncatalogued lot ofsix specimens, 17-26 mm SL, collected by V. E. Brocket al. at Johnston in 1963, was found in the referencefish collection of the University of Hawaii. This lot isnow BPBM 37413.The specimens are in poor condition, therefore not

designated as paratypes.All Hawaiian specimens of P. diaphanes were col-

lected at the island of Oahu except for one from Mid-way. The species may be expected at the otherislands of the archipelago.

This cardinalfish has been observed alive only atnight and only in caves where it is generally seen wellabove the substratum. Rotenone collections duringthe day have resulted in specimens from the depthrange of 3-25 m, all from caves, except one in a spurand groove zone.

The holotype, the 24.6-mm paratype, the 31-mmparatype, and a 32.8-mm specimen collected in July,1991 (BPBM 37252, not recorded as a paratype dueto bad condition) are fully mature females.Pseudamiops diaphanes is most closely related to P.

gracilicauda, described from the Marshall Islands byLachner in Schultz and collaborators (1953: 497, fig.84) from six specimens, 13.5-33.5 mm SL. P. gracili-cauda has since been reported from the SamoaIslands by Wass (1984: 14) and from Rapa by Randallet al. (1990: 19, fig. 24). It differs from P. diaphanes inhaving the edge of the preopercle smooth with twostout serrae at the angle (edge fully serrate indiaphanes), in having 15-16 pectoral rays (16-18 indiaphanes; only two of 18 specimens with 16 rays),and in color. P. gracilicauda has a dusky blotch on thepreopercle behind the eye and a sprinkling ofmelanophores along the base of the dorsal and analfins; these markings are not apparent on diaphanes.The edges of the scales of gracilicauda are faintly redin life, but not in diaphanes. Lachner gave the gill-raker count of gracilicauda, including rudiments, as 3+ 8 which would seem to be another meristic differ-ence. However, the gill-raker count of Lachner’s holo-type was checked by Jeffrey T. Williams who correct-ed it to 4 + 11 (Randall et al., 1990). Williams alsodetermined that the holotype has 1 weak opercularspine, not 2 as recorded by Lachner, and 2 stout ser-rae at the angle with an additional small one. If theRapa and Samoa specimens are correctly identified as

aqua vol. 3 no. 1 - 199829

John E. Randall

gracilicauda, this species appears to attain a largersize than diaphanes, the largest specimen of whichmeasures 32.8 mm SL. Randall et al. (1990) reportedfive Bishop Museum specimens from Rapa, 35-50 mmSL. One specimen from Tutuila, American Samoa(BPBM 17501) measures 41 mm. Figure 3 is an under-water photograph of P. gracilicauda taken in 7 m at nightat Kwajalein Atoll, Marshall Islands by the author; thefish was estimated to be 45 mm in total length.

Lachner described gracilicauda in the genus Gym-napogon, but he did so tentatively because of thepoor condition of the type specimens. One characterfavoring the placement by him in Gymnapogon wasthe apparent lack of scales. Although Smith (1954:784) doubted that gracilicauda lacked scales, hewrote, „If it should, however, prove that gracilicaudaLachner is truly scaleless, it unquestionably meritsgeneric distinction from Gymnapogon, and the newgenus Lachneria is provisionally proposed, its typegracilicauda Lachner, 1953.“ Gosline (1955) correct-ly concluded that Lachner’s specimens of gracilicaudawere scaled but lost them due to their deciduousnature; therefore there was no need to recognizeLachneria.

Smith (1954: 783) described the new genusPseudamiops, with P. pellucidus Smith (1954: 785, fig.2, Pl. XXIII,A-C) from Bazaruto, Mozambique as thetype species; he included gracilicauda in this genus.P. pellucidus is clearly distinct from gracilicauda anddiaphanes in having 9 anal soft rays, 14 pectoral rays,about 33 scales in longitudinal series, and prominentcanines 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, HawaiianIslands, Oahu, off entrance to channel at NE end ofKaneohe Bay, small cave, 29 m, rotenone, J. E. Ran-dall, E. H. Chave, and W. Hashimoto, 10 October1969.Paratypes: BPBM 10217, 32.8 mm, HawaiianIslands, Oahu, off Makapuu Point, one-fourth mile Eof Manana Island (Rabbit Island), from bilge of a boatraised from an initial depth of 160 m (boat sank onJune 4, 1970; on June 20 it was dragged to a depth of68 m; on October 14 it was dragged farther inshoreand raised to the surface), W .D. Madden et al.; BPBM14004, 26.6 mm, Tuamotu Archipelago, RangiroaAtoll, outside reef 1 mile E of Avatoru Pass, 60° drop-off, 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; USNM343982, 37.0 mm, Oahu, Makaha, under ledge, 44 m,rotenone, J. E. Randall, L. R. Taylor, and P. M. Allen, 16June 1973; BMNH 1997.2.4.2, 38.7 mm, Oahu, offKaneohe Bay, near entrance to ship channel at NW endof bay, cave at base of drop-off, 25 m, rotenone, D. W.Greenfield, K .R. Longenecker, and J. Mendez, 7 Sep-tember 1995; CAS 91614, 40.7 mm and AMS I.37828-001, 50.9 mm, same data as preceding.

DiagnosisDorsal 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 andlower edge with a protruding membranous lobe; pre-opercular ridge smooth; caudal fin forked with round-ed lobes; color light red with two dusky bars posteri-orly on body.

Description

Dorsal rays VI-I,9; anal rays II,8; all dorsal and analsoft rays branched, the last to base; pectoral rays 12(seven paratypes with 12, one with 11), the upper andlower 2 unbranched; pelvic rays I,5; principal caudalrays 17, the upper and lower unbranched; procurrentcaudal rays 6 (6-7, usually 6), the most posterior seg-mented; lateral line complete, the pored scales 24(plus 2 smaller pored scales on caudal-fin base); pre-dorsal scales 6; scales above lateral line to origin offirst dorsal fin 1; scales below lateral line to origin ofanal 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); pseudo-branchial 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 moderatelycompressed, the width 1.9 (1.75-2.1) in depth; headlength 2.5 (2.45-2.55) in SL; dorsal profile of head

aqua vol. 3 no. 1 - 1998 30

Review of the Hawaiian apogonids

Fig 4. Holotype of Apogon deetsie, BPBM 37400, 40.4 mm SL,off Kaneohe Bay, Oahu.

slightly convex to straight (except for rounded front ofsnout); snout length 4.1 (3.85-4.05) in head; eyediameter 3.0 (2.8-3.15) in head; interorbital width4.15 (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 extend-ing nearly to a vertical at posterior edge of orbit, theupper-jaw length 1.85 (1.75-1.9) in head; mouthslightly oblique, the gape forming an angle of about20° to horizontal axis of body; supramaxilla not pres-ent; a band of villiform teeth in jaws (maximum ofabout 10 rows in upper jaw and 5 in lower of holo-type); a single row of very small conical teeth onpalatines (15 teeth in row of holotype); small conicalteeth forming a V-shaped patch on vomer, in 1 to 2rows anteriorly and 3 rows posteriorly (vomerine teethlarger than those in jaws). Anterior part of tongueslender and spatulate with papillae on the upper sur-face. Longest gill rakers at angle and adjacent toangle on lower limb, its length half eye diameter inholotype.

A single, acute, sharp, opercular spine at level ofcenter of eye; upper edge of preopercle serrate, with18 (11-21) serrae; corner and ventral part of preoper-cular edge with a large membranous flap whichextends slightly posterior to serrate edge; lateralissystem of head with numerous close-set pores havingslightly elevated rims; 2 longitudinal rows of poresfrom front of snout to occiput, crossed by 27 (26-30)transverse rows; two suborbital rows of pores with thesame transverse pore system. Anterior nostril a shortmembranous tube slightly below center of eye about anostril diameter from groove at base of upper lip; pos-terior nostril a large vertically ovate aperture directly infront of center of eye, the internarial distance abouthalf pupil diameter; nasal chamber cavernous, theskin over the roof thin.

Scales weakly ctenoid and thin; scales present onnape, opercle, and preopercle, those on opercle larg-er than scales of body; no scales on occiput, interorb-ital, snout, or ventrally on head; no scales on finsexcept for small scales on about basal fourth of caud-al fin and a scaly process of 2 large scales extendingmidventrally from base of pelvic fins to slightly morethan 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 spinesstrong and sharp; first dorsal spine 4.25 (4.1-4.35) inhead; second dorsal spine longest, 2.45 (2.45-2.7) inhead; second dorsal fin separated by a median scale(indented posteriorly) from first dorsal fin, its originabove ninth lateral-line scale; spine of second dorsalfin abnormally short in holotype (2.5-2.8 in head ofparatypes); second dorsal soft ray longest, 1.45 (1.5-1.55) in head; origin of anal fin below or slightly post-erior to origin of second dorsal fin; first anal spineshort, 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 broadlyrounded lobes, 3.1 (2.8-3.15) in SL; caudal concavity3.45 (2.75-3.25) in head; pectoral fins long and point-ed, 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 afaint dusky bar below basal half of second dorsal finand a broader bar posteriorly on caudal peduncle andextending slightly onto base of caudal fin; fins paleexcept about outer fourth of caudal lobes which isdusky.

Color of holotype when first collected: body trans-lucent, the edges of scales broadly light red; twoblackish bars posteriorly on body as describedabove; head light red with an iridescent area of lightyellow and light blue over operculum; iris pale yel-lowish; fins transparent with pale pink rays exceptabout outer fourth of caudal lobes which is blackishand tips of second dorsal and anal fins which aredusky.

EtymologyThis species of Apogon is named deetsie in honor

of E. H. „Deetsie“ Chave, one of the collectors of theholotype, in recognition of her research on the ecolo-gy of apogonid fishes of the Hawaiian Islands. Thespecific name is treated as a noun in apposition.

RemarksApogon deetsie is known only from the Hawaiian

Islands and Rangiroa Atoll in the Tuamotu Archipel-ago. All specimens from the Hawaiian Islands werecollected off Oahu except for one of 62 mm whichwas taken by trawl by Paul J. Struhsaker from theR/V. „Townsend Cromwell“ off the north shore of Maui(20° 57.5’N, 156° 28.6’W) in 66-84 m; the trawl wastowed between 11 and 12 p.m on 7 June 1970. Thespecimen was deposited at the Honolulu Laboratoryof the National Marine Fisheries Service but later dis-carded due to its very poor condition. Meristic datawere taken by E. H. Chave.

This species occurs in moderately deep water,sometimes deeper than normal SCUBA-divingdepths, hence the probable reason for the few speci-mens that have been collected.Apogon deetsie is most closely related to an unde-

scribed species represented by a single specimencollected by the author and Gerald R. Allen in 18-37 mat Florida Island, Solomon Islands on 30 July 1973.This species has the same two broad blackish barsposteriorly on the body, but the ground color in life waslight yellowish, and there is no dark pigment on themedian fins. It differs further in having 13 pectoralrays, 5 + 14 gill rakers, and in lacking a broad mem-branous flap at the corner and lower margin of thepreopercle.

aqua vol. 3 no. 1 - 199831

John E. Randall

Six species of Apogon with two blackish bars post-eriorly on the body occur in the tropical westernAtlantic. Böhlke and Randall (1968: fig. 2) illustratedthe barred pattern of five of these, and Bö hlke andChaplin (1968: 240) the sixth. In view of the occur-rence of Apogon evermanni in the western Atlanticand the Indo-Pacific, A. deetsie was compared withthese six Atlantic species. A. phenax Böhlke and Ran-dall (1968: fig. 5; illustrated in color by Randall, 1996a:124) appears to be the closest relative among theAtlantic species. It differs from A. deetsie in the moreposterior position and usual wedge-shape of the firstblackish bar on the body, 2 scales above the lateralline to the origin of the dorsal fin, usually 7 predorsalscales, 5 + 13-14 gill rakers, and no protruding mem-branous lobe on the corner and ventral margin of thepreopercle. A. planifrons Longley and Hildebrand(illustrated by Böhlke and Chaplin, 1968: 245) is alsosimilar; it differs in having more pointed caudal lobes,2 scales above the lateral line to the origin of the dor-sal fin, 6 + 16-17 gill rakers, and no lower membran-ous preopercular lobe.

Apogon erythrinus Snyder, 1904Hawaiian 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).

DiagnosisDorsal rays VI-I,9; anal rays II,8; pectoral rays 13-15

(one of 20 with 13 and one with 15); lateral-line scales24; scales above lateral line to base of first two dorsalspines 3; predorsal scales 5 or 6; circumpeduncularscales 16; gill rakers 3-4 + 11-13 (one of 20 with 4 andone with 13); upper margin of preopercle serrate; cor-ner and lower edge of preopercle membranous andcrenate but not protruding posteriorly; preopercularridge smooth; body depth 2.5-2.8 in SL; head length

2.35-2.6 in SL; caudal-peduncle length 3.65-3.95 inSL; 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 smallblotch below anterior part of first dorsal fin. Attainsabout 45 mm SL.

RemarksApogon erythrinus appears to be endemic to the

Hawaiian Islands and Johnston Island. Hayashi andKishimoto (l983) have recorded it from the RyukyuIslands and the Philippines; however, these recordsneed confirmation.Apogon erythrinus is a close relative of A. crassiceps

Garman, 1903 (type locality, Fiji), widely distributed inthe Indo-Pacific except the Arabian region (Red Seato Persian Gulf) where another close relative, A. coc-cineus Rüppell,1838, is found.

Randall et al., 1990 reported on the examination ofthe holotype of Apogon crassiceps (MCZ 28214, 27mm SL). They concluded that it is the small, wide-ranging, transparent red species that has been con-fused 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 Winter-bottom 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 usual-

ly having 13 pectoral rays, 2 scales above the lateralline to the base of first two dorsal spines, and a moreelongate body on average (body depth 2.65-3.2 in SL).

The Bishop Museum has numerous specimens ofApogon crassiceps from Rapa (as reported by Ran-dall et al., 1990), Pitcairn Group, Society Islands,Tuamotu Archipelago, Marquesas Islands, LineIslands, Phoenix Islands, Caroline Islands, Kiribati,Marshall Islands, Minami-tori-shima (Marcus Island),Fiji, New Caledonia, Lord Howe Island, Great BarrierReef, Solomon Islands, Indonesia, Philippines,Ryukyu Islands, Ogasawara Islands, Sri Lanka, Sey-chelles, and Mauritius. A. coccineus differs in having3-5 + 13-15 gill rakers and a prominent, broad, longi-tudinal, blackish streak on the side of the caudalpeduncle.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 thesubstratum. It is a difficult fish to photograph in the wildbecause it quickly retreats to a hole in the reef when alight is shone in its direction. Chave noted that onlyApogonichthys perdix among the six common Hawai-ian apogonids is more sensitive to light. She found onlycrustaceans in the stomachs of 68 specimens exam-ined from Oahu. Hobson (1974) found food in eightspecimens from Kona, Hawaii, all crustaceans, ofwhich xanthid crabs were the major prey.

aqua vol. 3 no. 1 - 1998 32

Review of the Hawaiian apogonids

Fig. 5. Underwater photograph at night of Apogon erythrinus,about 45 mm TL, Black Rock, West Maui.

Apogon evermanni Jordan and Snyder, 1904Oddscale 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).

DiagnosisDorsal rays VI-I,9; anal rays II,8; pectoral rays 12; lat-

eral-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 preoperclefinely serrate, the corner and lower edge smooth,membranous, and not extending posterior to serratemargin; preopercular ridge smooth; body moderatelyelongate, the depth 2.7-3.7 in SL; head length 2.3-2.7in SL; dorsal profile of head straight except for round-ed front of snout; caudal peduncle long, 3.2-4.2 in SL;second and third dorsal spines subequal, 2.2-3.2 inhead; caudal fin forked with rounded lobes; red in lifewith a blackish stripe from front of snout, through eye,to end of opercle, a black spot at rear base of seconddorsal fin and an adjacent small white spot at base oflast dorsal ray and onto caudal peduncle. Smallerindividuals tend to be paler red; still smaller onestranslucent. Largest specimen examined, 121.5 mm.

RemarksBöhlke and Randall (1968) described Apogon

anisolepis from one specimen from the Bahamas andone from Curação, never suspecting that the specieshad been described from the Hawaiian Islands as A.evermanni in 1904. That realization came in 1969when the author, S. N. Swerdloff, and E. H. Chave col-lected 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 followinglocalities in the Indo-Pacific, in addition to Hawaii:Marquesas Islands, Marshall Islands, SolomonIslands, Molucca Islands, Cocos-Keeling Islands, Mal-dive Islands, Chagos Archipelago, and Mauritius.They added Cozumel Island, Mexico as an additionalrecord for the western Atlantic. A. evermanni isunknown from the Red Sea and Gulf of Aden; itappears to be replaced there by A. isus Randall andBöhlke. This species is very similar in color and morp-hology to A. evermanni but differs notably in havingthe scales of the body equal in size to the lateral-linescales.

Randall and Böhlke (1981: Table l) noted that thenumber of scales below the lateral line and the num-ber of circumpeduncular scales of Apogon evermanniis 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 beneathledges by day, and even at night is rarely seen outsideof caves. Like A. erythrinus, it is very difficult toapproach with a light at night. Specimens have beencollected in the depth range of 3-69 m, usually in morethan 25 m.

Apogon kallopterus Bleeker, 1856Iridescent 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.

DiagnosisDorsal rays VII-I,9; anal rays II,8; pectoral rays 13-14

(rarely 14); lateral-line scales 25; scales above lateral

aqua vol. 3 no. 1 - 199833

John E. Randall

Fig 6. Underwater photograph of Apogon evermanni, about 110mm TL, cave in Kona, Hawaii.

Fig 7. Underwater photograph of Apogon kallopterus, about80 mm TL, Oahu.

line to origin of dorsal fin 2; median predorsal scales5-6 (usually 6); circumpeduncular scales 12-14 (rarely12, usually 14); gill rakers 4-5 + 13-15; edge of preop-ercle and preopercular ridge serrate; body depth 2.5-3.3 in SL; head length 2.4-2.55 in SL; dorsal profile ofhead convex; caudal-peduncle length 3.65-4.35 in SL;third and fourth dorsal spines subequal, 1.7-1.95 inhead; caudal fin forked, with slightly rounded lobes;color in life light brown, the scale edges darker, withblue-green and yellow iridescence (especially atnight); a dark brown stripe from front of snout to eye,continuing more broadly and diffusely on head behindeye and along side of body (stripe on body absent ornearly so on large individuals); a dark brown spotabout size of pupil posteriorly on caudal peduncle, itslower edge on lateral line; leading edge of first dorsalfin broadly dusky to dark brown; a dark brown todusky band basally in second dorsal and anal fins, onsecond pelvic ray, and submarginally in caudal-finlobes. Largest specimen reported by Fraser and Lach-ner (1985), 122 mm SL.

RemarksApogon kallopterus is a wide-ranging Indo-Pacific

species, occurring from the Red Sea to South Africa,east to the Hawaiian Islands and Pitcairn Group; inthe western Pacific it is distributed from WakayamaPrefecture, Honshu to the southern Great BarrierReef. Bruce C. Mundy (pers. comm.) has offered anexplanation of the broad distribution of this cardinal-fish. He has caught small juveniles in tows in theopen ocean far from shallow water, thus indicatingthat this species can transform from the late post-larval (prejuvenile) stage and still survive in thepelagic 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 museumsduring the course of their study of the speciesof Apogon of the subgenus Pristiapogon.They reported it as occurring in a wide array of habi-tats from tidepools to reef environments as deep as60 m. Chave and Mundy (1994) reported submarineobservations to depths of 158 m in the HawaiianIslands.

This species is more inclined than other Hawaiianapogonids to move over sand and rubble substrataaway from the shelter of the reef when feeding atnight; it is seen closely oriented to the bottom. Hob-son (1974) examined the stomach contents of 24specimens from Kona, Hawaii; he found decapodshrimps in 12 fish, xanthid crabs in six, crab mega-lops in four, fishes in two, and mysids in two. Chave(1979) reported the food items eaten by 162 individu-als collected off Oahu as 74.9% crustaceans, 12.3%polychaetes, 6.0% fishes, 4.1% ophiuroids, and 2.7%gastropods.

Apogon maculiferus Garrett, 1863Spotted Cardinalfish (Fig. 8)

Apogon maculiferus Garrett, 1863: 105 (type locality,Hawaiian Islands).Amia maculifera Jordan and Evermann, 1905: 212,fig. 83.

DiagnosisDorsal rays VII-I,9; anal rays II,8; pectoral rays 13-14

(usually 14); lateral-line scales 25; scales above later-al line to origin of dorsal fin 2; predorsal scales 5; cir-cumpeduncular scales 14; gill rakers 5-6 + 15-16; edgeof preopercle serrate; preopercular ridge smooth insmall specimens, finely and irregularly serrate in largeones; body depth 2.6-3.3 in SL; head length 2.35-2.55in SL; dorsal profile of head slightly convex to straight(except for rounded front of snout); caudal-pedunclelength 3.75-4.05 in SL; third dorsal spine clearlylongest, 1.75-2.1 in head; caudal fin forked with slight-ly rounded lobes; color in life pinkish orange with iri-descence; longitudinal rows of small yellowish brownspots on body, one per scale, the row just above later-al line nearly joined to form a narrow stripe (spots faintventrally on body); a blackish spot at base of caudal fincentered on lateral line. Reaches 110 mm SL.

RemarksThis species is known only from the Hawaiian Islands;

it appears to be more common in shallow water at thenorthwestern end of the chain than in the main islands.Bishop Museum collections have been obtained fromdepths of 1-119 m. Chave and Mundy (1994) reportedsubmarine observations to depths of 153 m.

Chave (1979) observed that Apogon maculiferus for-ages mainly above the substratum, tends to formaggregations while feeding, and feeds more heavilytoward dawn.The food eaten by 132 specimens collected off Oahuconsisted of crustaceans (94%), fishes (4.3%), poly-chaetes (1.2%), and gastropods (0.5%).

aqua vol. 3 no. 1 - 1998 34

Review of the Hawaiian apogonids

Fig 8. Underwater photograph of Apogon maculiferus, about 90mm TL, Kaha Bay, Oahu.

Apogon taeniopterus Bennett, 1835Bandfin Cardinalfish (Fig. 9)

Apogon taeniopterus Bennett, 1835: 206 (type locali-ty, Mauritius).Apogon menesemus Jenkins, 1903: 448, fig. 19 (typelocality, Honolulu).Amia menesema Jordan and Evermann, 1905: 215,fig. 86, pl. 13.Apogon menesemops Lachner in Schultz and collab-orators, 1953: 455, fig. 78 (type locality, Bikini Atoll,Marshall Islands).

DiagnosisDorsal rays VII-I,9; anal rays II,8; pectoral rays

13 (rarely 12 or 14); lateral-line scales 25; scalesabove lateral line to origin of dorsal fin 2; medianpredorsal scales 5-6 (usually 6); circumpeduncularscales 16-18; gill rakers 5-6 + 16-18; preopercularedge and ridge serrate; body depth 2.65-3.1 in SL;head length 2.4-2.65 in SL; caudal-peduncle length3.55-4.4 in SL; third and fourth dorsal spines sub-equal, 1.95-2.35 in head; caudal fin forked withslightly rounded lobes; color in life pale purplish toreddish brown with iridescence, the edges of scalesdarker; longitudinal series of indistinct dark blotchesoften present below lateral line; a black band, edgedin pale blue, basally in second dorsal and anal fins,anteriorly on first dorsal and pelvic fins, and at upperand lower margin of caudal fin; black marginal bandsin caudal fin joined by a curved black bar, edgedin pale blue, about one-third distance from base offin; a narrow blackish stripe on side of snout. Thedark bands are more intense at night and there ismuch more iridescence, particularly blue-green.Subadults have a large oval blackish spot across theentire caudal peduncle at the base of the caudalfin (see Fraser and Lachner, 1985: fig. 12 of a60.8-mm specimen from Pohnpei). Largest spe-cimen reported by Fraser and Lachner (1985) 150mm SL.

RemarksRandall (1985) placed Apogon menesemus Jenkins

from the Hawaiian Islands, and A.menesemops Lach-ner from the Marshall Islands, in the synonymy of A.taeniopterus. Fraser and Lachner (1985) recognizedmenesemus as distinct on the basis of the black bar inthe caudal fin being complete in menesemus vs. inter-rupted 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 tae-niopterus with 13 pectoral rays (only one with 14).Randall (1996b) followed Fraser and Lachner in re-cognizing menesemus as a valid endemic Hawaiianspecies. Further study, however, has shown that spec-imens of A. taeniopterus from Pitcairn Island (BPBM16686, 5: 96-122 mm; BPBM 16898, 101 mm) haveidentical coloration, including a complete bar acrossthe caudal fin, as in menesemus in Hawaii. Further-more, there is an error in Table 3 of Fraser and Lach-ner; the count of pectoral rays of menesemus shouldbe 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 dis-

tribution. It occurs at islands of Oceania from the Pit-cairn Group, Marquesas, and Hawaiian Islands westto the Marianas and New Caledonia. There are as yetno records from the Indo-Malayan region, Australia,Taiwan, or Japan.There are four records from the Indi-an Ocean, all south of the Equator: Christmas Island,Cocos-Keeling Islands, Mauritius, andSt. Brandon’s Shoals (Fraser and Lachner, 1985:Fig. 10).

Bishop Museum specimens of this species havebeen collected from coral reefs in the depth range ofless than 1 to 42 m; it is most common in less than 10m. At night it is generally found about a metre abovethe substratum, feeding on zooplankton, but it alsofeeds at times on benthic prey. Hobson (1974) report-ed 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 specimensfrom Oahu; she found crustaceans (73%), fishes(24.2%), polychaetes (2.8%), and ophiuroids (0.3%).

Apogonichthys perdix Bleeker, 1854Waikiki Cardinalfish (Fig. 10)

Apogonichthys perdix Bleeker, 1854: 321 (type local-ity, 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 - 199835

John E. Randall

Fig 9. Underwater photograph of Apogon taeniopterus, about130 mm TL, cave at Moku Manu, Oahu.

DiagnosisDorsal rays VII-I,9; anal rays II,8; pectoral rays 14;

lateral-line scales 23; scales above lateral line to or-igin of dorsal fin 2; median predorsal scales 4; cir-cumpeduncular scales 12; gill rakers 2-3 + 10-13; pre-opercular edge membranous; preopercular ridgesmooth; supramaxilla present; no palatine teeth; bodydepth 2.5-2.9 in SL; head length 2.2-2.45 in SL; thirddorsal spine longest, 1.9-2.3 in head length; caudal finrounded; color in life orangish brown with brownblotches, dark brown spots, and whitish flecks, oftenwith white rims on scales of body; indistinct broaddark brown bands radiating posteriorly from eye; asmall white spot usually present at rear base of sec-ond dorsal fin; caudal fin with a translucent whitishposterior margin. Individual fish may alter their lifecolor greatly, sometimes with red dominating. LargestHawaiian specimen, 45 mm SL.

RemarksThis species occurs from the Red Sea south to Natal

and east to the Hawaiian Islands and Rapa; in thewestern Pacific from Honshu to New Caledonia. How-ever, there are many intermediate localities for whichthere is no record. It is generally found in shallow,well-protected coral reefs, often where there is con-siderable benthic algae. On coasts exposed to heavysurf, it is not apt to occur in less than 20 m. Chave(1979) noted that it hovers a short distance above itsplace of refuge in the reef while feeding at night. Shefound crustaceans (91.2%), fishes (7.8%), poly-chaetes (0.9%), and gastropods (0.1%) in the stom-achs 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.

DiagnosisDorsal rays VII-I,9; anal rays II,8; pectoral rays 12;

lateral line incomplete, ending beneath front of sec-ond dorsal fin, the tubed scales 9-11, followed by twoor three scales with a small pore; a midlateral row of 7or 8 scales on caudal peduncle with a small pore; lon-gitudinal scale series 22; scales above lateral line toorigin of first dorsal fin 1; median predorsal scales 3,with a pair of overlapping median scales anterior tofirst predorsal scale; circumpeduncular scales 12; gillrakers 4 + 10-12; preopercular edge and ridgesmooth; 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 con-vex; third dorsal spine longest, 1.65-2.05 in head; cau-dal fin rounded; pale gray to light yellowish brown withgolden and pale blue-green iridescence; edges ofscales narrowly dark; faint dark bars sometimes evid-ent on body, and indistinct brown bands may radiatefrom eye; outer part of anterior membranes of firstdorsal fin white. Attains 65 mm SL.

RemarksThe Bishop Museum has specimens of Foa brachy-

gramma only from the Hawaiian Islands. It is other-wise reported from the Philippines (Fowler and Bean,1930), Japan (Hayashi and Kishimoto, 1983, whoreviewed earlier records of the species from Japanesewaters), Natal (Gon in Smith and Heemstra, 1986),Maldive Islands (Randall and Anderson, 1993), andQueensland (Paxton et al., 1989). Not all of the aboverecords may be valid; a revision of the genus Foa isneeded.

Even more than the preceding species, Foa brachy-gramma is a species of protected habitats, particular-ly bays and harbors. However, it is known from relat-ively deep water; one collection came from a trawlhaul off Molokai in 78-134 m. It is common arounddead coral, sponges, and heavy algal growth in bayssuch as Kaneohe Bay, Oahu. It readily enters brack-ish environments and has been found in fresh water.

aqua vol. 3 no. 1 - 1998 36

Review of the Hawaiian apogonids

Fig 10. Aquarium photograph of Apogonichthys perdix, about45 mm TL, Kaneohe Bay, Oahu.

Fig 11. Underwater photograph at night of Foa brachygramma,about 45 mm TL, Kaneohe Bay, Oahu.

Chave (1979) observed that this species is usuallynear the bottom after dusk but rises during calmnights if there is half to full moon. Her analysis of thediet of 96 fish from Oahu consisted of crustaceans(90.7%), polychaetes (5.1%), fishes (3.2%), ophi-uroids (0.6%), and gastropods (0.4%).

Lachneratus phasmaticusFraser and Struhsaker, 1991Phantom Cardinalfish (Fig. 12)

Lachneratus phasmaticus Fraser and Struhsaker,1991: 718, figs. 1, 2 (type locality, Ka’u Loa Point,Kona, Hawaii).

DiagnosisDorsal rays VI-I,12-13; anal rays II,13-16; pectoral

rays 10; scales cycloid and highly deciduous (no scalecounts possible, and none given in original descript-ion); no lateral line; gill rakers 2-3 + 14-16; preopercu-lar edge smooth with 2 or 3 distinct spines at corner;preopercular ridge smooth; lower jaw slightly project-ing; slender incurved canine teeth in jaws and onvomer, in addition to villiform teeth; a row of slenderconical teeth on palatines; body elongate, the depth3.2-3.5 in SL; head length 2.35-2.50 in SL; caudal-peducle length 4.1-4.6 in SL; fin spines slender; sec-ond dorsal spine longest, 2.35-2.7 in head; caudal findeeply forked, the lobes pointed; transparent light redin life. Largest specimen, 73.6 mm SL.

RemarksThis species is known from three widely separated

localities in the Indo-Pacific region: Hawaii, Fiji, andthe Comoro Islands. Thirteen of the type specimens,including the holotype, were collected by the authorand Bruce A. Carlson from only 3-5 m at the rear partof a cave at the island of Hawaii that was totally darkduring 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 Islandparatype, 59.2 mm, was netted at a night light.

AcknowledgementsThanks are due the many collectors, in particular John

L. Earle and David W. Greenfield, whose specimensprovided us with the above descriptions and diagnoses.I am grateful to E. H. Chave and Gerald R. Allen for theircritical review of the manuscript. I also thank Jeffrey T.Williams for information on the type specimens of Gym-napogon gracilicauda and Arnold Y. Suzumoto andLoreen R. O’Hara for taking radiographs.

References

Bennett, E.T. 1835. Characters of several fishes fromIsle de France. Proc. Zool. Soc. London 3: 206-208.Bleeker, P. 1854. Bijdrage tot de kennis der ichthyol-

ogische 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 denMolukschen Archipel in het gevolg van den Gou-verneur-Generaal Duymaer van Twist. Act. Soc. Sci.Indo-Nederl. 1: 1-76.Bleeker, P. 1873-1876.Atlas Ichthyologique des IndesOrientales Nederlandaises, vol. 7: 126 pp., FrédericMuller, Amsterdam.Bleeker, P. 1876-1877. Atlas Ichththologique desIndes Orientales Nederlandaises, vol. 8: 156 pp.,Fréderic Muller, Amsterdam.Böhlke, J. E. & C. C. G. Chaplin. 1968. Fishes ofthe Bahamas and Adjacent Tropical Waters, xxiii +771 pp., Livingston Publishing Co., Wynnwood,Pennsylvania.Böhlke, J. E. & J. E. Randall. 1968. A key to the shal-

low-water west Atlantic cardinalfishes (Apogon-idae), 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 ben-

thic fish of the Hawaiian Archipelago, Cross Sea-mount, 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 adjacentregions. The fishes of the families Amiidae, Chandi-dae, Duleidae, and Serranidae, obtained by the Unit-ed States Bureau of Fisheries steamer „Albatross“ in1907-1910, chiefly in the Philippine Islands andadjacent seas. Bull. U.S. Natl. Mus. 100, vol. 10: ix +334 pp.Fraser,T. H. 1972. Comparative osteology of the shal-

low water cardinal fishes (Perciformes: Apogonidae)with reference to the systematics and evolution of thefamily. Ichth. Bull. 34: 1-105.

aqua vol. 3 no. 1 - 199837

John E. Randall

Fig 12. Underwater photograph of Lachneratus phasmaticus,about 80 mm TL, cave in Kona, Hawaii.

Fraser, T. H. & E. A. Lachner. 1985. A revision of thecardinalfish subgenera Pristiapogon and Zoramia(genus Apogon) of the Indo-Pacific region (Teleostei:Apogonidae).Smiths. Contr. Zool., no. 412: iii + 47 pp.Fraser, T. H. & P. J. Struhsaker. 1991. A new genus

and species of cardinalfish (Apogonidae) from theIndo-West Pacific, with a key to apogonine genera.Copeia 1991(3): 718-722.Garman, S. 1903. Some fishes from Australasia. Bull.Mus. Comp. Zoöl. 39(8): 229-241.Garrett, A. J. 1863. Descriptions of new species of

fishes. Proc. Calif. Acad. Sci. 3: 103-107.Gosline, W. A. 1955. The inshore fish fauna of John-

ston Island, a central Pacific atoll. Pac. Sci. 9(4):442-480.Gosline, W. A. & V. E. Brock. 1960. Handbook ofHawaiian Fishes, 372 pp., University of HawaiiPress, Honolulu.Hayashi, M. & H. Kishimoto. 1983. Fish fauna of Iri-

omote-Island, Ryukyu Islands III. Apogonidae(Apogoninae).Sci. Rept.Yokosuka City Mus., no. 31:15-46 (in Japanese).Hobson, E. S. 1974. Feeding relationships of

teleostean fishes on coral reefs in Kona, Hawaii.Fishery Bull. 72(4): 915-1031.Jenkins, O. P. 1903. Report on collections of fishes

made in the Hawaiian Islands, with descriptions ofnew species. Bull. U.S. Fish Comm. 22: 417-511.Jordan, D. S. & B.W. Evermann. 1903. Descriptions

of new genera and species of fishes from theHawaiian Islands. Bull. U.S. Fish Comm. 22:161-208.Jordan, D. S. & B.W. Evermann. 1905. The aquatic

resources of the Hawaiian Islands. Part I.— the shorefishes. Bull. U.S. Fish Comm. 23: xxviii + 574 pp.Jordan, D. S. & A. Seale. 1906. The fishes of Samoa.Bull. Bur. Fisheries 25: 173-488.Jordan, D. S. & J. O. Snyder. 1904. Description of a

new species of fish (Apogon evermanni) from theHawaiian Islands, with notes on other species. Proc.U.S. Natl. Mus. 28: 123-126.Paxton, J. R., D. F. Hoese, G.R. Allen & J.E.Hanley.

1989. Zoological Catalogue of Australia, vol. 7.Pisces, Petromyzontidae to Carangidae, xii + 665 pp.,Australian Government Publishing Service, Canberra.Madden,W. D. 1973. The collection of live fishes from

a salvaged vessel. Copeia 1973(1): 144-145.Randall, J. E. 1985. Guide to Hawaiian Reef Fishes,

79 pp., Harrowood Books, Newtown Square, Penn-sylvania.Randall, J. E. 1996a. Caribbean Reef Fishes, third

edition, 369 pp., T.F.H. Publications, Neptune City,New Jersey.Randall, J. E. 1996b. Shore Fishes of Hawaii, 216

pp., Natural World Press, Vida, Oregon.Randall, J. E. & R. C. Anderson. 1993. Annotated

checklist of the epipelagic and shore fishes of theMaldive Islands. Ichth. Bull. 59: 1-47.

Randall, J. E. & J. E. Böhlke. 1981. The status of thecardinalfishes Apogon evermanni and A. anisolepis(Perciformes: Apogonidae) with description of arelated new species from the Red Sea. Proc. Acad.Nat. Sci. Phila. 133: 129-140.Randall, J. E., P. S. Lobel & E. H. Chave. 1985.

Annotated checklist of the fishes of Johnston Island.Pac. Sci. 39(1): 24-80.Randall, J. E., C. L. Smith & M. N. Feinberg. 1990.

Report on fish collections from Rapa, French Poly-nesia. Amer. Mus. Novit., no. 2966: 1-42.Rüppell, E. 1835-1838. Neue Wirbelthiere zu derFauna von Abyssinien gehörig. Fische des rothenMeeres, 148 pp., Siegmund Schmerber, Frankfurt.Schultz, L. P. & collaborators. 1953. Fishes of the

Marshall and Marianas Islands. Bull. U.S. Natl. Mus.202, vol. 1: xxxii + 685 pp.Smith, J. L. B. 1954. Apogonid fishes of the subfami-

ly Pseudamiinae from South-East Africa. Ann. Mag.Nat. Hist., ser. 12, vol. 7: 775-795.Smith, J. L. B. 1961. Fishes of the family Apogonidae

of the western Indian Ocean and the Red Sea. Ichth.Bull., no. 22: 371-418.Smith, M. M. & P. C. Heemstra. 1986. Smiths’ SeaFishes, xx + 1047 pp., Macmillan South Africa,Johannesburg.Snyder, J. O. 1904. A catalogue of the shore fishes col-

lected by the steamer Albatross about the HawaiianIslands in 1903. Bull. U.S. Fish Comm. 22: 513-538.Wass, R. C. 1984. An annotated checklist of the fish-

es of Samoa. NOAA Tech. Rep., NMFS SSRF no.781: v + 43 pp.Winterbottom, R., A. R. Emery & E. Holm. 1989. An

annotated checklist of the fishes of the Chagos Arch-ipelago, central Indian Ocean. Roy. Ontario Mus. LifeSci. Contr. no. 145: vi + 226 pp.

aqua vol. 3 no. 1 - 1998 38

Review of the Hawaiian apogonids