BULLETIN OF MARINE SCIENCE, 25(3): 393-421, ]975

A REVISION OF THE INDO-PACIFIC ANGELFISH GENUS GENICA NTH US, WITH DESCRIPTIONS OF THREE NEW SPECIES

John E. Randall

ABSTRACT The Indo-Pacific pomacanthid genus Gellicallthus containing nine species is dis- tinguished chiefly by its emarginate to lunate caudal fin (often with produced lobes), relatively short teeth, and sexual dichromatism. Males have either bars or stripes on the body (black on all species except G. bel/us which has a yellow stripe) and lack black on the lobes of the caudal fin, whereas females (except the striped G. lamarck and the diagonally banded G. bel/us) lack dark markings on the body and all except G. spillus have a prominent black band in each lobe of the caudal fin. G. semicillctu.l' (Waite), known only from Lord Howe Island, is resurrected from the synonymy of G. melallospilos (Bleeker) of the western Pacific. G. caudibicolor (Lienard, in Sauvage) and G. zebra (Lienard, in Sauvage) are regarded as synonyms of G. caudovillatus (GUnther) of the western Indian Ocean; the latter, however, is not a synonym of G. melallospilos, as some authors have contended. G. macclesfieldiellsis Chan is referred to the synonymy of G. melanospilos. G. fucosus Yasuda and Tominaga is the probable female form of G. semi- fasciatus (Kamohara), which ranges from the Philippines to Japan. The distribution of G. melllllOspilos is extended to the New Hebrides, Loyalty Islands, Solomon Islands, Palau Islands, Fiji Islands, New Guinea, and the Ryukyu Islands; that of G. lamarck to the Solomon Islands; and that of G. watanabei to Pitcairn Island, Austral Islands, New Cale- donia, and Osprey Reef (Coral Sea). The following three new species are described: G. spillus (a relative of G. semicillctus) from the Pitcairn Group and Austral Islands; G. bel/us from Tahiti and Cocos-Keeling Islands; and G. persollatus from Hawaii. The male of the latter has not yet been collected.

Swainson (1839) established the poma- re-established as a genus of Pomacanthidae. canthid genus Genicanthus, noting that the In addition to the concave caudal fin, he body was more elongate than Holacanthus distinguished Genicanthus from Holacanthus and Pomacanthus and the caudal fin "deeply as follows: "Preorbital notched mesially, its lunated, the ends extended into filaments." hind margin free, serrated. Teeth in both He added "operculum spined as in Hola- jaws short. Scales on operculum in six to canthus." He mistakenly included Hola- eight rows." (Holacanthus listed as having canthus tricolor (Bloch) with H. lamarck about nine rows). Though many authors Lacepede in Genicanthus; tricolor is the type such as Fowler (1934), Weber and de Beau- specics of Holacanthus. fort (1936), Smith (1955), Norman (1957), Genicanthus went unrecognized as a valid Chan (1965), who named G. macclesfieldi- genus for many years. Bleeker (1857) ensis from the South China Sea, Munro described Holacanthus melanospilos from (1967), Araga (1972), and Randall (1973) Ambon, East Indies; Gunther (1860) named have followed Fraser-Brunner, others such the related H. caudovittatus from Mauritius; as Kamohara (1934, 1961), who described and Waite (1900) described H. semicinctus Holacanthus semi/asdatus from southern from Lord Howe Island. All of these species Japan, Watanabe (1949), Herre (1953), should have been placed in Genicanthus. and Yasuda and Tominaga (1970), have Not until the revision of the angelfishes reverted to Holacanthus. by Fraser-Brunner (1933) was Genicanthus In their excellent paper on "long-tailed" 393 394 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975 angelfishes, Yasuda and Tominaga described Holacanthus as Genicanthus, one or two two new species, G. fucosus from Japan and more rows may be counted on the former. G. watanabei from Okinawa. They wrote It is with the emarginate to lunate caudal that the pre orbital of Genicanthus is not fin, however, that Genicanthus is most readily always notched mesially and its hind mar- differentiated from all other angelfishes. gin is not always free, that the teeth are not Fraser-Brunner separated Genicanthus and always short, and they doubted the value of Holacanthus from Heteropyge (= Euxiphi- the number of vertical scale rows on the pops) in part as follows: "Lateral line ter- operde as a distinguishing character because minating at end of soft dorsal, sometimes a the count "may differ by one or two depend- separate portion on caudal peduncle." The ing on the investigators' counting method." lateral line was described as complete for They preferred to retain the long-tailed spe- Euxiphipops. Although the lateral line of cies in Holacanthus until an osteological Genicanthus and Holacanthus may be poorly study of the world representatives of the developed in the descending portion beneath family can be undertaken. the hind part of the soft portion of the dorsal While osteological research is certainly fin and one or two scales of the series in this needed to resolve confusion in the generic sector may lack pores, I would not describe it classification of pomacanthids, there does as incomplete for these genera. seem to be enough justification from external Genicanthus is unique among the poma- characters, the mode of life, and the striking canthid genera in the striking sexual di- sexual dichromatism to warrant the retention chromatism exhibited by all the species (ex- of Genicanthus as a genus. cept G. personatus for which the male is not I find Fraser-Brunner's separation of yet known). I first suspected that the color Genicanthus from Holacanthus largely cor- differences in Genicanthus were sexual when rect. The anteroventral notch in the pre- I speared four specimens of G. watanabei in orbital is present on the various species of Tahiti in 1969 and found a striped male and Genicanthus though not always as deep as three females with unmarked bodies. I ob- illustrated by Fraser-Brunner (1933: text- served that the two color forms occurred fig. 16), and it may be partially obscured by together in the same deep reef habitat; also, spines. Furthermore, the hind margin of the no small individuals in the striped pattern preorbital is free for about one-third to one- were seen. Similar observations were made half the distance from the lowermost edge to of G. melanospilos in the Palau Islands in the eye. However, a similar notch can be 1970. More convincing was the observation found on the preorbital of some specimens of of courtship of the barred male and plain- Holacanthus, and the posterior margin of bodied female of G. caudovittatus in the Gulf of Aqaba, Red Sea, in 1972, as well as this bone can be as free as that of Geni- noting an adult individual just developing canthus. the barred color pattern of the male. No Compared to Holacanthus, the teeth of small individuals of any species of Geni- Genicanthus are indeed relatively short. canthus have been found with the male color Their length is contained about five times in pattern, thus suggesting that sex reversal the diameter of the eye, whereas the length takes place, as has been demonstrated for of the teeth of Holacanthus is contained from various Serranidae, and Labridae. less than two to three times in the eye. The species of Genicanthus are poorly Admittedly, the vertical scales on the represented in the collections of the museums opercle are difficult to count because they of the world. In fact, some major museums do not lie in regular rows, and there may be have no specimens of the genus. This is small scales at either end. But if the same partly due to the occurrence of the species method is used to count the larger rows on in relatively deep water. Although one or RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 395 two species may occasionally be seen in as marek, G. melanospilos, G. watanabei, und little as 15 m, these fishes are not commonly G. semifasciatus. It is possible that G. bellus found in less than about 30 m. Also, they will ultimately be found there also, for Indo- arc associated with reefs or rocky bottom, nesia is contained within the present range oftcn ncar vertical discontinuities, hence are of Tahiti and Cocos-Keeling Islands. not apt to be taken in trawls. They have Only two species are definitely known from small mouths, thus are rarely caught on the Indian Ocean: G. caudovittatus in the hook and line. western part, ranging from Mauritius to the They swim higher above the bottom, Red Sea, a:Id G. bellus from Cocos-Keeling. gcncrally, than other angelfishes, probably The one record of a juvenile G. lamarck from reflecting their feeding to a significant degree Kenya cannot be confirmed due to loss of the on the larger animals of the zooplankton (the specimen. fcw stomach content analyses suggest that In southern Oceania there are three spe- pelagic tunicates are the dominant plank- cies: G. watanabei, G. semicinctus, and tonic animals consumed; benthic material G. spin us (the last two are an allopatric such as polychaetes, bryozoans, and algae pair) . The only record from northern have also been found in some stomachs, Oceania is G. personatus from Hawaii. however) . In contrast, the species of The apparent absence of the genus from Holacanthlls, Pomacanthus, Pygoplites, and the broad central region of the tropical Euxiphipops fecd mainly on sponges, and Pacific is somewhat puzzling. Some of the Centropyge on algae and detritus (Randall, island groups such as the Marshall Islands, 1967; Randall and Hartman, 1968; Randall, Mariana Islands, and Samoa Islands have MS). been well collected. At Enewetak in the In view of the morphological similarity, it Marshalls the author and associates have is plausible that Genicanthus has been de- made numerous deep dives, but no Geni- rived from Holacanthlls. The differences canthus have been sighted. Since G. wata- betwecn these two genera, such as the more nabei is known from northern latitudes elongate body, emarginate to lunate caudal (Japan and the Ryukyus) and southern fin and smaller teeth of Genicanthus, may be (Society Islands, Austral Islands, New Cale- related to the feeding by the species of donia and Osprey Reef), it would seem to Genicanthlls on zooplankton, with the con- be a good prospect to occur at intervening comitant need for greater mobility as they localities in Oceania. Its most tropical loca- venture well above the substratum. tion, however, has been at Osprey Reef If the postulated derivation of Genicanthus (14°S). is true, then G. personatus of the Hawaiian Specimens of Genicanthus were examined Islands may be the most primitive living at the Bernice P. Bishop Museum in Hono- representative of the genus. With its greater lulu (BPBM), British Museum (Natural body depth, caudal fin of least emargination, History) in London [BM(NH)], California and XIV dorsal spines, it seems closer to Academy of Sciences in San Francisco Holacanthus than any other species of Geni- [( CAS) and former Stanford University canthus. material (SU)], Museum National d'Histoirc It has long been noted that the Indo- Naturelle in Paris (MNHN), Rijksmuseum Malayan region has more species of most van Natuurlijke Historie in Leiden (RMNH), genera and families of tropical Indo-Pacific and Zo61ogischMuseum at Amsterdam. The marine fishes than the Indian Ocean to the specimens at the U.S. National Museum of west and the Pacific to the east. Our present Natural History at Washington, D. C. (US- knowledge of the distribution of Genicanthus NM) were unavailable due to being on loan. is consistent with this. Four species occur Specimens have been sent on loan by the in the Indonesian-Philippine area: G. la- Department of Zoology, University Museum, 396 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

University of Tokyo (ZUMT) and the Acad- diameter 2.4 to 4.3 in head; interorbital con- emy of Natural Sciences of Philadelphia vex, the width 2.6 to 3.6 in head; caudal (ANSP) , peduncle about as deep as long, the least Lateral-line scale counts were recorded depth 1.8 to 2.3 in head. Mouth smaU, ter- only for the types of the new species. The minal, the gape angling slightly downward, counts of the diagonal scale rows between the maxillary ending approximately beneath the upper end of the gill opening (disregard- anterior nostril; teeth setiform, in 3 to 4 rows ing the small scales in this region) to the in jaws, the tips tricuspid (central cusp mid-base of the caudal fin are more reliable. notably the largest); teeth relatively short Even then, there is often some irregularity for the family, their length 4.2 to 6.8 in eye in the scale rows anteriorly and posteriorly of adults; anterior nostril with a fleshy rim which may make counting difficult. When which expands to a pointed flap dorso-poste- the last two dorsal or anal rays share the riorly. Gill membranes narrowly attached to same basal element, they are considered as isthmus; branchiostegal rays 6; gill rakers one ray. Pectoral-ray counts include the short, 15 to 18 on first arch. No spines on upper rudimentary ray. Upper-limb gill- operclc (corners of opercle above pectoral raker counts are listed first; the raker at the base and at upper end of gill opening ob- angle is included in the lower-limb count. tusely rounded); a prominent spine at angle Standard length (SL) was taken from the of preopercle which may extend slightly pos- most anterior point of the upper lip to the mid- terior to a vertical at hind edge of opercular base of the caudal fin (end of hypural plate), membrane, the spine length 1.85 to 4 in head even though preorbital spines may extend (may be shorter in juveniles); upper pre- anterior to the lips. Depth was measured opercular margin serrate, lower margin with from the lowest point of the abdomen to the 1 to 5 small spines; subopercIe and anterior extreme base of the dorsal spine vertically edge of interopercle nearly always with small above it. Width of body was measured im- spines (some, especially those on subopercle, mediately posterior to the gill opening. detectable only with microscope); exposed Caudal concavity is the horizontal distance upper posterior edge of supracleithrum ser- between perpendiculars at the posterior ends rate; preorbital with 4 to 16 small spines, of the longest and shortest caudal rays. Cau- one or more of which may be enlarged; pre- dal peduncle length is the diagonal measure- orbital with semicircular notch or emargina- ment from mid-base of caudal fin to the tion on ventroanterior edge; posterior mar- rear base of the anal fin. Stripes are hori- gin of preorbital free for about one-third to zontal color bands, and bars are vertical one-half the distance from tip of lowermost markings. spine to eye. Scales longitudinally ridged and Characters common to all species of the strongly ctenoid; head, body, and fins entirely genus are not repeated in the species ac- scaled except lips to a varying degree, the counts. Data in parentheses in the descrip- spinous portion of dorsal and anal fins above tions of new species apply to paratypes when the basal scaled sheath, and the pectoral fins different from the holotype. except basally; scales on head and fins much smaller than those on body; diagonal scale Genus Genicanthus rows from upper end of gill opening to mid-

GCllicalltl/llS Swainson, 1839: 170, 212 (type base of caudal fin 45 to 54; approximate species, Holacallthus lamarck Lacepede, by vertical rows of large scales on opercle 6 to 8; subsequent designation, Jordan, 1919). some auxiliary scales present on nape, Description.-Body moderately deep, the thorax, lateral-line series, and partially on depth J.8 to 2.4 in SL, and compressed, the abdomen and back above lateral line for width 1.8 to 3.6 in depth; head length 3.4 to some species; lateral line complete, though 4,2 in SL; snout 2.6 to 3.6 in head; eye poorly developed in descending part beneath RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 397 soft portion of dorsal fin; lateral-line scales branes incised one-half or more length of spines; preorbital of adults with a stout with 2 diverging tubules (a single tubule in spine which may project anterior to lips; descending and peduncular part). Two pairs males with 8 to 13 black bars which do of prominent pores in lower lip. Dorsal not reach ventrally (except on caudal peduncle of G. spil1us); females without rays XIV or XV (one of 83 specimens with dark markings on head or body (though XV!), 16 to 19; pectoral rays 16 or 17 (one upper half of G. semicil1c/us is blackish) _ 8 with 15, and one with 18) (usually upper 3a. Caudal concavity of adults 1.55 to 5 in SL; males with or without dark bars (if 2 and lowermost unbranched); pelvic rays present, they extend onto ventral part of T,5; principal caudal rays 17 (upper and body); females without a black bar across lower unbranched); dorsal spines progres- caudal base ._ __ 4 3b. Caudal concavity of adults 4.0 to 6.4 in sively longer, the last 1.2 to 1.8 in head; SL; males with about 21 irregular narrow longest dorsal soft ray 3.1 to 6.4 in SL; first dark bars on upper half to two-thirds of 2 to 4 interspinous membranes of dorsal fin body; females with a large black triangle above each eye which are connected by incised one-half or more length of spines; a narrow black band across upper inter- third anal spine 1.1 to 1.6 in head; caudal orbital, broad blackish edges on upper pre- opercular and opercular margins separated fin varying from emarginate to lunate with by a whitish zone, and a broad black band produced lobes, the caudal length 1.15 to across base of caudal fin which is continu- 3.3 in SL, the caudal concavity 1.55 to 14 in ous with black on caudal lobes (southern Japan to Philippines) G. semifasc;alUS SL; pectoral fins 1 to 1.3 in head; pelvic 4a. Anal soft rays modally 16 or 17; males fin origin below lower pectoral base; pelvic with black or orange-yellow stripes on body fins often ending in a filament which reaches (females of G. lamarck also with dark stripes and females of G. belllis with diag- at least to anus and often slightly beyond onal black bands) 5 origin of anal fin. One predorsal bone. 4b. Anal soft rays modally 18, males with numerous dark bars, which extend to ven- Vertebrae 10 + 14. tral part of body, and a prominent black spot mid-ventrally on thorax in front of Key to the Species of Gel1ical1/hlls base of pelvic fins; females wi:hout dark markings on body except for anterior exten- la. Dorsal spines XV (two of 82 with XIV, sion of black band of each caudal lobe 7 one with XVI); depth of body (of speci- 5a. Basal one-half to three-fourths of upper lip mens 100 mm or more in SL) 1.9 to 2.43 scaled; first 4 dorsal interspinous mem- in SL; caudal fin moderately cmarginatc branes of adults incised one-half or more to lunate with produced lobes, the caudal length of spines; males and females with concavity of adults 1.55 to 8 in SL; eighth 3 to 5 horizontal black stripes on side of to tenth dorsal soft rays the longest; color body; no black margin on anal fin; caudal of females not as in 1b 2 fin with small dark spots except upper and Ib. Dorsal spines XIV; depth of body 1.77 to lower margins which are whitish in males 1.83 in SL; caudal fin emarginate, the and black in females; pelvic fins of males caudal concavity about 14 in SL; first 3 black, of females whitish (tropical western dorsal soft rays the longest; anterior head Pacific and Indian Oceans) G. lamarck except lips and all but posterior part of 5b. Lips not scaled; first 3 dorsal interspinous caudal fin black, in contrast to whitish membranes incised one-half or more length On rest of head and body; pelvic fins of spines; black stripes on body, if present, yellow orange (female; color of male 8 to 13 (or there may be 3 diagonal black not yet known) (Hawaiian Islands) _ bands); black margin on anal fin (except ______G. persona/lis, n. sp. male of G. bel/us); caudal fin without dark 2a. Scale rows from upper end of gill opening spots (except faintly on some females of to caudal base 45 to 48; first 3 or 4 dorsal G. be llus ) 6 interspinous membranes incised one-half or 6a. Males with 8 to 13 black stripes on about more length of spines; preorbital spines lower two-thirds of body; females without usually not extending anterior to lips; pec- dark markings on body but with black toral rays usually 16 (except G. bellus); transverse bands on interorbital and dor- dark bars, if present on males, more nu- sally On snollt and a black band in caudal merous than 12; females with One or more lobes (Ryukyu Islands, Amami {) Shima dark markings on head (except G. mel- Islands, Society Islands, Austral Islands, al10spi los) 3 Pitcairn Island, New Caledonia, and Os- 2b. Scale rows from upper end of gill opening prey Reef) . . G. wll/al/abei to caudal base 49 to 54; pectoral rays 6b. Males lavender gray with a mid-lateral usually 17; first 2 dorsal interspinous mem- orange-yellow stripe (pale in alcohol) and 398 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

an orange-yellow band along back at base on upper margin of preopercle 12 to 18; of dorsal fin; females with 3 diagonal dark bands on body (lowermost blue in life), depth of body 1.9 to 2.2 in SL; interorbital the middle one continuous with black band space 3 to 3.3 in head; caudal concavity 2.5 of lower caudal lobe, and 2 black bars to 4 in SL; longest dorsal soft ray 4.3 to 5.5 on head, the posterior one curving on nape to join broad black band at dorsal in SL; first 4 dorsal interspinous membranes base (Society Islands and Cocos-Keeling incised one-half or more length of spines; Islands) G. bel/us, n_ sp_ basal one-half to three-fourths of upper lip 7a. Longest dorsal soft ray 3.1 to 3.7 in SL; males without a large black area basally scaled. in dorsal fin; females without a dark bar Color in alcohol pale with 3 to 5 black dorsally on head; upper black caudal band stripes which continue onto head where they of females ending anteriorly on caudal peduncle (tropical western Pacific) _ converge and terminate at posterior or ven- ______G. me!anospilos tral edge of eye (fourth stripe, if present, may 7b. Longest dorsal soft ray 4.3 to 5.5 in SL; not reach head, may end on cheek, or ex- males with a large elongate black area basally and centrally in dorsal fin; females tend anterior to eye; fifth stripe, when pres- with a black bar from upper edge of eye ent, confined to lower or posterior part of to nape; upper black caudal band of adult body); spinous portion of dorsal fin and all females continuing onto back beneath soft portion of dorsal fin (tropical western In- but lower posterior part of soft portion black dian Ocean and Red Sea) G. caudovittatus (except margin which is narrowly pale); 8a. Interorbital space 2.84 to 3.1 in head caudal fin, posterior caudal peduncle, pale length; longest dorsal soft ray 3.9 to 4.75 in SL (110 to 195 mm SL); males light posterior part of dorsal fin, and soft portion bluish with about 12 black bars on upper of anal fin with small black spots. Males half of body, the unmarked lower half with black pelvic fins and no black band in as well as most of dorsal and anal fins and lobes of caudal fin dull yellow orange caudal lobes. Females and juveniles with (broad median part of caudal fin spotted pale pelvic fins and a pale-edged black band with black); females with upper half of in each caudal lobe; on some individuals the body blackish, lower half whitish, the caudal lobes broadly black (Lord Howe upper body stripe (which tends to be broader Island) G. semicinctus in juveniles and females) curves downward 8b. Interorbital space 2.6 to 2.74 in head posteriorly to join the black band of the length; longest dorsal soft ray 4.55 to 6.4 in SL; males light bluish with 8 to 12 dark lower caudal lobe. bars on upper half to two-thirds of body, In life the back above the upper black reaching ventrally only on caudal peduncle stripe is blue gray, the sides bluish white; and caudal base; females light bluish with- out dark markings (Pitcairn Group and males with a yellow area on nape. Herre Austral Islands) G. spinus, n. sp. and Montalban (1927) described the color of a Philippine specimen (male form) as Genicanthus lamarck (Lacepede, 1802) violet brown above, merging into sea green Figures 1-3 or blue green on sides, pale below, with a dilute roseate flush over all, becoming light Ho!acanthus lama/'ck Lacepede, 1802: 526, 530 (no type locality; from the Dutch collection). rose on breast and cheeks; 5 longitudinal Ho!acanthus !ama/'cki japoniclls Schmidt, 1930: black bands; a yellow spot on nape; eye 99, pI. Y, fig. 2 (type locality, Homan, Okinawa). yellow. Bleeker (1877, pI. 371, fig. 4) and Ho!acanthlls c!wpmani Herre, 1933: 19 (type Herre and Montalban (1927, pI. 16) have locality, Dumaguete, Oriental Negros, Philip- pine Islands). illustrated the male phase in color. Diagnosis.-Dorsal rays XV (one of 12 with Remarks.-Schmidt (1930) described the XIV),15 or 16, anal rays III,16 or 17 female form of G. lamarck from Okinawa as (usually 17); pectoral rays 16 (one with the subspecies japonicus. Herre (1933) 17); scale rows from upper end of gill named the female of G. lamarck a species, opening to caudal base 45 to 47; gill rakers G. chapmani. Fraser-Brunner (1933) fig- 4 + 12 (one of nine with 5 + 12); serrae ured and provided a key to four "varieties" RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 399

Figure 1. Gel/ical/lhll~' lamarck, male, 114 mm SL, Guadalcanal, Solomon Islands, BPBM 16097. of lamarck, two of which he gave new variety ensis" phase the upper black stripe is not names: trilineatus and amboinensis. His uniformly dark; one can see within the varieties lamarck and trilineatus display the lightly pigmented broad stripe a narrower male color pattern with 4% and 3 black darkly pigmented one. The narrower black stripes, respectively. The remaining two, part is straight whereas the broad dusky japonicus and amboinensis, are female forms, portion descends posteriorly across the lower the former with 3 black stripes and the latter stripes to join the lower black caudal lobe with 4. On some individuals in the "amboin- band. Fraser-Brunner has shown the broad

Figure 2. Gel/leal//fllls lamarck, female, 94 mm SL, Guadalcanal, Solomon Islands, BPBM 16097. 400 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

record is from Smith (1955) who took a juvenile at Shimoni, Kenya; he also observed the species at Aldabra. The identification of the Kenya fish was made in the field; the specimen was lost enroute to Rhodes Univer- sity. Gerald R. Allen collected a male speci- men (BPBM 13658, 147 mm SL) at Samarai, Papua-New Guinea in 10.5 m, which is unusually shallow for the species. The author collected four specimens at Guadalcanal, Solomon Islands, in 31 to 35 m (BPBM 16014, 54 mm SL; BPHM Figure 3. Gellicallihus lamarck, juvenile, 54 mm SL, Guadalcanal, Solomon Islands, BPBM 16014. 16097,94-114 mm SL). Genicanthus watanabei (Yasuda and dusky band in D of Text-fig. 18 by a stipple Tominaga, 1970) pattern. It is possible that this phase is Figures 4 and 5 exhibiting the first change in color from the Holacallihus caudovilallls Watanabe (11011caudo- female to the male; i.e., that the lightly villallls Giinther), 1949: 40, fig. 4 (Zamami, pigmented part of the stripe is in the Ryukyu Islands). process of disappearing. If a histological Holacallllllls watallabei Yasuda and Tominaga, 1970: 144, figs. 3,4,6,7, 11, and 12 (type examination were made of the gonad of such locality, Onna Beach, Okinawa-jima, Ryukyu an individual, it might show regressing Islands) . ovarian and developing testicular tissue. Diagnosis.-Dorsal rays XV (one of 14 spec- The holotype of G. lamarck was examined imens with XVI), 15 or 16 (usually 16); anal at the Museum National d'Histoire Naturelle rays III,14; pectoral rays 16; scale rows in Paris (MNHN AlO047, 123 mm SL), as was the holotype of G. chapmani at the Cali- from upper end of gill opening to caudal base fornia Academy of Sciences (SU 25504, 45 to 48; gill rakers 4 or 5 + 13 (one of ten 113.5 mm SL). with 12); serrae on upper margin of pre- G. lamarck is known in the literature from opercle 21 to 37; depth of body 2.05 to 2.4 Indonesia, Philippines, Ryukyu Islands, in SL; interorbital space 3.0 to 3.3 in head; southern Japan, and East Africa. The last caudal concavity 1.55 to 3.5 in SL; longest

Table 1. Fin-ray counts of the species of Gen;canthus

Dorsal Rays Anal Rays Pectoral Rays Species XIV XV XVI 15 16 17 18 ITI IV 16 17 18 19 15 16 17 18 lamarck 11 5 7 12 2 to 11 watanabei 13 9 5 14 14 14 bellus 3* 2 2 4 2 2 2 2 melanospilos 15 1 10 4 15 1 14 13 1 cal/dovillatlls 16 3 11 3 17 4 12 15 2 semifasciatlls 7 5 2 7 7 6 1 semicinctus 5 5 5 4 1 4 spinus 9 4 5 9 7 2 8 personatlls 4 3 3 3 4 • One with abnormal spinous portion of dorsal fin. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 401

Figure 4. Gellicalltlllls watanabei, male, 114 mm SL, Tahiti, Society Islands, BPBM 10804. dorsal soft ray 5.25 to 6 in SL; first 3 dorsal 13 black stripes which are slightly narrower interspinous membranes incised one-half or than pale interspaces (2 or 3 stripes angle more length of spines; lips not scaled. downward and broaden into basal portion of Color of males in alcohol: head, thorax, anal fin); a pale stripe of variable length and upper part of body brown; a broad zone (which may be broken) posteriorly in brown of lower part of body beginning anteriorly region of body just above black-striped zone; below lateral line and posteriorly on lower caudal fin light brown, the lobes a little third of caudal peduncle whitish with 8 to darker than rest of fin; dorsal fin black except

Figure 5. Genieanthus watanabei, female, 98 mm SL, Tahiti, Society Islands, BPBM 10803. 402 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975 lower posterior part which is brown below a SL; BPBM 16755, 107 mm SL) and one horizontal demarcation and the margin which from Raivavae, Austral Islands (BPBM is narrowly pale; anal fin with a pale margin, 12784, 128 mm SL). Gerald R. Allen and a broad black submarginal zone and a basal Walter A. Starck, II collected two at New section which is irregularly striped as on body Caledonia (AMS I.17512-002, 92 mm SL; but the black bands broader than pale inter- AMS uncat., 96 mm SL) and three at Osprey spaces; paired fins pale. Reef, Coral Sea (AMS 1.16513-001, 102- Females brown with a black bar across 118 mm SL). The specimens from Polynesia mid-interorbital space, a large black spot were taken by spearing in the depth range of nearly crossing anterior interorbital space, an 37 to 44 m. inverted "U"-shaped black mark on front Theholotype (ZUMT 52421,89 mmSL), of snout, the free ends of which cross upper a female, is at the University of Tokyo. lip, and some irregular black spots dorsally on head behind interorbital black bar; upper Genicllnthus bellus, New Species opercular membrane blackish; a prominent Plate 1; Table 2 black band in each lobe of caudal fin, the GenicGnthus sp. Randall, 1973 : 194 (Tahiti). rest of fin light brown basally shading to pale distally; dorsal and anal fins with a Holotype.-BPBM 16840, 105.7 mm SL, male, Tahiti, Society Islands, outside barrier reef off narrow pale margin and a broad submarginal district of Paea, vertical drop-off in 60 m, dip zone of black; paired fins pale, the upper nets, Clemens Classen, September 22, 1973. pectoral base with a blackish area which is Paratypes.-BPBM 16841, 95.7 mm SL, female, continuous with a larger black area in axil. same data as holotype; ANSP 127544, 2: 101.8 In life both sexes are light blue or bluish and 84.6 mm SL, male and female, respectively, gray, the back of the male a deeper blue than Cocos-Keeling Islands, Indian Ocean, north side of Turk Reef, 60° slope, spear, William F. Smith- the pale interspaces of the striped zone; Vaniz, March 9, 1974. uppermost black stripe of male continuous Description.-Dorsal rays XV,16 (15 or posteriorly with a bright yellow stripe of 16); anal rays III,17 (16 or 17); pectoral variable length on different individuals; cau- rays 17 (16 or 17); scale rows from upper dal lobes of male deep blue; black markings end of gill opening to caudal base 48 (46 to dorsally on head of female edged in bright 48); diagonal row of large scales above blue. lateral line to base of first dorsal spine about Remarks.-G. watanabe; is reported only 10; diagonal rows of large scales below from the Ryukyu Islands and the Society lateral line to base of first anal spine about Islands (Randall, 1973). The latter record 24; circumpeduncular scales about 27; gill is based on five specimens from Tahiti (BP- rakers 4 + 13 (4 + 12 to 5 + 12); vertebrae BM 10803, 86.5-98 mm SL; BPBM 10804, 10 + 14 (holotype only). 114 mm SL; BPBM 12784, 126 mm SL) Depth of body 2.12 (2.16-2.17) in SL; and two from the atoll of Tetiaroa (BPBM width of body 3.05 (2.68-3.05) in depth; 14982, 128-129 mm SL). Dr. Tominaga head length 3.64 (3.57-3.74) in SL; snout loaned a male specimen, 152 mm SL, taken length 3.2 (3.4 to 3.48) in head; diameter at Amami 0 Shima. 1he author has col- of eye 3.17 (3.13-3.15) in head; inter- lected, in addition, three specimens at Pit- orbital width 3.34 (3.2-3.28) in head; length cairn Island (BPBM 16449, 109-138 mm of preopercular spine 2.12 (2.07-2.39) in

~ Plate I. Holotype of Genican/IIUS persona/us, female, 91 mm SL, Oahu, Hawaiian Islands, BPBM 15509 (upper); Holotype of G. bel/us, male, 105.7 mm SL, Tahiti, Society Islands, BPBM 16840; lower caudal filament removed by another fish during sojourn in an aquarium (middle); Paratype of G. bel/us, female, 95.7 mm SL, Tahiti, Society Islands, BPBM 16841 (lower). RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 403 404 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Table 2. Measurements of the type specimens of Genicantlllls be/lus expressed as thousandths of the standard length

HO]Olype Paralypes BPBM 16840 BPBM 16841 ANSP 127544 Standard length (mm) 105.7 95.7 101.8 Greatest depth of body 472 463 442 Width of body behind gill opening 154 151 165 Head length 277 277 267 Snout length 86 80 79 Diameter of eye 87 89 85 Interorbital width 82 88 82 Length of preopercular spine 130 117 128 Least depth of caudal peduncle 131 131 130 Length of caudal peduncle 131 114 134 Snout to origin of dorsal fin 337 342 324 Snout to origin of pelvic fins 360 350 337 Snout to origin of anal fin 615 610 590 Length of caudal fin 748 450 472 Caudal concavity 521 235 252 Length of pectoral fin 236 254 233 Length of pelvic fin 289 279 275 Length of pelvic spine 173 188 174 Length of dorsal fin base 705 730 686 Length of first dorsal spine 76 75 85 Length of second dorsal spine 109 123 127 Length of third dorsal spine 144 153 147 Length of last dorsal spine 182 181 173 Length of longest dorsal soft ray 234 207 234 Length of anal fin base 337 343 334 Length of first anal spine 95 97 96 Length of second anal spine 147 151 154 Length of third anal spine 205 213 201 Length of longest anal soft ray 357 324 250 head; least depth of caudal peduncle 2.11 gin of supracleithrum and posttemporal (2.06-2.14) in head; caudal fin lunate with bones serrate. filamentous lobes, the caudal concavity 1.92 Lips unsealed, the outer surface and edges (2.97-4.25) in SL; length of pectoral fin 1.16 relatively smooth. Dentition typical of the (1.1-1.2) in head; last dorsal spine 1.51 genus, the upper teeth 6.3 in eye and the (1.54-1.55) in head; longest (the ninth) lower teeth 4.7 in eye. Longest gill raker dorsal soft ray 4.26 (4.28-4.83) in SL; third 5.8 in eye. anal spine 1.34 (1.31-1.32) in head. Lateral line complete, the pored scales Serrae on upper preopercular margin 15 44 (42 to 44); lateral-line scales with aux- (14 to 18); small spines on lower margin of iliary scales; auxiliary scales also present on preopercle 2 (2 or 3); small spines on lmvcr thorax, nape, and on scattered scales of ab- anterior margin of interopercle 3 (2); small domen and back above lateral line. Scales on spines on margin of subopercle 7 (0 to 8); mid-side of body of holotype with 36 to 40 spines on pre orbital 5 (5 to 7), the most ridges, ending in as many ctenii. anterior spine enlarged, directed horizontally Origin of dorsal fin above upper end of forward, and overlapping basal part of upper gill opening; origin of anal fin beneath base lip; a broad shallow notch in ventroanterior of eleventh dorsal spine; origin of pelvic border of preorbital; exposed posterior mar- fins beneath lower edge of pectoral base; RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 405 pectoral fins reaching a vertical at anus; orange-yellow band at dorsal base continuing filamentous tips of pelvic fins reaching to or as a brown band onto nape, ending at front slightly beyond origin of anal fin. of interorbital; a separate brown band pass- Color in alcohol light gray brown, the ing from front of eye to mid-base of upper edges of the scales darker than centers, with lip; a small brown spot behind and slightly a pale stripe about as broad as eye diameter above eye; edge of eye and margin of upper passing from upper end of gill opening to lip blue; lobes of caudal fin blue, the rest below mid-line of caudal peduncle (body of the fin light blue with elongate irregular darker above stripe than below); a second orange-yellow spots; dorsal fin above orange- pale band passing along base of dorsal fin yellow base bluish gray, the outer part of the which breaks into large irregular spots as it first few interspinous membranes mainly dull reaches and merges with lateral stripe; a dark orange yellow, the rest of spinous portion brown band passing diagonally upward and with two indistinct rows of diffuse dull posterior from above front of eye, becoming orange-yellow spots (only one large spot on pale-edged on nape, and narrowing and dis- first four interspinous membranes), the soft appearing into pale dorsal-base stripe be- portion with diagonal brighter orange-yellow neath first dorsal spine; a small dark spot stripes, one of which is continuous with mid-dorsally on snout at level of upper edge basal orange-yellow band and another with of eye and a second less distinct one above lowermost row of yellow spots in fin; nar- and posterior to eye; nostrils in a dusky band rower interspaces between diagonal posterior running anteriorly from eye; upper lip black- yellow bands in dorsal fin light blue; margin ish with a pale medial spot on upper margin; of fin light blue; anal fin grayish blue, shad- lower lip dusky especially near margin; ing outwardly to blue; paired fins whitish, the caudal fin pale, the lobes brown, the median axil of pectorals yellow. central portion with longitudinally elongate The 95.7-mm female (PI. 1) paratype col- brown spots with lighter centers (some ir- lected with the holotype was bluish white in regular and interconnected) ; dorsal fin gray- life with a curved black band broadly edged in ish brown with a large indistinct pale spot bluish white running from pectoral base and on each interspinous membrane about one- axil across opercle and nape, continuing as a third distance from base of fin, this row of broad band at dorsal base; a broad diagonal spots joining a pale band in lower fourth of black band originating in the curved black bar soft portion of fin, ending on eighth soft ray; on nape and passing to black band in lower soft dorsal posterior to eighth ray with three caudal lobe; a narrower dark brown band diagonal pale bands, the two posterior ones originating in pale interspace between above- dark-edged; distal triangular part of each mentioned diagonal band and band at base interspinous membrane pale; below this a of dorsal fin below sixth dorsal spine and dark band; anterior soft portion of dorsal fin passing to rear base of dorsal fin; a broad with a narrow pale margin and a dark sub- diffuse diagonal blue streak passing from marginal band; anal tin brownish gray; paired upper end of gill opening, beneath pectoral fins pale, the rays slightly dusky, the pectorals fin, and ending on lower body above anterior with a large pale spot at base. soft portion of anal fin; a broad black bar In life the holotype (PI. 1) was lavender edged in bluish white passing diagonally gray on head, thorax, and body above mid- upward from eye to nape; head except for lateral stripe, nearly white below; lateral pale-edged bars lavender brown; eye nearly stripe and band at base of dorsal orange rimmed with pale bluish; nostrils in a small yellow, edged in bluish white, as are the orange-yellow area, and some orange yellow irregular orange-yellow spots on caudal pe- at corner of mouth; upper lip with three duncle between posterior ends of the stripes, black bars separated by blue; a broad black and the orange-yellow spot at pectoral base; band, edged in pale blue, in each caudal lobe; 406 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975 rest of fin grayish blue with a few small go to the wall." He added that the dominant faint dusky spots; black band at dorsal base coral on the wall is the "pancake" or "saucer" broadens posteriorly into middle of soft por- type. tion of fin, its posterior edge rimmed with William F. Smith-Vaniz collected several bluish white; fin dusky blue-gray posterior to specimens at Cocos-Keeling besides the two band; main portion of fin above brown basal listed herein as paratypes; however, only the band dark orange brown with a bluish white two were available in time for this study. margin; anal fin grayish blue with a blue mar- The others were shipped by surface from gin, a broad submarginal band of yellow Cocos-Keeling to Philadelphia. orange, and a blue band below this; pectoral fins whitish, the rays a little dusky, the Etymology.-This species is named bel/us brown at the base containing two dull orange- from the Latin for beautiful, as it is the yellow spots, one above the other; pelvic fins most colorful of all the species of the genus. bluish white. The two paratypes from Cocos-Keeling Genicanthus melanospilos (Bleeker, 1857) Islands differ slightly in color from the two Figures 6-8 Tahitian type specimens and in having 15 Holacanlhus melanospilos Bleeker, 1857: 56 instead of 16 dorsal rays, and 16 instead of (tyre locality, Ambon, East Indies). 17 pectoral rays. The male from Cocos- Genicanlhus macclesfieldiensis Chan, 1965: 330, figs. 4, 5 (type locality, Macclesfield Bank, Keeling lacks the pale bluish borders on the South China Sea). yellow stripes, the spinous portion of the Diagnosis.-Dorsal rays XV,15 to 17 (usu- dorsal fin is yellow except for the margin, ally 16); anal rays III,18 (one of fifteen and the large orange-yellow spots on the with 17); pectoral raY5 15 to 17 (usually caudal peduncle are absent. The female 16) ; scale rows from upper end of gill open- (which has a deformed head and anterior ing to caudal base 46 to 48; gill rakers dorsal fin) has a broader and darker blue 4 + 12 (one of eight with 3 + 13); serrae lower diagonal band on the body and more on upper margin of preopercle 15 to 40; orange submarginally in the dorsal and anal depth of body 1.9 to 2.15 in SL; inter- fins. orbital space 2.9 to 3.3 in head (3.55 for Remarks.-It is not certain if the differences 32-mm juvenile); caudal concavity 1.85 to mentioned above between Tahitian and 3.6 in SL (14.5 in 32-mm juvenile); longest Cocos-Keeling specimens represent geo- dorsal soft ray 3.1 to 3.7 in SL; first 3 graphical variation or whether these are dorsal interspinous membranes incised one- within the range of individual variation of a half or more length of spines; lips naked single island population. except basal half or less of upper lip which The linking of the very different male may have small scales. from the diagonally black-banded female Color of males in alcohol pale brownish was achieved largely by seeing an aquarium with about 14 to 16 dark bars On body specimen of the latter just beginning its (narrower than interspaces) which reach alteration to the male form at an aquarium of ventrally except on abdomen where they Coral Fish Hawaii in Honolulu shortly after approach the mid-ventral line (bars on its shipment from Tahiti. Unfortunately it caudal peduncle may be indistinct and may died before transformation was complete, be broken into spots); 7 to 9 transverse dark and the fish was not preserved. bars on nape, interorbital, and dorsally on The collector of the Tahitian specimens, snout; a black spot about the size of eye Clem Classen, observed that these fish "seem mid-ventrally on thorax immediately in front to prefer the straight vertical walls. They of base of pelvic fins; median fins light brown are usually out 20 feet from the wall feeding with numerous small pale spots, the dorsal in midwater. As soon as you approach, they and anal fins with a narrow dark margin and RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 407

Figure 6. GelliculllI/lls l1lelUllospi!os, male, 89.5 mm SL, Palau Islands, BPBM 9499.

a pale submarginal band (a second band may fin light bluish gray with numerous small yel- be present on anal fin); paired fins pale. low spots (those basally on fin larger, thus the Color in alcohol of females light brown, fin more yellow there tban bluish gray), the lighter on nape and antero-dorsally on body lobes with narrow pale blue margins; dorsal and darker on interorbital, dorsally on snout and anal fins light bluish gray with numerous and upper lip; scales of most of body with the small yellow spots, a narrow pale blue mar- centers paler than edges; a pale elliptical area gin, dark submarginal line, a yellow band middorsally on head from posterior interor- below this, and then an unmarked bluish-gray bital to origin of dorsal fin; a broad black band zone (anal and anterior dorsal spines without beginning dorsally at front of caudal peduncle marginal and submarginal bands); paired and passing to distal end of upper caudal fins light bluish. lobe; a comparable band ventrally beginning In life females and juveniles are yellow about mid-length of peduncle; margins of dorsally (the median dorsal region of nape caudal fin narrowly pale; broad central region bright yellow), light brownish gray on sides of fin pale (central caudal peduncle also and ventrally with a lavender cast, the scales light on some individuals); dorsal and anal with a yellow spot in center; head and thorax fins light dusky with numerous narrow pale lavender gray; edge of eye pale blue except diagonal bands, a narrow dark margin, and dorsoanteriorIy; nostrils orange yellow; lobes a pale submarginal band; paired fins pale. of caudal fin with a broad black band, edged In life the male of Fig. 6 was light bluish in light blue except for filaments, the bands over the body and interspaces of barred part carrying forward onto upper and lower part of head; unmarked part of head lavender gray; of caudal peduncle; rest of fin and caudal bars dark brown, the ends of those on body peduncle bluish white; dorsal fin with narrow yellow orange, the bars and spots on cau- diagonal bands of yellow and light bluish dal peduncle mainly yellow orange; large spot gray (overall color more yellow than gray), ventrally on thorax black; upper part of pre- except posterior half of soft portion which opercular spine, edge of eye, and nasal region is solid light bluish gray; anal fin light bluish light blue, the nostrils orange yellow; caudal gray with a few faint yellow spots anteriorly; 408 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Figure 7. Gellicalllhus melanospilos, female, 63 mm SL, Palau Islands, BPBM 9605. both fins with a pale blue margin, dark the two are readily distinguished by the blue submarginal line and a band of yellow longer dorsal and anal soft rays of G. mel- below these (except anteriorly); paired fins anospilos and by color. whitish. G. melanospiIos is known in the literature only from Indonesia and Macclesfield Bank, Remarks.-Fraser-Brunner (1933) was in South China Sea. The author has recently error in regarding Genicanthus caudovittatus collected specimens at the Palau Islands (GUnther) and G. semicinctus (Waite) (BPBM 9499, 89.5 mm SL; BPBM 9609, junior synonyms of G. melanospiIos. Admit- 6: 56-92 mm SL), Viti Levu, Fiji Islands tedly, the allopatric G. caudovittatus is (BPBM 11397, 61.5 mm SL), Alite Reef, closely allied to G. melanospiIos; however, Solomon Islands (BPBM 15608, 32 mm SL), and Madang, New Guinea (BPBM 15860, 75 mm SL) in the depth range of 24.5 to 60 m. Dr. Y. Tominaga sent three specimens from the Ryukyu Islands on loan which represent the first record for these islands (ZUMT 52783, 149 mm SL, from Kikai Jima; ZUMT 52823, 135 mm SL, from Ishigaki lima; and ZUMT 52824, 106 mm SL, also from Ishigaki). These arc unusual for their large size-significantly larger than any seen by the author in morc tropical localities. Figure 8. Genicalllhus melanospilos, juvenile, 32 Gerald R. Allen collected specimens at mm SL, Alite Reef, Solomon Islands, BPBM Uvea, Loyalty Islands (AMS I.l7513-002, 15608. 87 mm SL) and the New Hebrides (AMS RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 409

Figure 9. Gellieall/flUs eaudol'illa/tls, male, 108 mm Sl, Gulf of Aqaba, Red Sea, BPBM 13875. uncatalogued, 8: 66-100 mm SL). Pierre to 3.25 in head; caudal concavity 1.85 to Fourmanoir has informed the author that 4.8 in SL (6.7 in 40-mm juvenile); longest diver Pierre Laboute has observed G. mel- dorsal soft ray 4.3 to 5.5 in SL; first 3 anospilos outside the barrier reef at New dorsal interspinous membranes incised one Caledonia near Dumbea Pass in 25 to 50 m. half or more length of spines; lips naked Yasuda and Tominaga (1970) have except basal third of upper lip which has shown that the original spelling for this spe- small scales. cies is melanospilos, not melanospilus. Color in alcohol of males light brown with The holotype of G. melanospilos is at the about 17 dark bars on body (those on caudal Rijksmuseum van Natuurlijke Historic at peduncle faint and may be broken in spots), Lciden (RMNH 5812, ] 32 mm SL, 190 mm reaching or nearly reaching mid-ventral line; TL). The holotype of G. macclesfieldiensis about 9 dark bars on nape, interorbital and Chan is in the British Museum (Natural dorsally on snout; a black spot about size of History) [BM(NH) 1965.3.23.1, 59 mm eye mid-ventrally on thorax just in front of SL]. base of pelvic fins; caudal fin light dusky with numerous small pale spots and a narrow Genicanthlls caudovittatus (Gunther 1860) pale upper and lower margin except basally; Figures 9-11 dorsal and anal fins similar except dorsal which has a large black area in basal two- Holacanthus caudol'ittatus Giinther, 1860: 44 thirds of fin between about fourth spine and (type locality, Mauritius). Holaeall/fllls zebra Sauvage, 1891: 263, pI. 32, fourth soft ray (also these fins with a dark fig. 2 (type locality, Mauritius). submarginal line which is better developed on Holacanthus caudibieolor Sauvage, 1891: 267, dorsal than anal); paired fins pale. pI. 33, fig. 2 (type locality, Mauritius). Color in alcohol of females and juveniles Diagnosis.-Dorsal rays XV (one of 13 with light brown, the edges of scales darker than XIV),15 to 17 (usually 16); anal rays III, centers; a broad black bar above eye which 17 to 19 (usually 18); pectoral rays l6; angles slightly posterior as it passes to mid- scale rows from upper end of gill opening dorsal line; a broad black band in upper lobe to caudal base 45 to 47; gill rakers 3 to of caudal fin (upper margin pale except 5 + 11 to 13; serrae on upper margin of basally) which continues dorsally onto caudal preopercle 28 to 46; interorbital space 2.7 peduncle and then passes diagonally upward 410 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Figure 10. Gel/icantl1us caudovittatus, female, 72 mm SL, Gulf of Aqaba, Red Sea, BPBM 13878. beneath base of soft portion of fin where it soft portion light bluish gray with small gradually fades to ground color of body; yellow spots, except for the large area of lower lobe of caudal fin with a comparable black centrobasally in fin; anal fin light black band which extends to mid-length of bluish gray with small yellow spots; both ventral part of caudal peduncle; region of fins with margins pale bluish, a dark sub- caudal peduncle and fin between black bands marginal line (almost black on dorsal), a pale; posterior margin of fin narrowly black- band of yellow below this, and a narrow ish; dorsal and anal fins light dusky with a zone of blue gray without yellow spots; narrow pale margin, dark submarginal line, paired fins whitish. pale band beneath this, and some pale spots In life females are light lavender gray, the anteriorly; paired fins pale. centers of scales lighter than the edges; the In life males are light bluish, the bars dark supraocular and caudal bands are black; eye brown (black dorsoanteriorly) but contain- faintly rimmed with pale blue; fins pale ing some scales which are more yellowish lavender gray, the dorsal and anal with some than brown, the bars and spots on caudal faint yellow spots in about anterior half, a peduncle more yellow orange than brown; a bluish white margin, faint dark submarginal few brown spots may be present on head line and a pale yellow band proximal to below eye; nostrils in a narrow horizontal this. band of yellow; mid-ventral spot on thorax Remarks.-Sauvage (1891) attributed the inky black; unmarked head and thorax descriptions of Holacanthus zebra and H. lavender gray; caudal fin bluish gray, the caudibicolor to Lienard, citing Mem. Soc. lobes slightly darker than rest of fin, with d'Hist. Nat. Maurice (1832) as the reference numerous small yellow spots, a narrow pale (but without any page numbers). Evidently blue upper and lower margin (except basally) this was only a manuscript at best. Fraser- and a narrow yellow posterior margin which Brunner (1933), Weber and de Beaufort does not extend onto filamentous part of (1936) and Smith (1955) listed Sauvage lobes; spinous portion of dorsal fin yellow; (1891) as the author of these two species. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 411

specimens in the Museum National d'Histoire Naturelle.

Genicanthus sel17ifasciatus (Kamohara 1934) Figures 12 and 13

Ho/acallthl/s semijasciatus Kamohara, 1934: 459, fig. 2 (type locality, Kashiwajima, K6chi Prefecture, Japan). Holacalll/;u.I' jucasus Yasuda and Tominaga, 1970: 141, figs. 1, 13 [type locality, Sanbon-ne, Miyake-jima, Japan, (about 33°05'N, 139°45' E)]. Figure 11. GCllicolltfluS cOl/dovittalus, juvenile, 41 mm SL, Gulf of Aqaba, Red Sea, BPBM 13371. Diagnosis.-Dorsal rays XV,15 or 16 (usu- ally 15); anal rays III,17; pectoral rays 16 or 17 (usually 17); scale rows from gill Attempts to find descriptions of these poma- opening to caudal base 46 to 48; gill rakers canthids by Lienard in any publication on or 4 or 5 + 12 or 13; serrae on upper margin of about 1832 in Washington, D. c., Paris, and preopercle 21 to 32; depth of body 2.06 to Port Louis, Mauritius, have been fruitless. 2.16 in SL; interorbital space 3.1 to 3.5 in Weber and de Beaufort (1936) were in head; caudal concavity 4.0 to 6.4 in SL; error in placing Genicanthus zebra (Sauvage) longest dorsal ray 4.75 to 5 in SL; first 3 in the synonymy of Centropyge bispinosus dorsal interspinous membranes incised onc- (GUnther) . half or more length of spines; basal one-halE G. caudovittatus is known from Mauritius, to two-thirds of upper lip scaled; lower lip Red Sea, Malindi (Kenya), and Pinda naked except median basal region. (Mozambique) . The last two records are Color of male in alcohol light brown with from Smith (1955) who recorded (and illus- about 21 irregular narrow dark bars on trated) the male form as melanospilus and upper half of body anteriorly and upper the female as caudovittatus. The author has two-thirds posteriorly (some bars short, collected 11 specimens in the Gulf of Aqaba, others may anastomose) and another 5 Red Sea, and nearly as many from Mauritius or 6 on nape; rest of body and head lighter in the depth range of 27.5 to 46 m. brown with an even paler zone on head On June 6, 1972, in the Gulf of Aqaba at enclosing eye, running above pectoral base, 45 m an individual of caudovittatus was seen and continuing to middle of body; upper lip which was intermediate in color between the with a large median area of dark brown; male and female forms but was not collected. caudal fin light brown with faint spots; dorsal On the following day a barred male was ob- fin light brown with broken longitudinal served in apparent courtship with a female. brown bands, the outer portion of inter- He swam in front of her, lay on his side, and spinous membranes pale; anal fin light rapidly fluttered his caudal fin and long brown; paired fins pale. caudal filaments; the female did not respond. Color of females in alcohol light brown, The holotype of G. caudovittatus is a the edges of the scales darker than centers; stuffed specimen in the female color form. It a large triangular black area immediately measures 105 mm SL and is catalogued at above eye connecting at apex on nape with the British Museum (Natural History) under marking of other side via a short transverse number BM(NH) 1860.6.22.2. black band; black pigment continuing down- The holotypes of G. zebra (MNHN ward from posterior corner of triangle around AI0044, 141 mm SL) and G. caudibicolor posterior and ventral part of eye; a band of (MNHN AI0060, 105 mm SL) are alcoholic black along upper margin of preopercle; a 412 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Figure 12. Geniean/llus semifascia/Us, male, 132 mm SL, Motobu, Wakayama Prefecture, Japan. Figure 13. Gel/iean/hus semifasciallls, presumed female, Miyake Jima, Japan. black bar or vertically elongate spot poste- body to caudal peduncle yellowish brown riorly on opercle and adjacent scapular region (the scale centers yellow, the edges brown), above level of pectoral base; a broad whitish lower half and caudal peduncle white; head zone on opercle (between dark bars of pre- with a black triangular area above eye, with opercle and posterior opercle) passing dor- a ventral extension behind eye; a black band sally on nape adjacent to posterior edge of on preopercular edge and a large vertically black triangle; front of snout and lips slightly elongate black spot on upper opercular edge, dusky; black on posterior third of caudal these two black areas separated by a broad peduncle, base of caudal fin and curving white zone that extends dorsally adjacent to broadly onto caudal lobes; large centroposte- black triangle above eye; a bright blue line rior part of caudal fin whitish with irregular encircling eye; a yellowish region below blue vertical rows of blackish dots forming lines ring, continuous with black extension from in posterior half of fin; scaled portion of triangle above eye; nostrils in an orange-yel- dorsal and anal fins dusky, naked parts pale; low spot; caudal fin with a broad black band paired fins pale. across base continuous with black in lobes of Color in life of a presumed male from a fins, leaving a crescentic centroposterior 35-mm color transparency provided by Fujio region of white; dorsal fin brownish yellow, Yasuda (reproduced in black and white darker posteriorly, with a blue margin and a herein as Fig. 12): bars dark brown, the yellow submarginal band; anal fin gray; interspaces light brown; lower part of body paired fins whitish. white; a broad irregular zone of bright yellow Hiyama and Yasuda (1971: figs. 19 and orange on head and anteriorly on body be- 27) have illustrated the two forms of G. tween upper barred and lower white portions semifasciatus in color. (orange encloses eye except for a nearly com- plete light blue ring at edge confluent with Remarks.-G. semifasciatus was described a light blue band running anteriorly from by Kamohara (1934). The holotype was lower edge of eye); orange zone narrows as destroyed in a war-caused fire in 1945. In a it passes posteriorly, ending at about middle paper on his type specimens, Kamohara of body (but this color may be seen at the (1961) designated a specimen 225-mm in lower ends of the next 4 or 5 bars beyond total length as a neotype (No. 8495, Bio- the orange zone); caudal and dorsal fins logical Laboratory, Kochi University, Shi- light brown with brown-edged yellow-orange koku, Japan). spots (smaller on lobes of caudal which are a The author has examined only one 132- little darker than rest of fin); anal fin yellow- mm male specimen which is unquestionahly ish gray; paired fins whitish. this species; it was collected off Motobu, Color in life of a female (from a 35-mm Wakayama Prefecture, Honshu (33°45'N; color transparency sent by Fujio Yasuda, 129°39'E) in 100-200 m. reproduced herein as Fig. 13): upper half of It seems likely, however, that G. fucosus RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 413

Figure 14. GellicalltJllIs semicillc/lIs, male, 195 mm SL, Lord Howe Island, BPBM 14809.

Yasuda and Tominaga is the female form of end of gilJ opening to caudal base 49 or 50; G. semijasciatus. Six specimens of G. jucosus gill rakers 4 or 5 + 12 or 13 (usually 4 + scnt on loan by Tominaga are females, as is J 3); serrae on upper margin of preopercle the holotype (ZUMT 52422, 128.5 mm 15 to 25; interorbital space 2.85 to 3.05 SL).1 No differences other than color could in head (3.13 in 50-mm juvenile); caudal be found to distinguish G. semijasciatus concavity 2.6 to 7.9 in SL (9.5 in 50-mm from G. jucosus. The loss of the black bar juvenile); first 2 dorsal interspinous mem- on the head and the black band on the branes incised one-half or more length of caudal lobes and the development of bars spines; longest dorsal soft ray 3.9 to 4.75 on the head and body of caudovittatus are in SL; all but edge of upper lip and basal comparable to the changes which fucosus half of lower lip scaled; anterior spine of would undergo to transform into semifas- preorbital of adults enlarged, often projecting ciatus. The blue ring around the eye, though slightly in front of snout. not as intense in semifasdatus as the jucosus Color in alcohol of males: head grayish form, is a common color character. The brown, the preorbital and upper lip a little distribution of the two forms, as given by darker than rest of head; one or two faint Araga (1972), Japan to the Philippines, is bars on upper posterior part of head; body the same. light yellowish gray with 9 to 13 black bars of 1 to I liz scales in width running from Genicanthlls semidnctlls (Waite, 1900) scaled basal sheath of dorsal fin about two- Figures 14 and 15 thirds distance down side of body (most posterior bars reach farther ventrally), the Holt/call/hus semicillc/Us Waite, 1900: 204, pI. 36 (type locality, Lord Howe Island). first bar on nape to upper end of gill opening Diagnosis.-Dorsal rays XV,17; anal rays faint, the last below rear base of dorsal fin or Ill, 18 or 19 (usually 18); pectoral rays 17 anteriorly on caudal peduncle; in addition, 2 or J 8 (usually 17); scale rows from upper or 3 broken bars or vertical rows of 2 or 3 black spots on caudal peduncle; thorax

1After this paper was submitted for publication, Osamu and pectoral base with brown blotches; Okamura of Kochi University examined four specimens of G. semiJascialus 135 to 158 mm SL, including the 150-mm caudal fin light gray with small black spots neotype, and determined that all are males. He also examined two G. fucoslIs, 119 and 123 mm SL, which are except lobes which are light yellowish gray both females. The two G. fucasus and the smaller G. semi- fUlda/liS are in the collection of Kochi High School. edged in darker gray; dorsal and anal fins 414 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Figure 15. Gel/ical//llus semicil/c/us, female, 156.8 mOl SL, Lord Howe Island, BPBM 14809. yellowish gray with a very narrow pale outer pigmented, and the demarcation to the pale margin and dark submarginal line, the dorsal lower part is higher on the body (approxi- with 2 to 5 small black spots posteriorly; mately from upper end of gill opening to mid paired fins pale, the rays a little dusky. caudal peduncle). The curved pale band Color in alcohol of females: head (except on the head encloses a black hemispherical lower opercle, subopercle and interopercle), area about two-thirds size of eye above and dorsal fin, and upper portion of body black- adjacent to the dorsoposterior part of the ish (on upper half of body anteriorly and all eye. A 110-mm specimen (BPBM 14809) of caudal peduncle posteriorly); lower part also has the pale curved band enclosing an of head and body pale (demarcation on body, entirely black area, but this marking is smaller especially posteriorly, rather abrupt); upper relative to the eye size and nearly centered opercular membrane and adjacent shoulder over the eye. region next to upper end of gill opening rn life the males are light gray (almost black; a faint pale band encircling all but white) on the barred upper part of the body, lower part of eye and enclosing a zone of the blotched thorax and pectoral base region, black adjacent to upper part of eye, and one the posterior black-spotted part of the dorsal or two black spots above this; caudal fin fin and the broad median black-spotted por- with lobes broadly black (continuous with tion of the caudal fin; the lower part of the blackish posterior part of body), the centro- body, dorsal and anal fins, and lobes of the posterior portion of fin pale with blackish caudal fin are orange yellow; head orangish dots; anal fin light gray; paired fins pale brown, bluish over mouth and chin, yellow (pectorals may have a faint dark spot on orange in small area from eye to nostrils; upper base) . outer margins of dorsal and anal fins and A juvenile (BPBM 14859, 50 mm SL) upper and lower edges of caudal fin pale is similar to the female pattern but the dark blue with a dark submarginal line; paired upper part of the body is not as strongly fins light gray. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 415

In life females are blackish dorsally and rays 16 one side, 17 on other (17); scale whitish ventrally; the pale band enclosing the rows from upper end of gill opening to caudal black sector adjacent to upper eye is bright base 51 (49 to 54); diagonal row of large blue; the spinous portion of the dorsal fin is scales above lateral line to base of first dark dull orangish with a poorly defined dorsal spine about 8; diagonal row of large blackish band beginning basally at front of scales below lateral line to base of first anal fin and ending distally on last two inter- spine about 23; circumpeduncular scales spinous membranes; soft portion of fin dark about 26; gill rakers 4 + 12 (4 or 5 + 12 to orangish, the rays blackish; anal and paired 14) ; vertebrae 10 + 14 (holotype only). fins whitish, the pectoral rays a little dusky; Depth of body 2.38 (2.1-2.43) in SL; lobes of caudal fin black, the centroposterior width of body 2.75 (2.52-2.96) in depth; part of fin whitish with black dots. head length 4.09 (3.91-4.2) in SL; snout length 2.77 (2.74-3.2) in head; diameter of Remarks.-Fraser-Brunner (1933) and eye 4.27 (3.63-4.12) in head; interorbital others have been mistaken in regarding width 2.61 (2.63-2.74) in head; length of semicinctus as a synonym of melanospilos. preopercular spine 2.29 (1.85-2.96) in Waite described semicinctus from a single head; least depth of caudal peduncle 2.01 specimen in the male color form (AMS I. (1.91-2.07) in head; caudal fin lunate with 4345, 160 mm SL, 195 mm TL). filamentous lobes, the caudal concavity 1.59 The five specimens at the Bishop Museum (2.88-6.75) in SL; length of pectoral fin were collected by the author, Walter A. 1.16 (1. 0 1-1.17) in head; last dorsal spine Starck, II, and Barry Goldman on rock-coral 1.69 (1.47-1.74) in head; longest (eighth to bottom in the depth range of 23 to 46 m. tenth) dorsal soft ray 5.35 (4.57-6.4) in Known only from Lord Howe Island, but SL; third anal spine 1.47 (1.43-1.52) in might be expected at least from Norfolk head. Island. Serrae on upper preopercular margin 27 (11 to 35); small spines on lower margin of Genicanthus spinus, new species preopercle 3 (2 to 5) ; small spines on lower Figures 16 and 17, Table 3 anterior margin of interopercle 13 (4 to 21 ) ; fIo[otype.-BPBM 16450, 203 mm SL, male, Pit- small spines on margin of subopercle 8 (5 to cairn Island, reef off Bounty Bay, 33.5 m, spear, 11); spines on preorbital 5 (5 to 9), the John E. Randall, January 9, 1971. most anterior spine enlarged, directed hori- PlIratypes.-BPBM 16452, 2: 182.1 and 199 mm zontally forward, anterior to lips; a large SL, female and male respectively, Pitcairn Island, notch ventroanterior]y on preorbital; exposed small head of coral and coral rock off Bounty Bay, 38.5 m, spear, John E. Randall, December 26, posterior margin of supracleithrum serrate, 1970; BPBM 16754, 156.2 mm SL, female, Pitcairn but no clear margin externally visible for Island, reef on north side off Gannet Ridge, 44 m, posttemporal bone and no serrations present. spear, John E. Randall, December 28, 1970; USNM 211815, 198 mm SL, male, same data as Lips scaled except for narrow marginal preceding; ZUMT 52987, 194 mm SL, male, same zones. Dentition typical of the genus, the data as preceding; BM(NH) 1974.4.24.1, 187 upper teeth 6.1 in eye, the lower teeth 6.8 in mm SL, female, same data as preceding; CAS 30456, 178.5 mm SL, male, Pitcairn Island, same eye. Longest gill raker 3.8 in eye. locality as preceding, 40-44 m, spear and rotenone, Lateral line complete, the pored scales John E. Randall, Dean B. Cannoy, James R. Hay- wood, R. Richard Costello, J. David Bryant, and 45 (43 to 47); auxiliary scales present on Steven R. Christian, January 6,1971; BPBM 12775, dorsal and ventral thirds of body except 147.5 mm SL, male, Raivavae, Austral Islands, posteriorly. Scales on mid-side of body of olltside barrier reef south of Motu Hoa, 52 m, spear, Dean B. Cannoy and John E. Randall, holotype with about 64 ridges ending in February 25, 1971. ctenii. Description.-Dorsal rays XV,17 (16 or Origin of dorsal fin slightly posterior to 17); anal rays III,18 (18 or 19); pectoral upper end of gill opening; origin of anal fin 416 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975

Figure 16. Hololype of Gellicalltlws SpiIlUS, male, 203 mm SL, Pitcairn Island, BPBM 16450. beneath base of twelfth dorsal spine; pectoral fin (anterior bars reach half way down side fins not reaching a vertical at anus; origin of of body while the caudal peduncle and caudal pelvic fins beneath lower edge of pectoral base bars are complete; faint irregular bars base; filamentous tips of pelvic fins reaching dorsally on head posterior to eye; caudal fin slightly beyond anus (to base of second with lobes brown, edged in slightly darker anal spine on paratype with longest pelvics). brown, the broad median part of fin light Color of holotype in alcohol light brown brown with a slightly elongate dark brown with 11 dark bars (averaging 2 scales in spot in middle (greatest diameter of spot width) on body, the first from base of third about half diameter of eye) and 4 faint dark dorsal spine and the last at base of caudal spots near margin; dorsal and anal fins light

Figure 17. Paratype of Gellicallthus spinus, female, 182.1 mm SL, Pitcairn Island, BPBM 16452. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 417

Table 3. Measurements of the type specimens of Genicanlhlls spinlls expressed as thousandths of the standard length

Ho1otype Paratypes

BPBM 16450 BPBM 16452 BPBM 16452 BPBM 16754 ZUMT 52987 Standard length (mm) 203.0 199.0 182.1 156.2 194.0 Greatest depth of body 421 423 413 478 420 Width of body behind gill opening 152 153 162 161 165 Head length 244 253 252 220 247 Snout length 88 91 92 88 83 Diameter of eye 59 61 66 67 61 Tnterorbital width 94 93 92 97 93 Length of preopercular spine 117 137 93 88 91 Least depth of caudal peduncle 121 ]22 114 135 120 Length of caudal peduncle 127 126 135 141 127 Snout to origin of dorsal fin 304 306 328 333 301 Snout to origin of pelvic fins 332 337 335 340 340 Snout to origin of anal fin 6]7 620 634 590 623 Length of caudal fin 823 550 329 393 515 Caudal concavity 630 347 148 157 314 Length of pectoral fin 210 215 218 249 221 Length of pelvic fin 286 288 247 310 284 Length of pelvic spine 171 ]47 163 178 158 Length of dorsal fin base 686 685 670 728 702 Length of first dorsal spine 67 56 71 72 71 Length of second dorsal spine 90 102 104 105 109 Length of third dorsal spine 131 125 122 125 137 Length of last dorsal spine 144 147 153 168 142 Length of longest dorsal soft ray 187 178 168 208 185 Length of anal fin base 345 341 315 355 353 Length of first anal spine 71 74 69 80 70 Length of second anal spine ]]7 118 115 132 114 Length of third anal spine 167 166 164 179 168 Length of longest anal soft ray 218 184 181 224 209 - - ~------_.- brown with a narrow pale margin and black black bars, the unmarked lower part of the submarginal line, the dorsal with a small body a little darker than the pale spaces black spot at rear base of fin and 2 faint between bars on the upper part; dorsal part dark spots in median part of fin; paired fins of head posterior to eye with irregular dark pale brownish. gray bars alternating with light blue; inter- The number of dark bars on body of male orbital region and upper snout dark gray; paratypes vary between 8 and 12. head below eye, thorax, and anterior ab- Female paratypes are entirely light brown domen pale bluish with gray blotches; on in alcohol, the head, caudal lobes, and caudal cheek the ground color is light blue just base slightly darker; outer third of median under eye, gray on the large scaled area part of caudal fin paler than rest of fin, with ventral to this, the blotches yellowish gray; faint dark dots; spinous portion of dorsal fin caudal fin pale bluish with a black spot with a very narrow pale margin, thin sub- in center and dusky yellow lobes which marginal dark line and a moderately broad are edged in brown, the upper and lower pale band beneath this (unsealed part of edges of fin narrowly light blue; dorsal fin); marginal pattern continues faintly and anal fins yellowish gray with a nar- into soft portion of fin. row pale blue margin and dark sub- In life the holotype was pale bluish with marginal line, the dorsal with a faint 418 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975 band of dusky yellowish beginning at base Paratypes.-USNM 211814, 100.2 mm SL, female, Kana coast of Hawaii north of Kailua, 84 01, of origin of fin and ending halfway to mar- dip net, Stan Ross, December, 1972; BPBM 15493, gin in anterior soft portion of fin; two 92.4 mm SL, female, Oahu, Hawaiian Islands, off diffuse dark spots posteriorly in soft portion Makaha, upper end of a small vertical discontinu- ity in 40 m, dip net, Anthony Nahacky, June 10, of fin in line with this band; a small black 1973; ZUMT 53027, 91.3 mm SL, female, Oahu, spot at rear base of fin; paired fins pale, the Hawaiian Islands, off Keaau Beach Park, Waianae pectoral rays a little dusky, the pelvics with Coast, 23 m, dip net, Eddie Tomihama, October 18, a bluish cast. 1973. In life the females are entirely light bluish; Description.-Dorsal rays XIV,17 (17 or the caudal lobes and the region around and 18); anal rays IV,16 (III,17); pectoral above the eye are slightly more blue; in rays 17; scale rows from upper end of addition to the dark dots posteriorly in the gill opening to caudal base 48 (46 to 47); central portion of the caudal fin, there are diagonal row of large scales above lateral some comparable dots in the lobes of the line to base of first dorsal spine about fin which are not apparent on the preserved 7; diagonal row of large scales below specimens. lateral line to base of first anal spine about 24; circumpeduncular scales about Remarks.-AII but one of the specimens of 24; gill rakers 5 + 12 (4 or 5 + 12); Genicanthus spinus were collected at the vertebrae 10 + 14 (holotype only). island of Pitcairn in SE Oceania (25°S, 0 Depth of body 1.77 (1.8-1.83) in SL; 130 W) in the depth range of 38.5 to 44 m. width of body 3.25 (3.1-3.33) in depth; The exception was the single specimen head length 3.54 (3.58-3.59) in SL; snout speared in 52 m off Raivavae, Austral Islands length 3.21 (2.83-3.02) in head; diameter (24°S, 148°W). One male was observed at of eye 3.89 (3.09-3.89) in head; interorbital nearby Rurutu in 58 m by the author. In width 2.95 (2.83) in head; length of pre- addition, several individuals were sighted at opercular spine 2.83 (2.74-4.02) in head; 55 m outside the reef of Ducie Atoll, Pitcairn least depth of caudal peduncle 2.05 (1.97- Group. Ducie is the easternmost and south- 1.99) in head; caudal fin emarginate, the ernmost atoll in Oceania (24°40'S, 124°47' caudal concavity 14 in SL; length of pectoral W). fin 1.25 in SL; last dorsal spine 1.26 (1.17- G. spin us is closely related to G. semicinc- 1.2) in head; longest (first) dorsal soft ray tus from Lord Howe Island. At the present 4.44 in SL; third anal spine 1.15 (1.1) in time these two species are separated by more head. than 3000 miles. Norfolk Island, the Ker- Serrae on upper preopercular margin 49 madecs, and Orne Seamount (16 fathoms) (31 to 39); small spines on lower margin represent widely spaced intermediate locali- of preopercle 1 (2 or 3); small spines on ties where Genicanthus of this complex might lower anterior margin of interoperc1e 3 (2 be expected. Study of specimens of the to 5); small spines on margin of suboperc1e genus from these localities would be of great o (0-3); spines on preorbital 8 (8 to 13), interest. none enlarged; a shallow notch ventro- Etymology.-G. spinus is named in reference anteriorly in lower margin of preorbital; ex- to the exceptionally large anterior preorbital posed posterior margin of supracleithrum and spme. posttemporal bones serrate. Basal two-thirds of upper lip scaled and Genicanthus personatus, new species median basal portion of lower lip scaled Plate 1 and Table 4 (scaled area set off by 2 pairs of pores); Holotype.-BPBM 15509, 91.0 mm SL, female, dentition typical of the genus, the upper Oahu Hawaiian Islands, off Magic Island, Hono- teeth 4.2 and the lower teeth 4.4 in diameter lulu, ~and and rubble on a sloping bottom in 24.5 01, dip net, David S. Norquist, June 25, 1973. of eye. Longest gill raker 5.9 in eye. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 419

Table 4. Measurements of the type specimens of Genican/hlls persona/lis expressed as thousandths of the standard length ------_-"'-""-'==----=--======--======Hotolype Paralypes BPBM 15509 USNM211814 BPBM 15493 Standard length (mm) 91.0 100.2 92.4 Greatest depth of body 562 560 546 Width of body behind gill opening 173 168 176 Head length 283 279 278 Snout length 88 99 92 Diameter of eye 98 91 96 Interorbital width 96 99 98 length of preopercular spine 100 102 70 Least depth of caudal peduncle 137 142 140 Length of caudal peduncle 126 130 115 Snout to origin of dorsal fin 367 364 364 Snout to origin of pelvic fins 378 396 368 Snout to origin of anal fin 633 647 619 Length of caudal fin 299 Caudal concavity 72 Length of pectoral fin 226 Length of pelvic fin 321 Length of pelvic spine 220 222 221 Length of dorsal fin base 745 728 753 Length of first dorsal spine 87 109 92 Length of second dorsal spine 146 156 162 Length of third dorsal spine 183 197 189 Length of last dorsal spine 220 232 239 Length of longest dorsal soft ray 225 Length of anal fin base 406 387 400 Length of first anal spine 124 129 119 Length of second anal spine 200 204 198 Length of third anal spine 245 254 254 Length of longest anal soft ray 255 ------~-

Lateral line complete, the pored scales (slightly in front of caudal base) straight, 43 (44); auxiliary scales present over entire the posterior with the black curving back- body. Scales on mid-side of body of holo- ward toward upper and lower margins. type with 24 to 27 ridges, ending in ctenii. In life (pI. 1) the body is light gray, each Origin of dorsal fin slightly anterior to scale with a crescentic spot of white; anterior upper end of gill opening; origin of anal head and caudal base black dorsal, anal, and fin below base of eleventh dorsal spine; pec- outer third of the caudal fins whitish with a toral fin just reaching a vertical at anus; bluish cast; pectoral fins whitish; pelvic fins origin of pelvic fins below lower base of yellow orange, whitish basally and on leading pectoral fins; pelvic fins reaching origin of edge. anal fin. Color of holotype in alcohol whitish, the Remarks.-This species is known from only edges of the scales a little darker than centers; four specimens from the Hawaiian Islands hcad in front of a vertical at posterior edge in the depth range of 24.5 to 84 m. All were of eye blackish, the demarcation darker and captured alive by divers who collect colorful more abrupt above eye than below; unsealed reef fishes for the aquarium trade. All were portion of lips whitish; fins whitish except maintained in aquaria before being brought basal two-thirds of caudal fin which is as specimens to the Bishop Museum. The abruptly black, the anterior demarcation 91.3-mm paratype is in such poor condition 420 BULLETIN OF MARINE SCIENCE, VOL. 25, NO.3, 1975 that it was not used for proportional mea- semi/asciatus used herein. I also wish to thank surements. Eroded or damaged fins of the Stan Ross, Anthony Nahacky, Eddie Tomihama, other two paratypes precluded some of the fin and David S. Norquist for collecting specimens of G. personatus, and Clemens Classen and William F. measurements of these specimens. Smith-Vaniz for the collection of G. bellus. The All four specimens are females. In view of latter's specimens are deposited at his institution, the marked sexual dichromatism of all other the Academy of Natural Sciences of Philadelphia. species of the genus, it seems likely that the Phil Lobel and James H. O'Neill kindly provided color photos of G. personatus in an aquarium, personatus male of G. will also have a dif- Homer K. Morgan sent comparable photos of the ferent color pattern. However, no diver has two color forms of G. bellus from Tahiti, and yet reported such a fish by underwater ob- William F. Smith-Vaniz sent transparencies of the servation (several individuals of G. persona- specimens of G. bel/us from Cocos-Keeling. tus have been sighted without being cap- Leighton Taylor and Helen A. Randall took X-rays of all the species of the genus. Pertinent informa- tured) . Of the known species of Geni- tion on literature or specimens was provided by canthus, this species most resembles the Gerald R. Allen, Chuichi Araga, Marie Louise "fucosus" form of G. semifasciatus. If there Bauchot, M. Boeseman, Pierre Fourmanoir, Paul is a different male color phase of G. persona- Gueze, Wolfgang Klausewitz, Margaret M. Smith, tus, it might be expected to have numerous and Alwyne C. Wheeler. poorly defined bars on the upper part of the body. LITERA TURE CITED As mentioned, Genicanthus seems more Araga, C. 1972. Marine angel fishes of the closely related to Holacanthus than any Japanese waters. Animals & Zoos, 9: 299-303. Bleeker, P. 1857. Achtste bijdrage tot de kennis other pomacanthid genus. Of the known del' vischfauna van Amboina. Acta Soc. Sci. species of Genicanthus, G. personatus is the Indo-Neer!., 2: 1-102. nearest in morphology to Holacanthus, thus 1877. Atlas ichthyologique des Ori- it may be the most primitive living member of entales Neerlandaises, ... Frederic Muller, the genus. If this species is truly confined et Cie, Amsterdam, 80 pp., 59 pis. Chan, W. L. 1965. Two new pomacanthinid to the Hawaiian Islands, it may represent allgelfishes from the Macclesfield Bank, South a relict population there. China Sea. Ann. Mag. Nat. His!., Ser. 13, 8: Anthony Nahacky reported that the para- 325-334, 5 figs. type he caught was first observed milling Fowler, H. W. 1934. The fishes of Oceania- Supplement 2. Mem. B. P. Bishop Mus., Chae- well above the bottom with a group of 11(6): 385-466, 4 figs. todon miliaris, a butterflyfish which feeds Fraser-Brunner, A. 1933. A revision of the primarily on zooplankton. The author dived chaetodont fishes of the subfamily Poma- with Mr. Nahacky at the same location but canthinae. Proc. Zoo I. Soc. London: 543-599, saw no more individuals of G. personatus. 29 text-figs., 1 pI. Gi.inther, A. 1860. Catalogue of the acantho- Probably the optimum depth for the species pterygian fishes in the collection of the British is below the deeper diving depths for man Museum. Vol. 2. Taylor and Francis, London, with air-filled SCUBA tanks. 548 pp. Herre, A. W. 1933. Twelve new Philippine Etymology.-Named personatus from the fishes. Copeia, no. 1: 17-25. Latin for Masked. ---. 1953. Check list of Philippine fishes. U. S. Fish Wild!. Serv., Res. Rept. 20, 977 pp. ACKNOWLEDGMENTS --- and H. R. Montalban. 1927. The Philip- Special thanks are due Dr. Yoshiaki Tominaga pine butterfly fishes and their allies. Philip. of the Department of Zoology, University Museum, Jour. Sci., 34 (1): 1-113, 24 pIs. University of Tokyo for sending on loan a valuable Hiyama, Y., and F. Yasuda. 1971. Living fishes collection of Genicanthus for this study. Dr. of the Japanese coastal waters. Kodansha, Tominaga abandoned his plans to revise G I!II i- Tokyo, 337 pp., 527 color iIIustr. canthus when he heard of the author's interest in Jordan, D. S. 1919. The genera of fishes, part the group. II, from Agassiz to Bleeker, 1833-1858. I am grateful to Dr. Fujio Yasuda of the Tokyo Stanford Univ. Pub!., pp. i-ix + 163-284 + University of Fisheries for the photographs of G. i-xiii. RANDALL: REVISION OF THE ANGELFISH GENUS GENICANTHUS 421

Kamohara, T. 1934. Additional notes on the Schmidt, P. J. 1930. Fishes of the Riu-Kiu fishes around Kochi city. Zoo!. Mag., 46 Islands. Trans. Pac. Comm. Acad. Sci. (552): 457-463, 3 figs. In Japanese. USSR, 1: 19-156, 8 text-figs., 6 pIs. 1961. Notes on the type specimens of Smith, 1. L. B. 1955. The fishes of the family fishes in my laboratory. Rept. Usa Mar. BioI. Pomacanthidae in the Western Indian Ocean. Sta., 8(2): 1-9, 7 pIs. Ann. Mag. Nat. Hist., ScI'. 12, 6: 377-384, Laccpede, B. G. 1802. Histoire nature lie des 2 pIs. poissons. Vo!. 4. Plassan, Paris, xliv + 728 Swainson, W. 1839. The natural history of pp., 16 pIs. fishes, amphibians & reptiles. Vo1. 2. Long- Munro, I. S. R. 1967. The fishes of New Guinea. man, Orme, Brown, Green & Longmans, Dept. Agri., Stock Fish., Port Moresby, New Paternoster, Row, and John Taylor, London, Guinea, xxxvii + 651 pp., 78 pIs. 452 pp., 135 figs. Norman, J. R. 1957. A draft synopsis of thc Waite, E. R. 1900. Additions to the fish-fauna orders, families and genera of Recent fishes of Lord Howe Island. Rec. Austral. Mus., and fish-like vertebrates. British Museum 3(7): 193-209, 1 text-fig., 3 pIs. (Natural History), London, 649 pp. Watanabe, M. 1949. Studies on the fishes of the Randall, 1. E. 15'67. Food habits of reef fishes Ryukyu Islands 1. Four unrecorded butterfly of the West Indies. Stud. Trop. Oceanogr. fishes. Bull. Biogeo. Soc. Jap., 16(8): 33-43, (Miami), 5: 665-847. 4 figs. 1973. Tahitian fish names and a pre- Weber, M. and L. F. de Beaufort. 1936. The liminary checklist of the fishes of the Society fishes of the Indo-Australian Archipelago. Islands. Occ. Pap. B. P. Bishop Mus., 24(11): Vol. 7. E. J. Brill, Leiden, xvi + 607 pp., 167-214. 106 figs. --- and W. D. Hartman. 1968. Sponge- Yasuda, F. and Y. Tominaga. 1970. Two new feeding fishes of the West Indies. Mar. Bio!., long-tailed pomacanthine fishes from M iyake- 1(3): 216-225. Jima and Okinawa-Jima, Japan. Japan. Jour. Sauvage, H. E. 1891. Histoire physique, natu- Ichthyo1., 16(4): 141-151,13 figs. relic et politique de Madagascar. Vo!. 16. Histoire naturelle des poissons. Imprimerie ADDRESS: Be/'/lice P. Bishop Museum, P. O. Box Nationale, Paris, 543 pp., 50 pIs. 6037, Honolulu, Hawaii 96818.