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SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM

Butvetin 202

FISHES OF THE MARSHALL AND MARIANAS ISLANDS

BY

LEONARD P. SCHULTZ AND COLLABORATORS:

WILBERT M. CHAPMAN, ERNEST A. LACHNER, AND LOREN P. WOODS

VOLUME 2

Families from Mullidae through Stromateidae

UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1960

For sale by the Superintendent of Documents, U. S. Government Printing Office Washington 25, D.C. = Price $2.25 (paper)

ADVERTISEMENT

The scientific publications of the National Museum include two series known, respectively, as Proceedings and Bulletin.

The Proceedings series, begun in 1878, is intended primarily as a medium for the publication of original papers based on the collections of the National Museum, that set forth newly acquired facts in biology, anthropology, and geology, with descriptions of new forms and re- visions of limited groups. Copies of each paper, in pamphlet form, are distributed as published to libraries and scientific organizations and to specialists and others interested in the different subjects. The dates at which these separate papers are published are recorded in the table of contents of each of the volumes.

The series of Bulletins, the first of which was issued in 1875, contains separate publications comprismg monographs of large zoological groups and other general systematic treatises (occasionally in several volumes), faunal works, reports of expeditions, catalogs of type speci- mens, special collections, and other material of similar nature. The majority of the volumes are octavo in size, but a quarto size has been adopted in a few instances. In the Bulletin series appear volumes under the heading Contributions from the United States National Herbarium, in octavo form, published by the National Museum since 1902, which contain papers relating to the botanical collections of the Museum.

The present work forms No. 202, volume 2, of the Bulletin series.

RemMIncTON KeE.ioea, Director, United States National Museum.

II

CONTENTS

Authorship of sections prepared by collaborators other than Leonard P. Schultz is indicated after the name of the group for which they are responsible. The appearance of my name as second author indicates that I furnished the ecological data and assisted in other ways but did pot write major portions of the manuscript.

Page ETO CUC CLOTS ee eee nee ee eee oe ee eee ees ae eee VIII Phylum Chordata—Cont., vol. 1 Subphylum Craniata—Cont., vol. 1 Superclass Gnathostomata—Cont., vol. 1 Class Osteichthys—Cont., vol. 1 Subclass Teleostom—Cont., vol. 1 Superorder Teleosteica—Cont., vol. 1 Order Percomorphida—Cont., vol. 1 Suborder Percomorphina—Cont., vol. 1 Family Mullidae (by Ernest A. UACHINER) eee oa ee oma 1 Genus Upeneus Cuvier & Valenciennes-_----- 3 Genus Parupeneus Bleeker 2252455252. 6 Genus Maulloidichthys Winhitley a2 2sseseo— 36 Family Pomacentridae (by Loren P. Woops and Lronarp PO SCHULTZ) Seen eae 47 Genus Amphiprion Bloch and Snyder-_-_-_ 47 Genus Dascyllus Cuv- LOT ey es ao ee ces 61 Genus Chromis Cuvier- 66 Genus Abudefduf Fors- Kile tat oT SaaS 79 Genus Pomacentrus acepedes - 5225-5222 104 Family, Labridaee. =.=. = 121 Genus Choerodon Bleekers-2 4 wes 125 Genus Pteragogus Petersaa sewer a eee 126 Pseudocheilinops, new PenUS esse sss 128 Genus Cirrhilabrus Temminck and Schle- Pele mets es 131

IV CONTENTS

Phylum Chordata—Continued Subphylum Craniata—Continued Superclass Gnathostomata—Continued Class Osteichthys—Continued

Subclass Teleostomi—Continued Superorder Teleosteica—Continued Order Percomorphida—Continued Suborder Percomorphina—Continued Family Labridae—Continued

Genus Cymolutes Giin-

Genus Epibulus Cuvier_ Genus Wetmorella Fow- ler and Bean________ Genus Xyrichthys Cuv- ION suse Poe eee Genus IJniistius Gill. - Novaculops, new genus_ Genus Hemipteronotus Lacepéde_._.......- Genus Cheilinus Lacepéde_2- 2 22S Genus Pseudocheilinus

Genus Labrichthys Bleeker a. 22 soaees Genus Labroides Bleeker. 222 eee Genus Cheilio Lacepéde_ Genus Coris Lacepéde_-_ Genus Thalassoma Swainsow. 202524 as Genus Gomphosus

Genus Macropharyngo- don Bleeker________- Genus Stethojulis Gtnther=- 3 -- ae Genus Hemigymnus Giuther 2.2 see Genus Anampses Quoy

and Gaimard_______- Genus Halichoeres Ruppells 22a. bee

Family Scaridies:22.00 22. ae Subfamily Scarinae________

Genus Chlorurus swain-

Genus Scarus Forsk&al__

Page

133 136

138 142 143 143 146 153 163 170 171 176 Waid 183 199 203 206 217 219 223 239 241

242 243

CONTENTS v

Page Phylum Chordata-—-Continued Subphylum Craniata— Continued Superclass Gnathostomata—Continued Class Osteichthys—Continued Subclass Teleostomi—Continued Superorder Teleosteica—Bontinued Order Percomorphida—Continued Suborder Percomorphina—Continued

Family Scaridae—Continued

Subfamily Sparisomatinae-. 250 Genus Calotomus Gil-

berks 2c eee us sone 250 Genus’ Leptoscarus Swainson! ease eee 251 Family Cirrhitidae: 220-222 3222 251 Genus’ Cirrhitoidea eri ksir) Sea 255 Genus Cirrhitus Lace- Pedewsc sao one ees 259 Genus Gymnocirrhites Srart ieee seen 262 Genus Paracirrhites Bleeker 2 w es eee 263 Family Mugiloididae__--------- 266 Genus Parapercis Bleckersans Svar a ats 268 Family Trichonotidae---------- 273 Genus Limnichthys AVR Gees crete any se 278 Genus Chalizodytes Sehuwitz7aeemee eae ae 280 Suborder Blenniinags ss 2.2 wee 281 Family: Glinidag. 25 2 281 Subfamily Tripterygiinae__._ 281 Genus Tripterygion Rissol 222 2 ae eee 283 Genus Helcogramma McCulloch and Waites 2.) vee ee 294 Hamily Blenntidaes =====2 === 301

Subfamily Salariinae (by LronarD P. ScHuuttTz and WiusBert M. CHapMANn)-. 302

Genus Exallias Jordan

and Evermann.-_--_-_ 305 Genus Cirripectes Swain-

SOT yeeros ee ee 307 Genus Rhabdoblennius

Wihitley sue eos 319

VI

CONTENTS

Phylum Chordata—Continued

Subphylum Craniata—Continued

Superclass Gnathostomata—Continued Class Osteichthys—Continued Subclass Teleostomi—Continued

Superorder Teleosteica—Continued Order Percomorphida—Continued

Suborder Blenniina—Continued Family Blenniidae—Continued Subfamily Salariinae—Cont.

Genus Entomacrodus Giles te ees ee Genus Istiblennius Wititleys = 522. ee Genus Salarias Cuvier_ Fallacirripectes, new PEN Gee eee yee eee Genus Alticus Cuvier and Valenciennes________

Praealticus, new genus_- Subfamily Blenniinae__ ____ Genus Petroscirtes Riip-

Genus Aspidontus Quoy and Gaimard________ Genus Runula Jordan and Bollman_______- Genus

Suborder:Ophidinas-< <_< -ese a ee Family Brolutidae_____________ Subfamily Brotulinae______

Genus Brotula Cuvier -

Subfamily Brosmophycinae_ Brosmophyciops, new

Family Carapidaes. 22-5 -- 2222 Genus Jordanicus Gil-

bertis: 2355-782 a See Genus Encheliophis Willers (22252 ese Genus Carapus Rafin- CSQUGy So .= eee eee

Suborder Schindleriina_________________ Family Schindleriidae__________ Genus Schindleria Gil-

Page

327

344 361

362 366 368 373 374 375 379 381 382 382 383 383 384 384

388 390

391 392 393 396 396

396

CONTENTS VII

Page Phylum Chordata—Continued Subphylum Craniata—Continued Superclass Gnathostomata—Continued Class Osteichthys—Continued Subclass Teleostomi—Continued Superorder Teleosteica—Continued Order Percomorphida—Continued suborder Callionymina: o- =... 22-22 397 Family Callionymidae_________-_ 397 Genus Diplogrammus GU a ge a are 399 Genus Callionymus Lin- TU ELOUS cree eles apene et pa 402 Genus Synchiropus Gill, 404 SuborderiScombring: = 25222 62 bee 410 Family Scombridae__________-_- 410 Genus Grammatorcynus Gr Se eA 411 Genus Acanthocybium Guile ae es 412 Genus Katsuwonus Ki- shinouyetes aaa 413 Genus Neothunnus Kishinowuyess: 202 -. 414 Genus Euthynnus Liit- Kemet s Smee ee ee 415 Genus Gymnosarda Gill_ 416 suporder Stromateinas.+ Jee oo ee eee 418 Family Stromateidae___________ 418

Genus Psenes Cuvier and Valenciennes__~__ 418

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INTRODUCTION

By LEONARD P. SCHULTZ

This is the second volume of U. S. National Museum Bulletin 202, “Fishes of the Marshall and Marianas Islands.” The first volume was published on December 15, 1953. The third volume is in preparation.

A list of fish collecting stations, maps, and other information perti- nent to the use of volume 2 were published in volume 1.

A few errors have been noted that we wish to correct:

p. 108, 3d paragraph last line should read 1949, not 1948

p. 166, since Euleptorhamphus is neuter the species name is viride, not viridis

. 259, figure 39, M. dunckeri is figure ‘a’ and M. brevirostris is figure “d” . 279, add p. 576 to Walbaum reference

. 314, in section 5a, 4th line from bottom should read scales cycloid or etenoid

. 869, synonymy, Forsk&l, not Fosk&l

. 386, ocellata, not ocellatus

. 387, last line, soft, not saft dorsal

. 407, 4th paragraph first line, Marston not Marsden

. 537, Monotazxis, not Montazis

‘pl. 26, A, name should be Epinephelus horridus (Cuvier and Valenciennes).

reser ar

FOIE to CSS

Except for the families by Ernest A. Lachner, I have supplied and mounted all the illustrations, and written the legends for them. If any errors were made, my collaborators should not be blamed.

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FISHES OF THE MARSHALL AND MARIANAS ISLANDS Volume 2

Family MULLIDAE: Goatfishes

By Ernest A. LACHNER

Members of the family Mullidae are readily recognized by the two long, unbranched barbels below the chin, their bodies being moderately elongate and often brilliantly colored. Goatfishes are mainly inshore, bottom forms, of small size, occurring in tropical and semitropical marine waters. The meat is sweet and is relished as a delicacy.

This report reviews all forms found in Oceania. The Mullidae of various areas of the Indo-Pacific other than Oceania have been reviewed by several authors, such as the early report by Bleeker (Verh. Akad. Wet. Amsterdam, vol. 15, pp. 1-40, 1875) and the more recent studies by Herre and Montalban (Philippine Journ. Sci., vol. 36, No. 1, pp. 95-136, pls. 1-6, 1928), Weber and de Beaufort (Fishes of the Indo-Australian Archipelago, vol. 6, pp. 361-408, figs. 75-79, 1931) and Fowler (U. S. Nat. Mus. Bull. 100, vol. 12, pp. 258-344, figs. 19-32, 1933). A revision of the genus Upeneus was published by Lachner (Proc. U. S. Nat. Mus. vol. 103, No. 3330, pp. 497-532, pls. 13 and 14, 1954). The many closely related but wide ranging species in the family, particularly those in the genus Parupeneus, and the incomplete nature of the collections from the area under study, necessitated a review of the species occurring in a broader geographical area. The result is a more reliable appraisal of the species.

Many species show differentiation only in coloration, such as the presence or absence of a spot or color mark, or its shape or location. There are few meristic and other morphological characters that aid in the separation of the species. We thus find, as a result of the numerous closely related species and their great range over the Indo-Pacific area, a confusing nomenclature. Many nominal forms have been described from the various insular and subfaunal areas of this vast region. Progress toward a stable nomenclature can be attained only in an appraisal of all forms in the family, including an evaluation of all types in extant.

Hl)

2 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

The primary characters forming the basis for the natural groups in the family are the differentiation of the dentition and the presence or absence of scales on the soft dorsal, anal, and caudal fins. These differences are tentatively summarized in the key to the genera and in the discussion under the generic accounts. The critical characters most useful in distinguishing the species are given in the keys to the species under each genus and in the discussions in the descriptive accounts of the species. The genera Upeneus, Mulloidichthys, and Parupeneus are reported for the Marianas, but only the latter two were taken in the Marshall Islands. The following tabulation shows the number of species in each genus from the three geographical areas:

Marshall Marianas

Islands Islands Oceania Unericuse ae eas. Chee SOAP ES 0 if 4or5 Miillovdichtliysee ee eae 3 1 4 Pariipeneus® .--bss2 2252 x tyes 5 4 15

The range of the data given for the following characters is common to all species reported herein for Oceania: Dorsal rays VIII-i,8 (Upeneus VII or VIII-i,8), anal rays IJ,i,6; pectoral rays 12 to 18; pelvic rays I,5; caudal rays, upper lobe-lower lobe, i,7-+6,1; vertical scale rows 27 to 39; scale rows above lateral line 3, rarely 2; scale rows below lateral line 5 to 7; scale rows around narrowest portion of caudal peduncle 14 to 16 (14 only in Parupeneus); total number of gill rakers on first right arch 19 to 42; scales ctenoid, with 4 to 9 radii; lateral line complete.

In order to determine the extent of infraspecific divergence the wide-ranging species were studied from areas other than Oceania when collections were available. The coloration, proportional, and meristic characters were recorded for each species. Wherever the data may be significant or of interest it is tabulated, segregated by locality, and included under the descriptive account of the species. The specimens studied are listed by geographical areas.

In general, there is very little infraspecific differentiation among the wide-ranging species in the numerous, scattered populations from the Red Sea to the Hawaiian Islands. Some divergence on a low racial level is exhibited by several species which differ mainly in the total number of gill rakers.

KEY TO THE GENERA OF MULLIDAE

la. Dentition complete, teeth on vomer, palatines, and on both jaws. Upeneus Cuvier and Valenciennes 1b. Dentition incomplete, the palatines and vomerine teeth, or the palatines, or those of the upper jaw, absent.

FAMILY MULLIDAE—LACHNER 3

2a. Upper jaw toothless. omer and palatines form a broad palatal patch. Mullus ! Linnaeus 2b. Upper jaw with teeth. 3a. No teeth on palatines or vomer. 4a. Teeth in jaws uniserial, large, stout, blunt-tipped, widely spaced, and arranged more or less in a regular row.__-_-_- Parupeneus Bleeker 4b. Teeth of upper jaw almost always in two rows anteriorly, the teeth of the outer row numbering 1 to 3 on each side, curved and directed posteriorly; teeth in lower jaw almost always in two or more rows; teeth stout, blunt-tipped, and rather widely spaced. Pseudupeneus ? Bleeker 4c. Teeth of jaws small, in a villiform band anteriorly. . Mulloidichthys Whitley 3b. Vomer with a few blunt teeth; none on palatines; jaws with stout, blunt teeth arranged uniserially laterally, and in two or three irregular TOWS ANteTIOL yee ke eee ee eee Upeneichthys ? Bleeker

Genus UPENEUS Cuvier and Valenciennes

Upeneus Cuvier and VaLencriennes, Histoire naturelle des poissons, vol. 3, p. 448, 1829 (type species, Mullus vittatus Forsk&l, designated by Bleeker, Arch. Néerl. Sci. Nat., vol. 11, pt. 2, p. 333, 1876.)

Hypeneus Acassiz, Nomenelatoris zoologici index, p. 190, 1846 (type species, Mullus vittatus Forsk&l, corrected orthography).

Upeneoides BunEexer, Verh. Bataviaasch Gen., p. 64, 1849 (type species, Mullus vittatus Forsk&l, designated by Jordau, Genera of fishes, pt. 2, p. 240, 1919).

This genus is characterized in having the dentition complete, an irregular or triangular patch of small villiform teeth on vomer, an elongate band of such teeth on palatines and in bands of narrow to moderate widths on both jaws. Caudal fin with dark, oblique bars in many species. Body elongate, somewhat compressed, and attain- ing a small size compared with other members of the family.

Several characters were common to all the species listed in the key below but may not be distinctive generically for they may apply to others in the family: Dorsal spines, VIII-i,8 (also VII-i,8 in three extralimital species), the first spine is minute and likely to be overlooked (see Lachner, Proc. U. S. Nat. Mus., vol. 103, p. 499, 1954); scale rows around caudal peduncle 16; scales ctenoid, with 4 to 7 radii; scales on soft dorsal, anal, and caudal fins; first elongate

1 Mullus Linnaeus, Systema naturae, ed. 10, p. 299, 1758(type species, Mullus barbatus Linnaeus, designated

by Bleeker, Arch. Néerl. Sci. Nat. vol. 11, p. 334, 1876).

This genus is limited to the Atlantic ocean.

2 Pseudupeneus Bleeker, Versl. Akad. Wet. Amsterdam, vol. 14, p. 134, 1862 (type species, Upeneus prayensis Cuvier and Valenciennes).

This genus is found in the Atlantic Ocean and in the eastern Pacific waters of the Americas,

3 Upeneichthys Bleeker, Verh. Akad. Wet. Amsterdam, vol. 2, p. 7, 1855 (type species, Upeneus porosus

Cuvier and Valenciennes).

This genus is represented by several species in the warmer waters of Australia and New Zealand and probably extending into Polynesia.

4 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

spine of dorsal fin flexible. Of the 9 species in the genus 4 or possibly 5 are known from Oceania. One species is known from the Hawaiian fauna, U. arge Jordan and Evermann, and one, U. vittatus (Forskal), from the Marianas Islands but not reported as yet from the Marshall Islands.

KEY TO THE SPECIES OF UPENEUS FROM OCEANIA

la. Total number of gillrakers range from 26 to 32; pectoral rays number 15 to 18; peritoneum brown to black. 2a. Caudal fin without dark bars; chin barbels long, 58 to 82 percent of head length in large specimens over 85 mm.; barbel usually extends pos- teriorly beyond vertical drawn through posteriormost point of pre- opercle; plate 75,A______-_- U. sulphureus ‘ Cuvier and Valenciennes 2b. Caudal fin with oblique dark bars on upper lobe; chin barbels short, 46 to 66 percent of head length; barbel when extended posteriorly not reaching vertical drawn through posteriormost point of preopercle. 3a. Lower lobe of caudal fin transparent to dusky, without dark, oblique bars; a pale to yellow median horizontal stripe on side of body, often faintly developed or obscure; plate 75,B. U. moluccensis 5 (Bleeker) 3b. Lower lobe of caudal fin with 2 or 3 dark oblique bars, the outer bar widest and more intensely colored; a light colored median and dorsolateral, horizontal stripe on body usually present. U. vittatus (Forsk&l) 1b. Total number of gillrakers range from 19 to 25; pectoral rays number 12 to 15; peritoneum silvery to transparent, sometimes with scattered, fine brownish spots. 4a. Seales smaller, 36 to 38 vertical rows on body; a faint, tan median and dorsolateral horizontal stripe on body, often completely faded; head and body lacking dark spots; tip of spinous dorsal fin without dark Spots plate: 75 aiys iy ples/seee es os eis U. arge ® Jordan and Evermann 4b. Seales larger, 28 to 32 vertical rows on body; a conspicuous dark brown median stripe on body; head and body with numerous dark brown spots, sometimes faded; outer portion of spinous dorsal fin with a large dark brown to black irregular spot; plate 75,E. U. tragula 7 Richardson

4 Upeneus sulphureus Cuvier and Valenciennes, Histoire naturelle des poissons, p. 450, 1829 (type locality, Antjer Straits of Sundra).

Occurs from East Africa through East Indies, Philippines and certain islands of western Oceania (Fiji, Lachner, Proc. U. S. Nat. Mus., vol. 103, p. 513, 1954); New Hebrides, Herre, Field Mus. Nat. Hist. Zool. Series, vol. 21, p. 210, 1936) but not yet known from the Marshall or Marianas Islands.

5 Upeneoides moluccensis Bleeker, Nat. Tijdschr. Nederl.-Indié, vol. 8, p. 409, 1855 (type locality, Amboina).

Reported from India through East Indies-Philippine area, northward to southern Japan and southward to Australia. Lachner (Proc. U.S. Nat. Mus., vol. 108, p. 515, 1954) doubtfully reports it from the Samoan Islands.

6 Upeneus arge Jordan and Evermann, Bull. U.S. Fish. Comm., vol. 22 (1902), p. 187, 1903 (type locality, Honolulu).

Reported from the Hawaiian, Phoenix, Palmyra, Caroline, and Gilbert Islands.

7 Upeneus tragula Richardson, Rep. Fifteenth Meet. British Asso. Adv. Sci., p. 220, 1846 (type locality, Canton).

This species is reported to be common in the western Indo-Pacific. It occurs from East Africa eastward through the Philippines to the Pelew and Solomon Islands, and from southern Japan to New South Wales, Australia.

FAMILY MULLIDAE—LACHNER 5

UPENEUS VITTATUS (Forskal) PLATE 75,C

Mullus vittatus ForsK&u, Descriptiones animalium .. ., p. 31, 1775 (type locality, Djedda, Red Sea).—Lacrrzpe, Histoire naturelle des poissons, vol. 3, p. 382, pl. 14, fig. 1, 1802——Suaw, General zoology, vol. 4, pt. 2, p. 616, pl. 89, 1803.

Mullus bandi Suaw, General zoology, vol. 4, pt. 2, p. 615, 1803 (type locality, Vizagapatam).

Upeneus vittatus Cuvier and VALENCIENNES, Histoire naturelle des poissons, vol. 3, p. 448, 1829.—Buenxer, Atlas ichthyologique . . ., vol. 9, pl. (2)392, fig. 3, 1877.—Fow er, U. 8. Nat. Mus. Bull. 100, vol. 12, p. 334, fig. 31, 1933; Mem. Bishop Mus., vol. 12, No. 2, suppl. 3, p. 95, 1949.—-SmirTH, Sea fishes of South Africa, p. 228, pl. 27, fig. 561, 1949.

Upeneus bitaeniatus Benner, Proc. Comm. Zool. Soc., vol. 1, p. 59, 1831 (type locality, Mauritius).

Hypeneus vittatus CANToR, Journ. Asiatic Soc. Bengal, vol. 18, p. 1017, 1850.

Upeneoides vittatus GUNTHER, Catalogue of the fishes in the British Museum, vol. 1, p. 397, 1859.—Day, Fishes of India, p. 120, pl. 30, fig. 2, 1878.— SauvaGE, Histoire naturelle des poissons, in Grandidier, Histoire . . . de Madagascar, vol. 16, p. 219 (not pl. 27, fig. 2), 1891.—HerRreE and MonTALBAN, Philippine Journ. Sci., vol. 36, No. 1, p. 105, pl. 4, fig. 1, 1928.

Upeneoides caeruleus Day, Proc. Zool. Soc. London, p. 194, 1868 (type locality, Madras) ; Fishes of India, p. 121, 1878.

Upeneoides vittatus Kuunzinaur, Verh. Zool.-Bot. Ges. Wien, vol. 20, pt. 1, p. 741, 1870 (error).

Upencoides philippinus Fowurr, Proc. Acad. Nat. Sci. Philadelphia, vol. 70, p. 37, fig. 15, 1918 (type locality, Philippines).

SPECIMENS STUDIED

Guam: 2 lots, 7 specimens, 74 to 155 mm. in standard length.

In addition 50 lots totaling 143 specimens were studied from Zanzibar, East Africa, eastward through the Philippines to the Low Archipelago and Marquesas Islands (see Lachner, Proc. U. 8. Nat. Mus., vol. 103, p. 516, 1954).

Description —The following counts and measurements were taken from the seven specimens from Guam. Dorsal rays VIII-i,8, the first spine minute; pectoral rays ii,14 to ii,15; vertical scale rows 34 to 36; scale rows above lateral line 3; scale rows below lateral line 7; total number of gillrakers 27 to 29; length of longest raker in longest filament averages about 1.2.

Peritoneum dark brown to silvery brown; preorbital scales absent; barbels extend beyond eye but not beyond preopercular margin; barbel length in percent of head length 51 to 58; third dorsal spine equal to or slightly greater than second.

Color in alcohol.—Head and body light tan to golden tan, darker above and lighter tan to silvery below; two or three faint, dark horizontal stripes dorsolaterally above middle of body.

Spinous dorsal fin with two blackish horizontal bars, one on outer and one near middle of fin; outer bar passes through outer third of second to fifth spines, and is colored an intense black; a whitish bar

6 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

between the two black bars; soft dorsal with three dusky to black bars, a short one near the posterior basal area, a long horizontal bar at middle of fin and a narrow dusky tip on the first to third rays; caudal fin with oblique, dusky to black bars, 3 or 4 on the upper lobe and 2 or 3 on the lower; tip of longest (outer) rays of the upper lobe are slightly touched in black; outer bar on lower lobe with more intense black and twice as wide as other bars on caudal; this particular bar never at tip of lower lobe; two, nearly horizontal bars extend from area near fork of caudal fin to areas just above and below mid-base of fin; pectorals, pelvics and anal fins transparent.

Range.—From the Red Sea, East Indies and Philippines eastward in the islands of Oceania to the Low Archipelago, and from Japan southward to Australia but not reported from the Hawaiian Islands.

Genus PARUPENEUS BLEEKER

Parupeneus BLEEKER, Versl. Akad. Wet. Amsterdam, vol. 2, p. 344, 1868 (type species, Mullus bifasciatus Lacepéde).

Dentition incomplete; teeth in the jaws uniserial, stout, blunt- tipped and widely spaced, arranged in a more or less regular row; no vomer or palatine teeth; scales around narrow portion of caudal peduncle 14.

The following characters were common to all species of Parupeneus and may apply to some of the other genera in the family: Scales on caudal fin, absent on dorsal and anal fins; dorsal fin rays VITI-i,8, the first spine small; pectoral rays 14 to 18; scales above lateral line 3; scales below lateral line 5 to 7; second dorsal spine (first elongate spine) of spinous dorsal fin either flexible or rigid near tip.

This genus inhabits the warm littoral marine waters of the Indo- Pacific region from the African east coast eastward to Easter Island. It is absent in the Atlantic fauna and the eastern Pacific region of the Americas. Five species were taken in the Marshall Islands and four from the Marianas. There are probably more than 15 species in the area of Oceania.

The following specimens of Parupeneus, representing at least three species almost all taken at night, were not specifically identified, as the coloration typical of the adults was not developed:

Bikini Atoll: 8 stations, 368 specimens, 20 to 59 mm. Eniwetok Atoll: 4 stations, 149 specimens, 21 to 55 mm. Rongelap Atoll: 2 stations, 158 specimens, 33 to 58 mm. Rongerik Atoll: 1 station, 5 specimens, 32 to 35 mm.

The critical characters useful in distinguishing the species of Paru- peneus occurring in Oceania are given in the key. Several of the meristic and proportional characters are summarized and tabulated in tables 59 through 62. Several characters differ in their increase rate of development with increase in body length, notably the length

FAMILY MULLIDAE—LACHNER i of the head, barbel, and snout. A regressive development occurs in the diameter of the eye; it decreases with an increase in body length. These regressions are shown in the frequency distributions in the descriptive accounts of the species P. barberinus, tables 65 and 67; P. bifasciatus, tables 69 and 70; P. trifasciatus, tables 72 and 73; P, pleurostigma, tables 74 and 75; and P. cyclostomus, table 77. T his difference in rate of development has lead to much of the confusion in species recognition. Many of the nominal forms are size variants, the very large specimens of several species look, superfically, totally unlike their respective subadults.

TasBLE 59.—Number of gill rakers in species of Parupeneus from various Indo- Pacific localities

Total number of gill rakers

Species

24 | 25 | 26 | 27 | 28 | 29 | 30} 31 | 32 | 33] 34 | 35 | 36 | 37 | 28 | 39 | 40 | 41 | 42 War beriniesee =) een | eee Aiea | SLA Heed Len | feed Feil LD ia oh ee ere een eS eelee us Pe (ae | ee pleunotaentas 2 |eo ssa ale see pees eee he on) La Ieee ee ee ooo esa [memo mn fea Onentaliguce nee | Sete eee a ee eee ee a hae Apia Bi pie aot S fs fee oe eee AACE OTETIUELA oes | ea ets |e |e |e oan eel | | cs 3s Ae Set | RS = OR{ OS CUTIES a ee | ees |e |e eel eed ee a Eee Ta Sue elOm hehe aaalieeot Saale eae MIVA DBCARUILS eae ee fee | eee | te | ee eae ee 2h bikOe |) v6nlaaes 1 TAS SCICIUS eee ee eee | eee | eee ee eae ee eee 2 3] 9} 23} 19 | 27} 28) 11 4 barberinoides eas |p 225) =| a TUS ASS Hees Sel ese tab | el a pera 3 Sue sce eee DIUT OSG EGS ses |e ee | eel eles (PO) | eLOL | Sal ee eee S ee fae eeee Ui CUS =e Da | ieee Leta Oo) oases eme | eel| e t B By 2 ca ee cyclostomus_._____|----|----|---+ Te Selon ett. | Lea e bE eas Tinteiig ies ee eee 2 Ware ae See Queue ok RAE Se ce we Sakis | Pee DOTDRYTCUS ssa |S eas | oee eee eee ol od 9 a1 lar a Sf eee tes ee See CUT SONETINUG = eee | beens | ee eee ee eee | aoe ae ily: 4on | ie Tm Pee es et eet a fle eee crassilabrig= = 20s tae Sa alee lee ee es ie OE EN ie |) 2 | SR 1 1 oh ee an

Tape 60.—Total number of pectoral fin rays in species of Parupeneus from various Indo-Pacific localities

Pectoral fin rays

Species 14 15 16 17 18

barberints. = 22-2 ee eo ot ee eR 1 15 15 pleurotaenia.. =<. == -- ss eses hae hee cee as [eee eee ee es 1 4) {|Pow ete | SERS coe OVLORLLIS So ee Se oe i (| | el | ee 8 AIG LOT OTC TIVES a a ce re ee | eae ee 3 Bil ee eR CN (Red aT YS pe ees ee el ee eS 4 Ot eee eee al ee eee ee AVILES CLOLU Soe ee Ma ee eS ee a NES |e ae ae 1 13 Die ees PrifCSCtOiUs ene oo cee ARN ee oe enh ee ta 5 Don iene ean sens cee OarDertzO Ides kae en ae i Siar ea! Was cae 2 be Re LPL Bs 13 TUL fae ey ne | eae Re DICUT OSLO ING a mee eee eee ee AU ewe Ge oe ean, 2 14 Dye ec Aes RIDUACUS eo ee ae es eS eS eS ee tk 2 FL7 pees ee | ee ee CUCLOSTO MUG ans tc NS nse Ak, heen ore eth Ae aes eps See ea 26 Oi eee eee ULE CAG se eS BR eNO ee ees Ouse e oe te ees DOrDnyr esi ce on Rae OLE = Se 2 41 sho] ears eee | Cts oa ee CUTUSONE TUS ea ee ee a ee 2 {Ohi | aces en | bebe ee

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FAMILY MULLIDAE—LACHNER

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10 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

Populational divergence on a racial level is evident in three wide- ranging species, P. cyclostomus, P. trifasciatus, and P. bifasciatus. The East Indies and Philippine populations of these species have a lower number of gill rakers than specimens from Oceania (see tables 68, 71, and 78 in descriptive accounts). No other meristic, propor- tional or color character showed any perceptible divergence in the various subfaunal areas of the Indo-Pacific region, but larger and more adequately distributed collections are needed for many of the species.

KEY TO SPECIES OF PARUPENEUS FROM OCEANIA, INCLUDING CLOSELY RELATED EXTRALIMITAL FORMS la. Body with dark or light horizontah stripes; total number of gill rakers range from 26 to 36, fewer than 34 in all but one species.

2a. Peritoneum dark brown to black; second dorsal spine flexible near tip, not

pungent; gill rakers range from 26 to 31; a conspicuous, black hori- zontal stripe passing from snout through eye, on dorsolateral portion of body, and ending on upper portion of caudal peduncle; a large, circular black spot at base of caudal fin, its diameter more than %

that of eye, with the lateral line passing nearly through its center. P. barberinus (Lacepéde)

2b. Peritoneum pale, light to silvery.

3a. Second dorsal spine flexible near tip in the adults, not pungent; gill rakers fewer, range from 29 to 31; barbel of average length, extend- ing to margin of preopercle; a median and a dorsolateral light stripe on body; a light or pale saddle posterior to dorsal fin, followed by a dark brown saddle, which is occasionally divided into two dorsolateral spots; body depth in standard length about 3.1 to 3.3 in-edults2o2 et eee a ee P. pleurotaenia ® (Playfair)

8 Mullus pleurotaenia Playfair, in Playfair and Gtinther, Fishes of Zanzibar, p. 41, pl. 5, fig. 3 (not fig. 4), 1866 (type locality, Zanzibar). Mullus dispilurus Playfair, in Playfair and Gunther, Fishes of Zanzibar, p. 41, pl. 5, fig. 4 (not fig. 3), 1866 type locality, Zanzibar, Island of Pemba) (questionable allocation). Upeneus dispilurus Day, Fishes of India, p. 125, pl. 31, fig. 3, 1878 (questionable allocation). Pseudupeneus fraterculus Fowler, U. S. Nat. Mus. Bull. 100, vol. 12, p. 302, fig. 25, 1933.

Five specimens (USN M 138602 to 138605 and 138651, Albatross) taken at Cataingan Bay and Jolo Market, Philippine Islands, were examined.

The nomenclatorial problems concerning this species are complicated, and the application of Mullus pleurotaenia Playfair, is tentative. Playfair (1866) confused his descriptions of pleuroteenia and dispilurus with his figures, probably by assigning the incorrect figure numbers. Day (1878) examined Playfair’s types and remarked that faint light horizontal stripes were visible on dispilurus as well as pleurotaenia. On the strength of Day’s observation I have placed Mullus dispilurus Playfair in the above synonymy, but as & questionable allocation. The color description that Day gave for his specimens from ‘“‘Sind,” such as the small spots on the scales and the absence of light horizontal stripes, suggests that he had still another species.

Sauvage (Histoire naturelle des poissons, in Grandidicr, Histoire . . . de Madagascar, vol. 16, p. 225, pl. 27, fig. 3, 1891) placed Mudllus pleurotaenia Playfair in the synonymy of Upeneus fraterculus Cuvier and Valenciennes (Histoire naturelle des poissons, vol. 7, p. 524, 1831, type locality, Mahé Island, Seychelles Island, Indian Ocean) and Fowler (U. S. Nat. Mus. Bull. 100, vol. 12, p. 302, fig. 25, 1933) followed this procedure. However Sauvage’s figure 3 shows 2 dark dorsal] saddles on the body which is totally unlike the species in question. His figure closely resembles P. bifasciatus (Lacepéde). Fowler’s figure 25 is fairly characteristic of our specimens. The collections listed above represent those reported on by Fowler from Cataingan Bay and Jolo Market. The inclusion of Upeneus fraterculus Sauvage in the synonymy of P. fraterculus by Fowler is certainly erroneous. Further, there is no validity in applying the name Upeneus fraterculus Cuvier and Valenciennes to our specimens, as no evidence is presented by them that their speci- mens possessed light stripes on the body.

Pseudupeneus ischyrus Snyder (Proc. U. S. Nat. Mus., vol. 32, p. 90, fig. 2, 1907, type locality, Tokyo, Japan) may also represent this species. Snyder’s specimen was undoubtedly considerably faded for he indicates and illustrates remanents of a dark and light dorsolateral body stripe.

FAMILY MULLIDAE—LACHNER tt

3b. Second dorsal spine not flexible, pungent; gill rakers numerous, 36; barbel short, almost extends to posterior margin of eye; 8 or 9 narrow dark horizontal stripes on body; light post-dorsal fin saddle very faint; body deep, depth in standard length about 2.6 in adults; faint traces of 5 dark dorsal blotches on body; plate 78,E__P. orientalis ® (Fowler) 8c. Second dorsal spine flexible near tip, not pungent; gill rakers range from 31 to 33; barbel of average length, extends beyond posterior margin of eye, almost reaching margin of preopercle; a conspicuous black horizontal stripe from snout through eye to dorsolateral portion of caudal peduncle; a distinct round spot dorsolaterally on caudal peduncle, about % diameter of eye, the lateral line just touching its lower margin; a light, post-dorsal fin saddle sometimes visible, followed by a small faint brown saddle; body depth in standard length about 3.4 to 3.8 in adults____P. macronemus ” (Lacepéde) 1b. Body with wide, moderate to intensely developed, dark vertical bands or saddlelike bars, sometimes encircling body; gill rakers numerous, range from 34 to 42. 4a. Three widely spaced blackish transverse bands over dorsal portion of body and extending ventrally to belly but not encircling it; anterior band extends from about the anterior half of the base of the spinous dorsal fin, middle band passes through base of soft dorsal fin, and posterior band passes over caudal peduncle just anterior to procurrent rays of CELUI Chet teers eae hie ee Ree far ee sages a SRE eo ee cio P. bifasciatus (Lacepéde) 4b. Four blackish transverse bands nearly encircling the body; first, and anterior- most, widest, passes over body dorsally between occiput and origin of spinous dorsal fin and extends ventrally through base of pectoral fin; second is separated from third by a scale or two and passes between the spinous and soft dorsal fins; third passes through anterior half of soft

9 Pseudupeneus orientalis Fowler, U. S. Nat. Mus. Bull. 100, vol. 12, p. 294, fig. 23, 1933 (type locality, Cooks Bay, Easter Island). Pseudupeneus multifasciatus Kendall and Goldsborough, Mem. Mus. Comp. Zool., vol. 35, p. 122, 1912 (based on holotype of P. orientalis). Upeneus multifasciatus Fowler, Mem. Bishop Mus., vol. 19, p. 228, 1928 (that portion based on Kendall and Goldsborough, 1912).

This species is known by the holotype, USNM 65639, taken by the Albaiross at Cooks Bay, Easter Island.

10 Mullus macronemus Lacepéde, Histoire naturelle des poissons, vol. 3, pp. 383 and 404, 1802 (type locality

not given).

Mullus auriflamma Lacepéde, Histoire naturelle des poissons, vol. 3, p. 400, 1802.

Upeneus lateristriga Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 3, p. 463, 1829.

Upeneus lateristriata Valenciennes, in Cuvier, Régene animal . . ., ed. 4, page opp. pl. 19, 1836. (spelling error).

Upeneus lateristriga Valenciennes, in Cuvier, Régne animal . . ., ed. 4, pl. 19, fig. 3, 1836.

Parupeneus macronema Bleeker, Verh. Akad. Wet. Amsterdam, p. 24, 1875; Atlas ichthyologique . .., vol. 9, pl. 391, fig. 3, 1878.

Apogon amherstinus Day, Fishes of India, pt. 1, p. 124, 1878 (a manuscript name; = barberinus?).

Eight specimens (USNM 47602, 49329, 147587, and 147538) from the Red Sea were examined.

Although this species has been reported from the Red Sea to the Hawaiian Islands, I cast doubt on its presence east of the East Indies and Philippine Islands. Fowler (Copeia, No. 112, p. 83, 1922) merely listed it from Hawaii. No one has since reported on this material. Poorly preserved specimens of this species may easily be confused with P. barberinus. Vaillant (Bull. Soc. Philomath., Paris, ser. 7, vol. 11, p. 69, 1866) reported macronemus from Tahiti, but his description is also inadequate and he may have had one of Several species.

Day’s (1878, p. 124, pl. 31, fig. 1) description and illustration of macronemus lacks the necessary details to determine with assurance what he had. His fig. 1 resembles barberinus not macronemus. His confusion is illustrated by his statement, ‘‘The similarity between this species {[barberinus| and U. macronemus is so great that I have not considered it necessary to figure both.” Thus, Apogon amherstinus on his page 124, probably a manuscript name of Blyth’s, may either represent macronemus or barberinus.

12 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

dorsal fin; fourth band or saddlelike mark, is present just before base of caudal fin, passing dorsally over peduncle; plate 77,B. P. multifasciatus 1! (Quoy and Gaimard) 4c. Five black, transverse bands over dorsal portion of body and extending ventrally to area of belly but not encircling it; first is located just anterior to spinous dorsal fin; second, wider than first, passes through middle of spinous dorsal fin; third passes between spinous and soft dorsal fins, and is the narrowest; fourth more intensely black than those anterior to it, passes through anterior portion (5 anterior rays) of soft dorsal fin; fifth band passes over caudal peduncle just before base of Gada fink ee eee sehen ay eaters P. trifasciatus (Lacepéde lc. Body with a large (larger than several scales) conspicuous dark or (and) light spot, or patchlike mark, on side; gill rakers range from 24 to 32. 5a. Gill rakers more numerous, total from 27 to 32; peritoneum light, trans- parent or slightly silvery. 6a. Side of body with a very large, somewhat rectangular, patchlike dark mark extending from end of head to a vertical drawn ventrally from origin of soft dorsal fin, where it abruptly ends; a small spot or W- shaped mark about the size of the eye on caudal peduncle just above lateral line and below area posterior of soft dorsal fin; barbel longer, extending beyond posterior margin of preopercle; pectoral rays usually 15 (table 60); plate 77,D_.-_P. barberinoides ” (Bleeker) 6b. Side of body below end of spinous dorsal fin with a black, somehwat cir- cular, spot about 4 scale rows in diameter; this black spot followed posteriorly by an elongate oval light spot below and slightly anterior to base of soft dorsal fin, its horizontal length about equal to length of base of soft dorsal fin; barbel shorter, not extending to posterior margin of preopercle; pectoral rays usually 16 (table 60). P. pleurostigma (Bennett)

U Mudlus multifasciatus Quoy and Gaimard, Voyage autour du monde. . . sur, . . . 1’ Uranie et la Physi” cienne, Zoologie, p. 330, pl. 59, fig. 1, 1824 (type locality, Hawaiian Islands). Parupeneus bifasciatus Weber and de Beaufort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 386, 1931 (in part). Pseudupeneus multifasciatus Jordan and Evermann, Bull. U. 8. Fish Comm., vol. 23, 1903, part 1, p. 256, pl. 22, 1905.—Fowler, U. S. Nat. Mus. Bull. 100, vol. 12, p. 300, 1933. Upeneus velifer Smith and Swain, Proc. U.S. Nat. Mus., vol. 5, p. 130, 1883 (type locality, Johnston Island).

Nine lots, totaling 28 specimens, from the Hawaiian Islands and 2 specimens (USNM 26822, type of Upeneus velifer, and 157360) from Johnston Island were studied.

This species is apparently confined to the Hawaiian and Johnston Islands. The various authors that dis- tinguished trifasciatus and bifasciatus from multifasciatus also report the latter species only from these Islands. Fowler (Acad. Nat. Sci. Philadelphia, Monogr. 2, p. 284, 1938) listed it from Tahiti, Society Islands, but gives no specific data or source of material (one specimen, Parwpeneus trifasciatus, USNM 82966 taken at Tahiti by the Wilkes exploring expedition was misidentified as multifasciatus previous to this report).

P. multifasciatus appears to replace almost completely trifasciatus and bifasciatus in the Johnston and Hawaiian fauna (see discussion of range under trifasciatus and bifasciatus).

It does not seem logical to apply any other name to this species, in view of its restricted distribution, especially when it is compared with such closely related forms, trifasciatus and bifasciatus. However, the figure of multifasciatus by Quoy and Gaimard is crude and their description is incomplete.

12 Upeneus barberinoides Bleeker, Nat. Tijdschr. Nederl.-Indié, vol. 3, p. 263, 1852 (type locality, Wahai, northern Ceram).—Herre and Montalban, Philippine Journ. Sci., vol. 36, No. 1, p. 120, pl. 4, fig. 3, 1928. Parupeneus barberinoides Bleeker, Neder]. Tijdschr. Dierk., vol. 1, p. 234, 1863; Atlas ichthyologique . . .

vol. 9, pl. (2) 392, fig. 5, 1877. Pseudupeneus barberinoides Fowler, U. S. Nat. Mus. Bull. 100, vol. 12, p. 276, fig. 20, 1933.

Thirteen lots were examined: Philippine Islands, USNM 138639 to 138649, 22 specimens, Albatross; Oki- nawa, USNM 71734, 2 specimens, Albatross; Tonga Islands, USN M 65995, 3 specimens, Albatross.

This species has been reported from the East Indies, Philippines, southern Ryukyu Islands and several Island groups of Oceania.

FAMILY MULLIDAE—LACHNER 13

5b. Gill rakers fewer, total from 24 to 27; peritoneum dark brown in adults, silvery brown in small specimens; a light oval spot about 7 scale rows in length below end of soft dorsal fin, the lateral line passing through its lower portion; a distinct circular black spot on caudal peduncle anterior to caudal fin and more of it above lateral line than below, and its diameter greater than orbit; barbel long, extending beyond posterior margin of preoperele; plate 76,A~ 2) 22h ok P. indicus ® (Shaw) 1d. Sides of body chiefly plain, lacking conspicuous stripes, bands, spots or marks; at most, scales may have small light spots or the caudal peduncle may have a light or dark saddle (weak dark saddles through the dorsal

fins present in one species); gill rakers, range from 27 to 39. 7a. Second dorsal spine flexible near tip, not pungent; gill rakers fewer, range from 27 to 32; barbels long, extending beyond posterior margin of

preopercle.

8a. Seales lacking light circular spots; a prominent light spot dorsally on caudal peduncle posterior to soft dorsal fin, its length more than twice diameter of eye (spot present in about 50 percent of speci- mens, faded in others); head longer, 32 to 37 percent of standard length; barbels very long, extending to posterior margin of head; snout longer and pointed, eye smaller, eye in snout 2.2 to 5.1 in specimens larger than 75 mm______---- P. cyclostomus (Lacepéde) 8b. Seales on the body with a light circular spot, slightly smaller than pupil, these forming about 5 faint horizontal rows; light spot on caudal peduncle absent; head shorter, about 30 to 32 percent of standard length; barbels shorter, not extending to end of head; snout shorter

13 Mullus indicus Shaw, General zoology, vol. 4, pt. 2, p. 614 (type locality, Indian Seas) (on Rahtee goo-

livinda Russell, . . . Fishes collected at Vizagapatam on the coast of Coromandel, vol. 2, p. 42, pl. 157, 1803, Vizagapatam). Upeneus indicus Gunther, Catalogue of the fishes in the British Museum .. . , vol. 1, p. 406, 1859.—Day,

Fishes of Malabar, p. 28, 1865; Fishes of India, pt. 1, p. 126, pl. 31, fig. 4, 1875.—Herre and Montalban, Philippine Journ. Sci., vol. 36, No. 1, p. 115, pl. 2, fig. 1, 1928.

Parupeneus indicus Bleeker, Verh. Akad. Wet. Amsterdam, vol. 15, p. 27, 1875; Atlas ichthyologique ... , vol. 9, pl. 394, fig. 5, 1878.— Barnard, Ann. South African Mus., vol. 21, pt. 2, p. 589, 1927.—Weber and de Beaufort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 394, 1931.

Pseudupeneus indicus Fowler, U.S. Nat. Bull. 100, vol. 12, p. 287, fig. 22, 1983.—Smith, Sea fishes of southern Africa, p. 230, pl. 27, fig. 567, 1949.

Upeneus russelli Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 3, p. 465, 1829 (on Rahtee yoolivinda Russell, 1803).

Upeneus waigiensis Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 3, p. 466, 1829.

Upeneus malabaricus Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 3, p. 467, 1829 (type local- ity, Malabar).

Parupeneus malabaricus Weber and de Beaufort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 395, 1931.—Schultz, U. S. Nat. Mus. Bull. 180, p. 130, 1943.

Upeneus griseofrenatus Kner, Sitzb. Akad. Wiss. Wien, vol. 58, pt. 1, p. 305, pl. 3, fig. 7, 1868 (type locality, Fiji).

The following 32 USNM lots totaling 55 specimens were examined: East Indies, 1 specimen; Philippines, 23 lots, 40 specimens; China, 1 specimen; Japan and Okinawa, 8 lots, 4 specimens; New Guinea, 1 specimen; Admiralty Islands, 1 lot, 4 specimens; Fiji Islands, 1 specimen; Samoa, 1 lot, 3 specimens.

This species is widely distributed from the east African coast eastward to islands of Oceania.

An examination of the coloration, size, and location of the light body spot and dark caudal spot, meristic characters (tables 59 and 60), and body proportions (tables 61 and 62), did not indicate that more than one species may be present in our material. Such differences as the location of the eye and length of snout and barbel considered by Weber and de Beaufort (Fishes of the Indo-Australian Archipelago, vol. 6, p. 397, 1931) to distinguish between indicus and malabaricus can be attributed to individual variation or differences in stages of development, a condition exhibited in most of the species of Parwpeneus. Alcoholic specimens 40 to 50 mm. in length have developed the adult coloration, but the eye is larger and the snout considerably shorter than in larger specimens. An inspection of the data in table 63, as well as the data for other species, reveals that as the body increases in length. the eye proportionately decreases in size and the snout increases, thereby considerably altering the position of the eye in respect to head length.

Vol. 2

NATIONAL MUSEUM BULLETIN 202

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FAMILY MULLIDAE—LACHNER LS

TaB Le 64.— Total number of gill rakers in Parupeneus indicus, by locality

Gill rakers Locality 24 25 26 27 Basteindteseee i. LoS 08 seer es eh ee eee he EE | ee See Aes eR ee IPH lip pines ese 2 2 ah A PD OE EDEL EE AO ASA, Fae 1 2 8 6 © nize Se ee oe ee eC ER Dee Daler up et Mee IMM RE TE ul Live Leet eae ae el 1 Tapan-OKIN aw ae oe ere EA ee OS Re we eA Dee dat ce Sapo tse see PACT rea yee eh ee te RP URE ey Bee Oh Dead a np 1 2 1 IRE ener, RUE UE Re Wa MOEA EME WEEE RRP ee SE LINE Me ROR SS Ame Se LE eS Le fel Verea Ue hee Sammons eee eS) Bete Se Eh Ee CENA EO Spore Se ET eee COE de ue ot ee 2

and rounded, eye larger, eye in snout 1.9 to 2.4 in specimens larger than 87 mm.; plate 76,C____P. luteus “ (Cuvier and Valenciennes) 7b. Second dorsal spine not flexible near tip, pungent; gill rakers intermediate in number, ranging from 31 to 35; barbels either short, extending to about the posterior margin of eye, or long, nearly reaching end of head. 9a. Barbels shorter, extending to posterior margin of eye or slightly beyond; pectoral rays almost always 15 (table 60); a light spot or saddle on caudal peduncle just posterior to soft dorsal fin, slightly larger than eye; light spot partially or completely divided into halves along the dorsal midline in some specimens; a dark saddle usually posterior of light spot; three weak bars through dorsal fins in some specimens; a faint dark stripe through eye, faded on body; a small dark spot just behind eye; body coloration generally pale or dusky. P. porphyreus § (Jenkins) 9b. Barbels longer, extending clearly beyond posterior margin of preopercle, nearly reaching end of head; pectoral rays usually 16 (table 60); a light area dorsally on caudal peduncle followed by a faint to moderately developed dark saddle; head and body otherwise uni- formly light golden or pale; plate 76,D. P. chrysonemus # (Jordan and Evermann)

14 Upeneus luteus Cuvier and Valenciennes, Histoire naturelle des polssons, vol. 7, p. 521, 1831 (type locality, Mauritius).—Day, Fishes of India, p. 125, pl. 31, fig. 2, 1878.—Herre and Montalban, Philippine Journ. Sci., vol. 36, No. 1, p. 114, pl. 5, fig. 1, 1928.

Mullus luteus Playfair, in Playfair and Giinther, Fishes of Zanzibar, p. 41, 1866.

Parupeneus luteus Bleeker, Verh. Akad. Wet. Amsterdam, vol. 15, No. 3, p. 32, 1875; Atlas ichthyologi- que... , vol. 9, pl. (4) 394, fig. 1, 1878.—Weber and de Beaufort, Fishes of the Indo-Australian Archi- pelago, vol. 6, p. 401, 1931.

Pseudupeneus luteus Fowler, U. 8. Nat. Mus. Bull. 100, vol. 12, p. 313, 1933.

Five specimens (USNM 138599 to 138601) from the Philippine Islands were examined: This species has been reported from the east African Coast to the Philippines and New Guinea. I have seen no specimens or convincing records of its occurrence in Oceania.

18 Pseudupeneus porphyreus Jenkins, Bull. U. S. Fish Comm., vol. 22, p. 454, fig. 22, 1903, (type locality, Honolulu, holotype, USNM 50705, examined).—Jordan and Evermann, Bull. U. S. Fish Comm., vol, 23, pt. 1, p. 262, fig. 110, 1905.—Fowler, U. S. Nat. Mus. Bull. 100, vol. 12, p. 310, fig. 26, 1933.

Upeneus porphyreus Fowler, Mem. Bishop, Mus., vol. 10, p. 228, pl. 20,A, 1928

Fourteen lots, totaling 72 specimens, from the Hawaiian Islands were examined.

This species may be endemic to the Hawaiian Islands.

16 Pseudupeneus chrysonemus Jordan and Evermann, Bull. U.S. Fish. Comm., vol. 22, p. 186, 1903 (type locality, Hilo; Honolulu; 6 cotypes, USNM 50676, 126548-9, were examined).—Jordan and Evermann, Bull. U.S. Fish. Comm., vol. 23, pt. 1, p. 258, pl. 21, 1905.— Fowler, U.S. Nat. Mus. Bull. 100, vol. 12, p. 312, 1933. Upeneus chrysonemus Fowler, Proc. Acad. Nat. Sci. Philadelphia, p. 648, 1930.

Upeneus taeniatus Fowler, Mem. Bishop Mus., vol. 10, p. 229, 1928 (based on type).

Ten lots (USNM 126548, 126549, 50676), totaling 12 specimens, from the Hawaiian Islands were studied. This species appears to be restricted to the Hawaiian Islands.

16 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

7c. Second dorsal spine not flexible near tip, pungent; gill rakers numerous, about 37 to 39; barbels intermediate in length, extending to posterior margin of preopercle; body pale, margins of fins dusky, soft dorsal and anal with dark spots on basal half, with about 4 faint narrow, dark bars on outer portion__-_--- P. crassilabris (Cuvier and Valenciennes)

PARAPENEUS BARBERINUS (Lacepéde)

PuatE 78,D Mullus barberinus Lacrr®pE, Histoire naturelle des poissons, vol. 3, p. 406, pl. 13, fig. 3, 1802 (type locality, near Moluccas). SPECIMENS STUDIED

Bikini Atoll: 6 stations, 22 specimens, 32 to 175 mm. in standard length.

Eniwetok Atoll: 2 stations, 3 specimens, 34 to 169 mm.

Rongelap Atoll: 3 stations, 5 specimens, 66 to 109 mm.

Rongerik Atoll: 1 specimen, 162 mm.

Kwajalein Atoll: 1 specimen, 132 mm.

Guam: 3 lots, 24 specimens, 37 to 146 mm.

In addition 54 lots totaling 153 specimens were examined from: East Indies and Philippines, 41 lots, 124 specimens; Okinawa, 2 lots, 2 specimens; Caroline Islands, 2 lots, 12 specimens; New Guinea, 1 specimen; Solomon Islands, 2 lots, 3 specimens; Gilbert Islands, 1 specimen; Samoan Islands, 1 specimen; Admiralty Islands, 1 lot, 7 specimens; Tuamotu Islands, 1 specimen; Pacific Ocean, 2 lots, 2 specimens (Wilkes exploring expedition).

Description.—The following counts and measurements were taken from 8 of the specimens from Bikini Atoll, 65 to 144 mm. in standard length (other counts and measurements in tables 65-67): Dorsal rays VIII-i,8; anal rays 11,6; pectoral rays ii,14 to 16 (30 specimens); vertical scale rows 28 to 29; scale rows above lateral line 3, below lateral line 6.

Body depth 3.5; head length 2.8 to 3.0; length of caudal peduncle 3.6 to 4.1; length of longest pectoral ray 4.1 to 4.7; all in standard length. Eye 4.6 to 5.5; length of upper jaw 3.0 to 3.4; depth of caudal peduncle 2.5 to 3.2; length of snout 1.8 to 2.2; length of barbel 1.4; all in length of head. Eye in snout 2.4 to 3.3 Gill rakers, including rudiments, 6 or 7+-1+-20 or 21, total (89 specimens) 27 to 29, longest raker about two-thirds longest filament, 2 to 2.3 in eye.

Teeth of upper and lower jaws stout, blunt tipped, and in a single row; no teeth on vomer or palatines; small scales on caudal fin, absent on dorsal and anal fins; cheek and opercle scaled; barbels reach vertical drawn touching posterior edge of preopercle; first dorsal spine minute, second spine about nine-tenths length of third spine; tip of second spine of spinous dorsal flexible in adults; peritoneum dark brown to black; scales ctenoid, with about 4 to 8 radii on the anterior field.

Color in alcohol.—Head and body light or whitish on cheeks, oper- cles, and area on body below horizontal stripe, but the scales in some

FAMILY MULLIDAE—LACHNER Vz

specimens are bordered in dusky; dorsal portion of snout, head, and body above horizontal stripe light tan to dusky tan; a blackish hori- zontal stripe faintly developed from tip of snout, on each side, to eye, well developed from eye through anterior portion of lateral line and extending to upper portion of caudal peduncle, where it ends just beyond soft dorsal fin; width of horizontal stripe about two-thirds diameter of eye; a large, circular, black caudal spot, located at mid- base of caudal fin; the posterior margin of spot reaching end of hypural; lateral line passing through middle of spot; some dusky pigment on midportion of membrane of soft dorsal fin, forming a faint horizontal streak; some dusky pigment on membrane of spiny dorsal fin; remainder of fins transparent.

Color in life —The following color notes were taken from a Koda- chrome transparency of a specimen about 12 inches in total length, loaned to the writer by Dr. Donald Strasburg, U.S. Fish and Wildlife Service, Honolulu, Hawaii. Head and body light silvery; lips and lower cheeks orange; black stripe from snout through eye dorso- laterally to area below middle of soft dorsal; area above black dorso- lateral stripe yellowish; a black basicaudal spot; spinous dorsal fin bluish-orange near base, bluish outer portion; soft dorsal fin light bluish; base of pectoral fin rose; pectoral fin transparent; pelvic fins light orange; anal fin light dusky to blue; caudal fin ight blue; barbels pale blue; iris orange to light silvery.

Range.—This species is widely distributed. Our material extends from the East Indies to the Low Archipelago, but it has been reported frequently in the literature from the Red Sea eastward to the Hawaiian Islands.

Remarks.—No measurable differentiation was found in populations from several major island groups of the Indo-Pacific region.

Interesting and important phenomena in the morphology of this species are the change and rate of change in length of the head, barbel, eye, and snout, with increase in body length. Although specimens about 30 mm. in length have a coloration typical of the adults, the head, barbel, and snout are considerably shorter. Small specimens, 30 to 50 mm., show a rapid increase in the length of head, barbel, and snout in respect to body length, whereas in specimens greater than 50 mm. this differential rate of development is reduced almost completely except in the length of snout, which increases steadily with increase in body length. The diameter of the eye shows an interesting regres- sion when related with length of body for it decreases uniformly with increase in body length.

These data demonstrate how the large specimens may look com- pletely ornate; with their small eye and extremely long snout; espe- cially so in specimens having the coloration faded. Specimens such

18 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

as these may represent several of the nominal species of Parupeneus considered in the doubtful status.

P. barberinus superficially resembles macronemus but the dark spot on the peduncle is much larger and located at the midbase of the caudal fin rather than dorsolaterally on the caudal peduncle. The characters given by Smith (Sea fishes of southern Africa, p. 229, 1949) to distinguish between these species, such as the elongation of the last rays of the soft dorsal and anal fins, is completely erroneous for this condition represents an age character in both species. His illu- stration of macronemus (fig. 565), lacks detail and accuracy in the size and location of the caudal spot.

TABLE 65.—Length of head and barbel as a percentage of standard length in Paru- peneus barberinus from various Indo-Pacific localities, by size group

Standard Head length Barbe] length length in Se es a pe arn oe ae || ae J mm, 31 32 33 34 35 36 37 38 39: |,18 | 19 | 20 | 21 | 22 | 23 | 24.) 26 || 26 30- 49_..___ 2 2 obese i a | es al a 2 1 1 2 DR ah ee ee SOR; 69-2 22s FSi Sea ee | es 1 3 5 Dyes. Ele ee (BSE eee 1 1 2 4 2 1 OU OO eee |e eee per ee ee | eee ee 2 A ed bere be) ee fe eee | Mo een ek = sas 90-109). - == 2 |e = 1 2 Bees ee ee tes | eee | ee | ek | 1 3 1 1 Resa es PIQSIZOF ee steel ee ] Ai Se € 3) Bie ok es BS coe SRI. oe EE | See Cae Se Sse sn Ee ae TAQ S49 a See se ee eee 1 4 4 re é aaaee canoe ae 1 2 3 2 ee 1 - 5 O10 Pee ele | ee ee ots | ee | ee | eee | ee Co 1 is Ie sao W7OFI89 + BS sees Sah. SAN eee 2 |S Be Sess | Soo See ee eee BS = Djs Se 190-209-2225 2 eo he eS eae Sa ae 2 ee ae ee a eee ee ee Be Sees 220 ee A | Eee cape eee | eee | nl | mene ae ee |e | eee eee ees ee ee eee eee |p P30 24g Eee pL eS | SERS eee eee Lien eae ee 1p] E sae Ee | Pe er ae ee Lassa e224 1 |

TaBLE 66.—Numober of gill rakers in Parupeneus barberinus, by locality

Upper limb Lower limb Total ! Locality Pcs eee ee ee | eee 6 7 8 19) |) SO 21 3) 226 23) 26 2c |) 28) 298 sO) | a East Indies,

Philippinesi< 222232 11/}) 25 1 2 32) EON) WS 5 1 3 7 6 | 10 6 Okinawa\sland= 522-22 = 2|5. 222 “Pg = sift a tl a a a 1 eee 2 aes Lig sat ee es eee Marshall Islands_-_._-_____ 14 deliss eases 8 6] 10 Di | eae | ee 4 10 45) | ort A | ee Marianas Islands__.._-__- 20 LP pA Soe eas 8 5) Ag oy |e eee 5 11 Boe a RARE Solomon Islands_________- Tipe Seen fos eal ie at | AE cel le Peete RS oe eee alae 8 Vighie 4 LCs 2 GilbertiIslands22- = 2.2.22 Lon| Scced | CR Se ates G1) | ee Pe ce |S | a Jul ee ges > eel eee Samoan Islands____________ Dv eee) [three UR eee Memes Ae pete lh pe Paws) 2 sees | 0 al ee Tuamotu Archipelago_____ Dy |e Sa |e = ol Dy staal Oo ee rele J le eet |S ae

1 Raker at the angle of the arch was included only in the total count.

FAMILY MULLIDAE—-LACHNER 19

TABLE 67.—Diameter of eye and length of snout as a percentage of standard length in Parupeneus barberinus from various Indo-Pacific localities, by size groups

Standard Eye Snout length inmm.,

PARUPENEUS BIFASCIATUS (Lacepéde)

PLATE) 77,A

Mullus bifasciatus LacErrD®, Histoire naturelle des poissons, vol. 3, pp. 383 and 404, pl. 14, fig., 1802 (no locality).

Parupeneus andrewsii Reaan, Proc. Zool. Soc. London, p. 403, pl. 65, 1909 (type locality, Christmas Island, Indian Ocean).

Parupeneus bifasciatus WEBER and DE Bravurort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 386, 1931 (in part).—Scnuurz, U. S. Nat. Mus. Bull. 180, p. 130, 1943.

Pseudupeneus bifasciatus Fowuier, U. 8. Nat. Mus. Bull. 100, vol. 12, No. 1513, p. 291, 1933; Mem. Bishop Museum, vol. 12, No. 2, p. 95, 1949.

Upeneus bifasciatus Herre and MonrauBan, Philippine Journ. Sci., vol. 36, No. 1, p. 118, pl. 6, fig. 2, 1928.

Upeneus semifasciatus Macunay, Proc. Linnean Soc. New South Wales, vol. 8, 1883, p. 263, 1884 (type locality, Hood Bay, New Guinea).

SPECIMENS STUDIED Guam: 1 specimen, 59 mm. in standard length. Rota Island: 1 specimen, 142 mm. In addition 43 lots totaling 91 specimens were examined from: East Indies

and Philippines, 33 lots, 57 specimens; Phoenix and Samoan Islands, 8 lots, 32

specimens; Society Islands, 2 lots, 2 specimens.

Description.—The following counts and measurements were taken from the two specimens from the Marianas Islands (those taken of certain critical characters from specimens other than the Marianas Islands are listed in tables 68-70): Dorsal rays VIII—i, 8; anal rays I, i, 6; pectoral rays ii, 14; vertical scale rows 28; scale rows above lateral line 3, below lateral line 7, around caudal peduncle 14.

Body depth 2.9 and 3.3, head length 3.1, length of caudal peduncle 3.6, 3.7; length of longest pectoral ray 4.2 and 5.0; all in standard

20 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

length. Eye, 4.2 and 4.9, length of upper jaw 2.6 to 3.0, depth of caudal peduncle 2.7 to 2.8, length of snout 1.8 and 2.3, interorbital width 3.5 and 4.0, length of barbel 1.7 and 1.9; all in length of head. Eye in snout 1.7 and 2.6. Interorbital width in eye 0.7 to 1.0. Gull- rakers, 8+1-+28, total 37; longest raker about 1.2 to 1.3 in longest filament.

Teeth of the upper and lower jaws uniserial, blunt, and spaced a moderate distance apart; no teeth on vomer and palatines; scales on caudal fin, absent on dorsal and anal fins; barbels extend to about the posterior margin of eye, sometimes reaching just before posterior margin and sometimes extending beyond this margin; second dorsal spine about 1.3 in length of third dorsal spine, second dorsal spine pungent, the tip not flexible; peritoneum transparent, lateral line complete; scales ctenoid, with about 5 to 7 radii in the anterior field ; last rays of soft dorsal and anal fins slightly elongated.

Color in alcohol._—Head and body light tan to dusky; 3 dark saddles or traverse bands on body: The anterior saddle extends from about the anterior half of the base of the spinous dorsal fin to the ventro- lateral portion of the belly; the middle saddle, a bar, extends from the base of the soft dorsal fin almost to the base of the anal fin; the posterior saddle extends from the dorsal portion of the caudal pedun- cle, just before the procurrent rays of the caudal fin, to just below the lateral line; these saddles, especially the anterior one, may be greatly faded in some specimens. The spiny dorsal fin is transparent, with some dusky near the tip; the soft dorsal is marked with 5 to 6 alter- nating transparent and dusky horizontal stripes; the caudal is dusky to transparent; the pectoral is transparent; the pelvic fin is trans- parent to dusky, with the outer margin of the fin white; the anal fin is light to dusky, with more dusky located near the tips of the rays, and traces of possibly 3 dusky stripes.

Range.—This species has been reported from Madagascar (Sauvage, Histoire naturelle des poissons, 7n Grandidier, Histoire . . . de Mada- gascar, vol. 16, p. 221, 1891) eastward through the Indian Ocean, East Indies, Philippines and island groups of Oceania east to the Society Islands and northeast to the Hawaiian Islands (Fowler, Mem. Bishop Mus., vol. 10, p. 227, 1928). It has not been reported from Japan. Since bifasciatus has been considerably confused with irifasciatus and multifasciatus by various authors, the specimens in- volved in the reports of Sauvage and Fowler should be reexamined. These localities represent certain east and west extremes in the distribution, and the material may involve different species.

Remarks.—The snout and barbel elongate considerably with in- crease in body length and the eye becomes proportionately smaller.

FAMILY MULLIDAE—LACHNER Dit

There may be a tendency for specimens from Oceania to have a greater number of gillrakers than those from the East Indies and Philippines.

TaBsLeE 68.—Number of gill rakers in Parupeneus bifasciatus, by locality

Upper limb Lower limb Total ! cality

718 | 9 | 26 | 27 | 28 | 29 | 30] 31 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 East Indies-Philippines_---| 9 | 28 1 510) 6 6 AMS ied Sa o0F | 12 5 2 lee ee ee Manianasislands=s==-4--—-|".-= 20) eee Ewe 2/222 S Qe] see eee oe Aa ae Pil eel bs Ate ele Phoenixaisiandasssoe-— sean = 3 hg eater ale gs Zale 2a ee oe eee eee ee aye 3 SS | es Samoantislands: =—2= 2445 Sel nee | ee ae rata BA ie | ae By lse2idl aes 1 sa ee Bacletylslandsses===2—== as |=-=—|—==— Di | nate Alec etre [a = 1 Mere Steels we Pea | eee 1 1

1 Raker at the angle of the arch was included only in the total count.

TABLE 69.—Diameter of eye and length of snout as a percentage of standard length in Parupeneus bifasciatus from various Indo-Pacific localities, by size-groups

Standard Eye Snout length |_ par bie: = in mm,

TaBLe 70.—Length of head and barbel as a percentage of standard length in Paru- peneus bifasciatus, from various Indo-Pacific localities, by size groups

Standard Head length | Barbel length

length me == in mm.

605792 = 1 1 Seesaw 3 Vy peo 1 2 4 1 Li Rese | ees ae ee 80-092 set ien see 1 3 5 To leeooe J ease 6 1 2p eee see tae wee BUH 100 ©) sete | renee Oren | seen sete | cas | nce | cue | fo | ete eee | trae ear | sneer A201 SQees = S| Benen | eeT ee 1 BD Sy ek | a ee SES eee 1 Le eee ee eee 140-159 REE Ee oa 1 2 Te ae a ee 1 2 SD hee ee ee LOOSE Obeen en |e 1 3) |aseo= Dees eae acces |ereeaean ae 1M eee 2 2 eae TSO—L9922 = 2 | SESE SESE 2 3 AS See eee eee ee ee ee 1 Gy zee 1 ANY 21 See nee eee eee 1 2 eee | ee eee | Pe es | ee eee | 2 Dao se ASO eee | eee ree | ere | ee Sf | es | ee | ee ee ee eee eee ee ee On| sean A025 OES ames ee eee ete | en 1 1 | aaa | ae | ee | | ee ee ee 2

29 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

PARUPENEUS TRIFASCIATUS (Lacepéde) PuatE 77,C

Mullus trifasciatus LacEPEpDE, Histoire naturelle des poissons, vol. 3, pp. 383 and 404, pl. 15, fig., 1802 (no locality).

Parupeneus trifasciatus WEBER and DE BEaAvurort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 382, 1931.—Scuuutz, U. 8. Nat. Mus. Bull. 180, p. 130, 1943.

Pseudupeneus moana JorRDAN and SxEAuxg, Bull. U. 8. Bur. Fish., vol. 25 (1905), p. 274, 1906 (type locality, Apia, Samoa).—Snyper, Proc. U. 8. Nat. Mus., vol. 32, p. 89, fig. 1, 1907.

Pseudupeneus trifasciaius FowuER, U. 8. Nat. Mus. Bull. 100, vol. 12, No. 1513, p. 295, fig. 24, 1933. Mem. Bishop Museum, vol. 12, No. 2, p. 95, 1949.

Upeneus atrocingulatus Kner, in Steindachner and Kner, Sitzb. Akad. Wiss. Wien, vol. 61, p. 443, 1870 (type locality, Savay).

Upeneus moana HrrrE and Monta.san, Philippine Journ. Sci., vol. 36, No. 1, p. 124, pl. 4, fig. 2, 1928.

Upeneus trifasciatus GUNTHER, Journ. Museum Godeffroy, vol. 2 (Fische der Siidsee, pt. 3), p. 59, pl. 44, fig. B (not C), 1873.

SPECIMENS STUDIED

Bikini Atoll: 24 stations, 72 specimens, 43 to 184 mm. in standard length. Eniwetok Atoll: 1 station, 2 specimens, 92 and 106 mm.

Kwajalein Atoll: 1 specimen, 112 mm.

Likiep Atoll: 2 lots, Univ. Washington, 2 specimens, 81 and 90 mm.

Rongelap Atoll: 6 stations, 30 specimens, 46 to 150 mm.

Rongerik Atoll: 5 stations, 31 specimens, 45 to 146 mm.

Rota Island: 1 lot, 2 specimens, 164 and 193 mm.

Guam: 2 lots, 3 specimens, 49 to 53 mm.

Saipan: 1 specimen, 52 mm.

In addition, 49 lots totaling 110 specimens were examined from: East Indies and Philippines, 37 lots, 54 specimens; Japan and Ryukyu Islands, 2 lots, 4 specimens; Formosa, 1 specimen; China, 1 specimen; Caroline Islands, 1 specimen; Phoenix and Samoan Islands, 4 lots, 46 specimens; Society Islands, 3 lots, 3 specimens.

Description.—The following counts and measurements were taken unless indicated otherwise, from 5 specimens from the Marshall Islands, ranging in length from 47 to 162 mm. (Counts and measure- ments of specimens from other Indo-Pacific localities are also included in tables 71-73): Dorsal rays VIII-i,8; anal rays 1,16; pectoral rays (28 specimens), ii,13 and 11,14; vertical scale rows, 22 specimens, 27 to 29; scale rows above lateral line, 3, below lateral line 6, around caudal peduncle (16 specimens), 13 and 14.

Body depth 3.5 to 4.3; head length 2.9 to 3.3; length of caudal peduncle 3.8 to 4.1; length of longest pectoral ray 3.9 to 4.6; all in standard length. Eye, comparatively larger in smaller specimens, 3.6 to 5.8; length of upper jaw 2.4 to 3.0; depth of caudal peduncle 2.9 to 3.3; length of snout 1.8 to 2.8; interorbital width 3.8 to 4.7; length of barbel 1.3 to 1.6; all in length of head. Eye in snout 1.3

FAMILY MULLIDAE—LACHNER 23

to 3.3. Interorbital width in eye 0.7 to 1.1. Gillrakers, 8 to 10+1-+28 to 31, total 37 to 42 in 25 specimens from the Marshall Islands; longest raker about 1.2 to 1.3 times longer than longest filament.

Teeth of the upper and lower jaws uniserial, blunt, stout, and spaced a moderate distance apart; those in the upper jaw number about 20 to 24, in the lower jaw, 18 to 23; no teeth on vomer and palatines; scales on caudal fin, absent on dorsal and anal fins; barbels extend beyond eye, reaching posteriormost (upper) portion of margin of preopercle; second dorsal spine about 1.4 in length of third dorsal spine, pungent, the tip not flexible; peritoneum transparent; lateral line complete; scales ctenoid, with 5 to 7 radii in anterior field; last ray of dorsal fin greatly elongate, about 1.5 to 2.0 times longer than other rays of fin, last ray of anal fin about 1.3 times longer than other rays of fin.

Color in alcohol.—Head and body light tan to dusky; barbels tan; a diffuse dark brown stripe from snout to eye, often completely faded, ends just posterior to eye in the form of a dark brown spot or blotch; the characteristic color pattern of this species is the presence of 4 to 5 dark brown to black dorsal saddles on the body, each extending from the middorsal area below the lateral line to the ventrolateral portion of the body; the two anterior saddles are often greatly faded and almost obscure; the first saddle (anteriormost), is located just before the spinous dorsal fin and is about 1 to 1.5 times wider than eye; the second saddle is wide, about 1.5 to 2.0 times diameter of eye, and passes through middle of spinous dorsal fin; the third saddle is narrow, slightly less than diameter of eye, and passes between spinous and soft dorsal fins; the fourth saddle, darker and more conspicuous than those anterior to it, passes through the anterior portion (first 5 rays) of the soft dorsal fin and is slightly wider than diameter of eye; the fifth saddle (posteriormost) passes over caudal peduncle just before base of caudal fin, its width is about 1.3 times greater than eye and it, too, is dark and conspicuous; a white saddle, located between the fourth and fifth dark saddles, is present on caudal peduncle; this white mark is variable in size, extending usually to the lateral line, but in some specimens extends below the lateral line to ventral portion of peduncle; pectoral fin transparent; spinous dorsal with margin of first spine dusky, remainder transparent; soft dorsal with a black bar just above and parallel to base, but extending to tip of membrane of last ray, remainder of fin transparent; margin of outer ray of pelvic fin slightly dusky, remainder of fin transparent ; anal fin with 3 narrow light tan stripes on outer half and parallel to base, remainder of fin transparent; outer margins of caudal fin edged with black, remainder of fin slightly dusky to transparent.

422142—60—_3

24 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

Although great variation exists in the intensity of color on the body, the pattern is uniform.

Color in life-—From kodachrome transparency, snout brown to pupil, with 3 or 4 blue lines from eye to tip; one blue line passes through iris of upper part of eye and ends just posterior to eye; opercle with a brownish purple patch; bar from snout to eye very diffuse and dusky; spot behind eye black and conspicuous; base of pectoral fin purple brown; saddles gray to black; scales with diffuse light blue margined in yellow, the bluish becoming circular spots towards base of caudal fin; pectoral fin light yellow; spinous dorsal light orange near base, becoming light yellow towards tips; basal half of spinous dorsal with black rays; the membrane dusky with dashes of yellow; upper half of fin with 4 narrow yellow stripes parallel to base, remainder of fin transparent; pelvic fin with 3 or 4 alternating blue and yellow stripes, the base of fin orange and its outer edge tipped in black; anal fin with 3 yellow wavy stripes on outer hali more or less parallel to base, the basal half with yellow stripes broken into irregular spots; basal portion of anal fin with purple background, outer part transparent; rays of caudal yellow, yellow to purple near base; outer margin of lobes black, the membrane of fin transparent.

Range.—This species has been reported from the Indian Ocean and ranges eastward through the East Indies, Philippines, and Islands of Oceania to the Society and Tuamotu groups. It is known from southern Japan and southeastern China. It has not been reported in the Hawaiian Islands. Fowler (Acad. Nat. Sci. Phila- delphia Monogr. 2, p. 284, 1938) reported it from Johnston Island, but his specimen should be reexamined, for it possibly may represent P. multifasciatus.

Remarks.—In applying the name trifasciatus to this species, I have followed the reasoning of Weber and de Beaufort (1931, pp. 384-385) wherein trifasciatus (Lacepéde, 1802, p. 404, pl. 15) is distinguished from bifasciatus (Lacepéde, 1802, p. 404, pl. 14). However, the crude manner in which Lacepéde’s figures of these two species were com- pleted and his inadequate description certainly invites suspicion concerning this distinction. Jordan and Seale (1906, p. 274) were so positive Lacepédes species were identical that they described their Samoan specimens as a new species Pseudupencus moana, which is here placed in the synonymy of trifasciatus.

No evidence was found from examination of specimens over a wide area of the Indo-Pacific to support the naming of subspecies of trifasciatus on the basis of color pattern, as Weber and de Beaufort have done. A significantly lower number of gill rakers was found in specimens from the East Indies and Philippine Islands (average about 37) than in those from the Islands of Oceania (average about

FAMILY MULLIDAE—-LACHNER 25

39). These data only suggest racial differentiation, for the overlap in the frequency distributions of the gill raker counts from these two general areas (see table 71) is too great to recognize them as subspecies.

TABLE 71.— Number of gill rakers in Parupeneus trifasciatus, by locality

Upper limb Lower limb Total !

Locality

7 | 8 | 9 | 10} 27 | 28 | 29 | 30 | 31-] 32 | 35} 36.) 37 | 38 | 39} 40 | 41 | 42 East Indies-Philip-

PINES: — S85 re ee Se 6 | 31 Bee Le LOR ELO Deel eeel oS 9 | 18 8 3 1 | ese ees=

Japan-Ryukyu NSInNOSae eee. ee | ase Oy ecko Oh eee NE I tage ie al eee ae aE 1 1 Lasse Formosa-Hong Kong-|_---] 2 |_---|_---]---- TAA ERS SE oil | Sees | alee te eS Lei ete fF Ee oe eee ee BoninWslandsee2- 22222) 2-2 1 ee eB ee | ee PR ea oe EE ot Ay pees eee @arolinedisiands).--=_|2---|--2— Ms] ee eee |e Dp} | Uf Pe EN A |S SR le dO |e Seo a Marshall Islands_.__-_|__-- Tay LEU lie tS es | aN ek OO athe Dee ose |e ee De ASE OVE | Bele. 1 Marianas Islands--_-__|__-_|-_-- Dre ie |e ee PA ee eal ed | (emer poser Ue tle a 2 | ese es Samoan Islands.-_____|--_-- NST R24 a) | ees | ee Dy) DD i5 |Z kn awe eee eee | een eee TAASE PLE eo 2 Society Islands------.}----|---- AN Ee See | Pie ee Ses a eee ee ee | a eee ei) 2 1 Narcuesaspls am cds saa= |i glee | ee eens een ee CAV |e | ace en ef Lee [re | eee S| ee ees

1 Raker at the angle of the arch was included only in the total count.

TaBLe 72.—Diameter of eye and length of snout as a percentage of standard length in Parupeneus trifasciatus from various Indo-Pacific localities, by size growps

Standard Eye Snout length in ews mim, 5 6 7 8 9 10 | eA Dy tS plan |e eek eel fall Sine |e 20 lara

AQ FORE a Se | eae 2 A: oe ee 3 | | eae | Wee weer | rene) Aol ares yce| JONOS A AS el Oa Pa 60=) 79. 4 hee se ea 6 Orlin Ss ieee | es Eee 5 ON eee Te eg a aie SO POO ees i] Sh els mea 3 Di eee se Sete |e alleen ee 2 2 1 ee tee ne eee SCS OST eee cla | Pi FBS a aD la Bl fa ge | ol ph a pepe 8 P20=139 2302 | kee 2 ‘Weis | Cee ees ee ae le es Se a AYRES Sek 2 1 1 Ly [see eee 140-159____}_---_-_ 2 oy Re ee Map ee BR Se a el 2 Pee Se ee 1GO=1 7 Oo) ees 8 OS es ea se le | |W | 5 | Reka een 180=109..25 -|L2e FS) | Saag VS Mee Re (We OU te 1 (a 2 1 1 1 200-219____ 1 DV | pesme Ne Ta Nee | a eA ae NID ge ce | Ae es ee | Re Sblsaoee seco

TABLE 73.—Length of head and barbel as a percentage of standard length in Paru- peneus trifasciatus from various Indo-Pacific localities, by size groups

Standard Head length Barbel length length pea sche Jb 5 in mm,

30 | 31 | 32 | 33 | 34] 35 | 36 | 37 | 38 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 29 | 30

40-50 ee na A ELA tee) ees ees SLi | 2eS PM De || ron | eee eco ces| wrlon i eeeseee | a pare ee ee ae GOSTO2 ESE SAL ea bale 28 UAE RQ fe esa sees |S eet | ee | ara ef a We SP eae el Or aal eeel eo ale ae fol Utada le Ss Sal bee Boce|esoh epee es ees eee | ee | ee |e el ee te Zee ioe |e eel ae | eee ALE HL epee | PLES Ie ON TA a ee ARE AE ek Ee | ee Sa ae eee 120-1 3OE ase | pee BR ca eee ee Re hk Se ee Al DE SeA Aye ee N40 150 8- — a2 |eecn| = 2s De Pe gD ee a a ef en eee I Sees | bs ees 1 TGOTI(GEe 4322 | sees rome WG OF |e | ER) ke ES ee A Beer Cee 2 Ee |e | a 2 TS0=100 Sees eee |e Th S39] ee EES So De eS ee eee Lil, ayia 1

26 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

PARUPENEUS PLEUROSTIGMA (Bennett) Puate 77,E

Upeneus pleurostigma BENNETT, Proc. Zool. Soc. London, vol. 1, p. 59, 1831 (type locality, Mauritius) Herre and Montrasan, Philippine Journ. Sci., vol. 36, p. 122, pl. 5, fig. 2, 1928.

Mullus pleurostigma PLAYFAtR, in Playfair and Giinther, Fishes of Zanzibar, p. 40, 1866.

Parupeneus pleurostigma BubEKeR, Atlas ichthyologique ... , vol. 9, pl. 393, fig. 3, 1878.—Werser and pe Beravrort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 398, 1931.

Pseudupeneus pleurostigma JeNKtNs, Bull. U. S. Fish. Comm., vol. 22, p. 456, (1902) 1903.—SnypxEr, Bull. U. S. Fish. Comm., vol. 22, p. 527 (1902) 1904.—Jorpan and EverMANN, Bull. U. S. Fish. Comm., vol. 23, pt. 1, p. 260, fig. 108 (1903) 1905.—Fow er, U. S. Nat. Mus. Bull. 100, vol. 12, p. 275, 1933.

Upeneus brandesi BuerKerR, Nat. Tijdschr. Nederl.-Indié, vol. 2, p. 236, 1851 (type locality, Banda Neira).

SPECIMENS STUDIED

Bikini Atoll: 9 stations, 44 specimens, 48 to 169 mm. in standard length.

Rongelap Atoll: 2 stations, 85 specimens, 49 to 104 mm.

In addition, 10 lots totaling 19 specimens were examined from the following localities: Philippine Islands, 2 lots, 5 specimens; Okinawa, 1 lot, 2 specimens; Hawaiian Islands, 7 lots, 12 specimens.

Description —The following counts and measurements were taken from 6 specimens unless indicated otherwise, all from the Marshall Islands and ranging in standard length from 80 to 151 mm. (counts and measurements of specimens from other areas of the Pacific are also included in tables 74-76): Dorsal rays VIIJ-i,8; anal rays I,i,6; pectoral rays ii,13 to ii,15(18 specimens) ; vertical scale rows 28; scale rows above lateral line 3; scale rows below lateral line 6; scale rows around caudal peduncle 14.

Body depth 3.7 to 3.9; head length 2.9 to 3.0; length of caudal peduncle 3.7 to 3.9; all in standard length. Eye 4.5 to 5.3; length of upper jaw 2.8 to 3.1; depth of caudal peduncle 3.5 to 3.9; length of snout 1.9 to 2.1; width of interorbital 4.5 to 5.1; length of barbel 1.4 to 1.7; all in length of head. Gill rakers 6 or 7+1-+21 to 23, total 29 to 32 (18 specimens); longest raker only slightly smaller than longest filament.

Teeth of the upper and lower jaws uniserial, blunt, stout and widely spaced; those in upper jaw about 32 and in lower jaw about 26; no teeth on vomer or palatines; scales on caudal fin, absent on dorsal and anal fins; barbels extend beyond vertical, touching posterior margin of eye but do not reach vertical touching posteriormost edge of preopercle; first dorsal spine about one-sixth length of second spine, second spine about three-fourths length of third spine, its tip not flexible; peritoneum light; lateral line complete; scales ctenoid, with

FAMILY MULLIDAE—LACHNER 27

4 to 6 radii in anterior field; pectoral fin pointed, posterior branch of last ray of soft dorsal fin slightly elongate, that of anal conspicuously elongate, about one-third longer than length of sixth anal ray.

Color in alcohol.—Head and body tan, dusky to dusky tan above, light below; barbels light; the characteristic color mark is a blackish irregular to rectangular spot or blotch located on the midbody area below the end of the spiny dorsal, or its center at a point where the longest ray of the pectoral fin touches the lateral line, this spot almost divided into halves by the lateral line, sometimes twice as deep as wide, greatest depth equal to length of snout; located immediately posterior to dark spot is a very light spot extending from half a scale row below base of soft dorsal fin to about half a scale row below lateral line; this light spot equal to or slightly longer than length of base of soft dorsal fin; a diffuse, dusky saddle is located posterior to last two rays of soft dorsal fin, extending to one scale row below lateral line and half the distance to end of caudal peduncle, very faint or completely obscure in some specimens; basal third of soft dorsal fin is black, remainder of fin light or transparent; all other fins are light or transparent. The young in transformation stages, taken at night, are dusky to blackish with only a faint dorsolateral spot (see remarks below).

Color in life-—The following color notes were taken from a koda- chrome transparency photographed by Dr. Donald Strasburg from a specimen about 6 inches in total length: Head and body light silvery, suffuse with orange-red on lips, snout, operculum, and dor- sally on head, and in lesser amounts dorsolaterally on body; a black spot dorsolaterally below end of spiny dorsal fin; spiny dorsal reddish; soft dorsal with an intense black basal bar on lower half of fin, the upper portion dusky to pale tinted with some orange; anal fin dusky, with some orange; caudal fin light orange at base, becoming lighter toward outer portion; barbels light; iris orange with some black, pupil deep black.

Range.—P. pleurostigma is a wide-ranging species that has been reported from the east African coast eastward to the Hawaiian Islands.

Remarks.—As is characteristic in several species of Parupeneus, the length of snout and barbel in pleurostigma increase with growth and the eye becomes smaller. The most abrupt change occurs at a length of 40 to 60 mm., probably associated with the transformation from the pelagic stage.

The color of 81 specimens, 49 to 56 mm. in length, from Rongelap Atoll, taken July 1, 1946, at night, is dusky to blackish, with the fins transparent to dusky. The black, dorsolateral spot on the body is faint, barely discernible in many of the specimens. The spot varies

a8 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

in size, just developing in some specimens. Another 157 specimens, chiefly smaller in size but ranging from 33 to 58 mm. in length, were taken with the above collection and are similar in coloration and morphological characters but lack the distinctive dark dorsolateral spot on the body. These are probably also pleurostigma, representing a stage just prior to the formation of the spot.

TasiLe 74.—Diameter of eye and length of snout as a percentage of standard length in Parupeneus pleurostigma, from the Marshall and Hawaiian Islands, by size groups

Standard Eye Snout

Jength

in mm,

5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

40-50 Re eA e Sa ee 4) she Bee 5 te ee Nie Se een ol ee ee | ee no ae ee ees 60=/9 22-22 |SssSa | 1 | Ste eee ee eee ea ee 2 Sse eae 1 Dae Leyte. Veen 8 ee lh ee 8000 bee Seem same 1 DI a here eee eS | ee | ee | cee | 2 A) ee A ee eee | ee 1OOH11 9S | aes oes 2 2 1 Se es | Eee See ee ee | See eee ee 2 eS | ee ek ees L20-139EE S| 2 oe ered 3 3a sb Se ern Ss Se Sa ees ees Soe | eee ee 1 4 Jes _ 2 eee 140-159. . .|---.- 2 i ee NC ee ied ee eS Se Soa oes oie, |e 2 Dale Se |e ae T60=HN/ Ose o See a sles Eee aS ea me eck aw ela epee cree ee cee ewe ale tae erp 9 |e eee 180-199___ Dy | ah ek ea ee eee eee yell oe ee a eee | Sa e eeee Ea) Se i eess =| rsa! ze ee 200-219 _ . . Dope aA Sa ee |e | SO Se ein eee ee Lo) ee eee 220-239 Ss a|C ee Dy eee ees ee ae ea bn ce JS SSS Re ee Sere aesee |e ose scene eee | ane 1

TaBLe 75.—Length of head and barbel as a percentage of standard length in Paru- peneus pleurostigma, from the Marshall and Hawaiian Islands, by size groups

Standard Head length Barbel length length ade ee Se 2S in mm. | |

98 | 29} 30!) 3 || 32 | 33: | 341 835 |) 15) 16) 17) 18 | 19") 20°} 20.) 22) 23 [324 sobs) 26

40-592 3522225) 1 |e (Dye te are et ea 3 1 ie Po | eR SS et | ees CORO eas ia se ae eileen ew 1 Pe Eee cee te a a 2.) eee Ee pees aes ee 80-09 Ss ee es See sy se mete > a Naan ied tiem cee fee Ue at a De Se i] Seen |eecel soem LOOK OSes 25 3S Pe ale Ae tee ra eA Rig | te ee | eet ae | ee 2 TA Se ec ee ieee L20-1S9k Ss = te ees ale eS 2 S80 2a Salad 2 3422S) E- Sea a eee es 1 1 2 1 hl zeud 40-1592 se See eee (Bees | al eee Re De hs ee | BP ee eas 1 1 1 | en SS |e NGO=U7 92 S32 252s ees pce ere al Bae SEA See ot ew Sa ee ee | ae |e 1) | 222 Esa Eee | aoe 180-199. 228) 22) eesti eee Mee all eee at Sel” See Bal | Se Sea eam hs by (ee fess fs ml ee 200-21 Os sees |e eee ea ek2 ol a ea ee AS SE ee ee eS | ee | Pf St a Ste Se Se 20 LOO eee eee |e eee | eee eens, ||| ee | | ee a] Ee a I a ea | ae ieee ee He | eee 1

Tasie 76.—Numober of gill rakers in Parupeneus pieurostigma, by locality Upper limb Lower limb Total 1 Locality ee 6 7 8 21 22 23 28 29 30 31 382

Okinawa Islands--._- 1 1 Ws |e as Dic ee lc Bicones 1 L222 22 | eee Se Marshall Islands- --_- 4 13 1 1 8 9 Ese Ss - 4 6 7 1 Hawaiian Islands_-_-__- 3 4) eae 1 4 2 1 1 4 1A ooo ee

1 The raker at the angle of the arch was included only in the total count.

FAMILY MULLIDAE—LACHNER 29 PARUPENEUS CYCLOSTOMUS (Lacepéde) Puiate 76,B

Mullus cyclostomus Lacerkp®, Histoire naturelle des poissons, vol. 3, pp. 383, 404, pl. 14, fig. 3, 1802 (type locality not given).

Upeneus cyclostomus Cuvinr and VALENCIENNES, Histoire naturelle des poissons, vol. 3, p. 472, 1829.—Hurre and Montaxzan, Philippine Journ. Sci., vol. 36, No. 1, pp. 127, pl. 6, fig. 3, 1928.

Pseudupeneus cyclostomus JonpAN and Seaxe, Bull. U. 8. Bur. Fish., vol. 25 (1905) p. 275, 1906.—Fow.esr, U.S. Nat. Mus. Bull. 100, vol. 12, p. 304, 1933.— Smrru, Sea fishes of Southern Africa, p. 230, 1949.—Fow.erR, Mem. Bishop Mus., vol. 12, No. 2, supp. 3, p. 95, 1949.

Mullus chryserydros LAcePmpE, Histoire naturelle des poissons, vol. 3, pp. 384, 406, 1802 (type locality, Mauritius).

Upeneus chryserydros Cuvier and VALENCIENNES, Histoire naturelle des poissons, vol. 3, p. 470, 1829.—Herre and MontaBan, Philippine Journ. Sci., vol. 36, No. 1, pp. 127, pl. 5, fig. 3, 1928.

Upeneus chryserythrus GUNtaER, Journ. Mus. Godeffroy, vol. 2 (Fische der Siidsee, pt. 3), p. 60, pl. 45, fig. A, 1873.

Parupencus chryserydros BLEEKER, Atlas ichthyologique . . ., vol. 9, pl. 393, fig. 2, 1878.—Wepser and pr Beavrort, Fishes of the Indo-Australian Archipelago, vol. 6, p. 404, 1931—Scuutrz, U. 8. Nat. Mus. Bull. 180, p. 180, 1943.

Pseudupeneus chryserydros JoRDAN and EvprMAnn, Bull. U. 8. Fish Comm., vol. 23 (1903), pt. 1, p. 255, 1905.—Jorpan and Sate, Bull. U. 8. Bur. Fish., vol. 25 (1905), p. 275, 1906.

Uneneus oxycephalus Bueexur, Act. Soc. Sci. Indo-Néerl., vol. 1, p. 45, 1856 (type locality, Manado, Celebes).

Parupeneus zanthospilurus BuerKker, Verh. Akad. Wet. Amsterdam, vol. 15, No. 3, p. 37, 1875 (type locality, Amboina).

Upeneus saffordi SEALE, Occ. Pap. Bishop Mus., vol. 1 (1900), No. 3, p. 71, 1901 (type locality, Guam).

Pseudupeneus aurantiacus Smaue, Oce. Pap. Bishop Mus., vol. 4, No. 1, p. 48, fiz, 14, 1906 (type locality, Tubuai, Austral Islands).

SPECIMENS STUDIED

Bikini Atoll: 11 stations, 13 specimens, 47 to 270 mm. in standard length. Eniwetok Atoll: 1 station, 2 specimens, 111 to 194 mm.

Rongelap Atoll: 4 stations, 8 specimens, 51 to 89 mm.

Rongerik Atoll: 1 station, 2 specimens, 59 and 77 mm.

Rota Island: 1 specimen, 172 mm.

In addition 20 lots totaling 28 specimens were examined from the following localities: East Indies and Philippines, 8 lots, 14 specimens; Admiralty Islands, 1 specimen; Phoenix and Samoan Islands, 3 lots, 4 specimens; Johnston Island, 1 specimen; Marquesas Islands, 2 lots, 2 specimens; Hawaiian Islands, 5 lots, 6 specimens.

Description.—The following counts and measurements were taken from 7 specimens, unless otherwise indicated (counts and measure- ments of specimens from other localities are also included in tables 77 and 78); dorsal rays VIII-i,8; anal rays I[,i,6; pectoral rays 11,14 to ii,15 (29 specimens); vertical scale rows 27 to 29, scale rows above

30 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

lateral line 3; scale rows below lateral line 7; scale rows around caudal peduncle 14.

Body depth 3.6 to 4.0; head length 2.8 to 3.0; length of caudal peduncle 3.6 to 4.1; length of longest pectoral rays 4.1 to 4.7; all in standard length. Eye, highly variable with size, comparatively smaller in the larger specimens, 4.9 to 7.1; length of upper jaw 2.3 to 3.4; depth of caudal peduncle 3.0 to 3.3; length of snout 1.7 to 2.0; width of interorbital 3.8 to 5.1; all in length of head. Eye in snout 2.4 (in smallest specimen) to 4.2 (in largest). Width of interorbital in eye, variable, 0.5 to 1.0. Length of barbel in length of head 1.1. Gill rakers, including rudiments, 6 to 8+-1+22 to 24, total 29 to 32 in 16 Marshall Islands specimens; longest raker about 1 to 1.2 in long- est filament.

Teeth of upper and lower jaws stout, blunt tipped, widely spaced, and in a single row; no teeth on vomer or palatines; scales on caudal fin, absent on dorsal and anal fins; cheek and opercle scaled; barbels long, reach end of head or to base of pelvic fins in most specimens; first dorsal spine minute, second dorsal spine almost as long as third, flexible at tip; peritoneum transparent; lateral line complete; scales ctenoid, with 5 to 6 radii in anterior field; last rays of soft dorsal and anal fins somewhat elongate.

Color in alcohol.—Body and head pale to light tan in most of the smaller specimens, and tan with some dusky in the larger ones. Fins are transparent in smaller specimens, transparent to slightly dusky in larger ones. Barbels are colored light tan. Body, head, and fins of 2 smaller specimens darker, dusky to brown. Light blotch on dorsal portion of the caudal peduncle posterior to soft dorsal fin present in about 40 percent of the specimens from the Marshall Islands. This spot is conspicuously developed in 2 specimens, faint in 2 specimens, questionable in 2, and not perceptible in all others; when present, its size is variable, usually extending half distance from end of base of soft dorsal to procurrent caudal rays, in smaller speci- mens, to about three-fourths of this distance, in larger ones.

Color in life—A brownish blue and a yellow color phase have been observed in life by Strasburg who recently collected in the southern Marshall Islands. These color phases correspond with the dusky (dark) and light color phases observed in some of our smaller preserved specimens. It cannot be determined at this time whether these phases are associated with size and maturity, or sex, or reproduction, or whether perhaps two distinct forms are involved. All our larger specimens have a uniform dusky to tan coloration.

The following color description taken in life at Wake Island by John E. Randall, University of Miami, from a specimen 313 mm. in length (USNM 167602) tentatively referred to this species: Body

FAMILY MULLIDAE—LACHNER Si

brilliant irridescent blue; head olivaceus, with an olivaceus area extending back from the head a short distance on the body; occasional olivaceus spots on body, especially posteriorly; a concentration of olivaceus blotches dorsally on caudal peduncle; these definitely not appearing as yellow in life, in the water or out; bright blue lines radiating from red eye; dorsal, anal, and caudal fins yellowish with lengthwise bright blue lines.

Range.—This species has an extensive geographical range, having been commonly reported as far west as the Red Sea and Hast Africa. Our material was taken from the East Indies eastward to the Hawaiian Islands.

Remarks.—This species represents one of the most complex specific problems among the Mullidae, owing to ontogenetic changes in certain morphological structures, such as the length of snout and the size and location of the eye, and to the probable color forms. The length of the snout and the size of the eye show the greatest change with increase in body length, the snout becoming relatively longer and the eye smaller (table 77). The larger specimens thus look quite different. The light spot, or saddle, just posterior to the soft dorsal fin is variable in its size and intensity of development in the adults, and sometimes it is not discernible. Certain specimens from the same locality also may represent two color forms, one having a darker body (bluish in life) and another having a light body (yellowish in life). Consequently the six nominal species in the above synonymy may represent different growth stages or different color forms of one stage.

The possibility that more than one of the nominal forms may represent a species is not yet satisfactorily ascertained, since sufficient comparative material was not available from some of the critical areas of the Indo-Pacific, such as the Indian Ocean and Madagascar. Additional data from living specimens are also needed from all areas of the Indo-Pacific to determine exact color and color patterns, so that possible specific differences may be detected.

The literature referring to this problem is confusing, because the names of two or more of the nominal species have been freely inter- changed, the descriptive accounts are not critical, the illustrations are often poorly drawn, and few attempts have been made to develop a concept of the probable mode of differentiation. Interesting dis- cussions concerning various nominal species have been presented by Giinther (1873, p. 60), Weber and deBeaufort (1931, p. 406), and Fowler (1933, p. 306).

Lacepéde (1802) originally described two of the most controversial nominal species, cyclostomus and chryserydros. Subsequent interpre- tations of these forms, coupled with the descriptions of additional ones, has lead to a confusion in the interpretation of possible valid species.

32 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

The figure of cyclostomus by Lacepéde (1802, pl. 14, fig. 3) shows no light spot dorsally on the caudal peduncle.

Bleeker (1878, fig. 2) recognized chryserydros as a species with an elongate light spot or saddle extending from the posterior base of the soft dorsal to the procurrent caudal rays, whereas he illustrated (1878, fig. 5) the nominal form Pseudupeneus xanthospilurus with a light spot extending only half this distance. He also illustrated his chrys- erydros with a shorter snout, bluish horizontal stripes on the head, and gold and blue stripes on the soft dorsal, whereas the figure of zanthospilurus has a longer snout and lacks the color markings. Both Schultz, who collected many of our Marshall Island specimens, and Strasburg, observed no stripes on the head or any other outstand- ing color markings on the head or body in live specimens, but Stras- burg did observe a dusky and a reddish color phase.

From the same subfaunal area from which Bleeker’s specimens were taken, Weber and de Beaufort (1931, pp. 404, 407) recognized chrys- erydros and cyclostomus as probable species, but had no specimens of the latter. In an adjacent area Herre and Montalban (1928, pp. 123, 127) recognized from the Philippines cyclostomus and chryserydros as distinct forms based on two specimens of each. They illustrate cyclostomus as yellow-orange and lacking the light caudal saddle and chryserydros as having a purplish body. Fowler (1933, p. 304) lists all his material from the Philippines under cyclostomus, but regarded (p. 309) one specimen (disposition not stated) as zanthospilurus. He also recognized (p. 309) the nominal species chryserydros from Mauri- tius, but he had no specimens.

Sauvage (Histoire naturelle des poissons, in Grandidier, Histoire ... de Madagascar, vol. 16, p. 226, pl. 26, figs. 3 & 4, 1891) dis- tinguished the two nominal forms cyclostomus and chryserydros but illustrated no light dorsal saddle on the caudal peduncle. Smith (1949), who recently has carried out considerable collecting in an area adjacent to Madagascar, recognized only cyclostomus.

One or two forms have also been recognized from areas of Oceania. Jordan and Seale (1906, p. 275) noted two species from Samoa, chryserydros and cyclostomus; the general color of the latter was observed as “clear red without markings” and a “pale saddle on the tail.’ From the same area Schultz (1943, p. 130) recognized one species. Jordan and Evermann (1905, p. 255) reported one form in the Hawaiian fauna and Fowler’s more recent list of fishes of Oceania (1949, p. 95) also reports only one form.

The nominal species Upeneus orycephalus Bleeker, U. saffordi Seale, and Pseudupeneus aurantiacus Seale, are only tentatively con- sidered as synonyms of cyclostomus.

33

FAMILY MULLIDAE—-LACHNER

ee rr a eee SOSREEEESTT FUESESEEIET GREET TEESEENEEDT BERGER [CSSESET TEENY [RESET] (GUEST

Pee aera eee al | eae ee ie seis le oa ia Aa eae ea Si sae lees acel sqealie | elie I |-77--]---7"]-~ 668-098 eee | ara leee seen nema meal occa [aac ie eeee lps nate mee ge eee ee ed oie aie fal aan lie gl) eee el ealioge le 2 al Bee lle eae ga 698-028 Ber ieee [esis Sele ee ee | ae eee Toe | es we eaten a eee lee nin le aalee, cle see ie cal ee lee al a I |--77-|--618-082 Saee (peal ese elena loose |Pee" (ser ale Senco a Vee ere es tal ber les elise aes Toda Seer ee ee cE oe [egies aoe I |-77-7]""622-08% Foo ee lle ca oe | ee ee Te Qe lone (Go | pee Ten Ale ra a Fe foil? 235| me saliegee|, esl ee Ta go sol ee Ts alles | nee -00e Toma BOs at Were Seir seal cies oie aes Co USapl te lel ee [cane Se te Mt el ates |S eee ea lees aah lees aes 2 |@ |. |---7-|-766t-09r Tose staal fatale ceca || ale Ose ep lige soe aoe sclera SM BP ae ae ES el faa TSho| es TE \oieslaee ai = [-209T-0er ong | ees peale seme ele es lame al Paes) lee (prea ele (ee [oe lod a (aT lear [realises ea nie eee NIC eee Netra e608 Ties os less tometer =" [seer lapy sales |e osle ees eae ee eee A eee pee ek — | ee Teele |e mee a Tee [ec RIT pte tieal

‘TU UT 339] qnoug oA poqued peoH prepueyg

sdnoib az1s fig vvwna0g fo spunjsr pun saurddywyd ay} wouf snwioysopso snousdnieg u2 yjbua) punpunys fo abnjuaosad v sp 11910 fo dajawnryp pun jnous pup “aging ‘poay fo yybuaT—'}) ATAV I,

34 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

The number of gill rakers in East Indies and Philippine specimens averaged less than those from Oceania; and the light pedunclar saddle was conspicuously developed in all specimens from the East Indies and Philippines, whereas this was true of only 40 percent of the specimens from the Marshall Islands. These data suggest that the divergence between these two geographical areas is on a racial level. These differences could not be correlated with the dark and light color phases discussed above.

A species, P. chrysonemus, with a superficial appearance of the cyclostomus complex and known only from the Hawaiian Islands, is distinguished from the latter mainly by its comparatively shorter snout, larger eye, and higher number of gill rakers (see tables 59 to 62).

TaBLe 78.—Number of gill rakers in Parupeneus cyclostomus, by locality

Upper limb Lower limb Total! Locality ae 5 6 it 8 DN ire 22) | P23, Wo 24s 27 eon 200 | OO olen mos East Indies-

Philippinesssoss=2 2 252 1 11 2) 4 4 Giulia sae 1 2 4 de eee eae Marianas Island= 2.2 2252 )22222 15 |e Sa ae | eee | Paes Di) 2 OE oe | ee eee Pesce Marshall Islands---------- 1 8 6 1 ees 3 8 yj] eel ees 1 3 10 2 Samoan Islands----------- ae 1 Golesees esse 1 3 1 | | eee 1 | ease Johnstoniisland ==. 22-2222. |= aS o8 Dyce stissese LN Seca ees eS eC eee ae dieses eee Hawaiian Islands_......---|----- 3 Be feessa} ste 1 3 2h | eee ae eee 3 2 1

1 Raker at the angle of the arch was included only in the total count. PARUPENEUS CRASSILABRIS (Cuvier and Valenciennes) Puate 76,E

Upeneus crassilabris Cuvier and VALENCIENNES, Histoire naturelle des poissons, vol. 7, p. 523, 1831 (type locality, New Guinea).—Smirx and Swany, Proc. U. S. Nat. Mus., vol. 5, p. 129, 1882.—Jorpan and EverMann, Bull. U. 8. Fish. Comm., vol. 23 (1903), pt. 1, p. 259, 1905.

Pseudupeneus crassilabris FowuLER, U.S. Nat. Mus. Bull. 100, vol. 12, p. 315, 1933.

SPECIMENS STUDIED

Bikini Atoll: 1 specimen, 203 mm. in standard length.

In addition, 1 specimen (USN M 26824), 265 mm. in length, from Johnston Island, was also examined.

Description.—The following counts and measurements refer to the specimen collected at Bikini Atoll, those in parentheses refer to the Johnston Island specimen: Dorsal rays VIII-i,8 (VIII-i,8); anal rays 1,i,6 (1,i,6); pectoral rays ii,14 (ii,15); vertical scale rows 28 (27); scale rows above lateral line 3 (3), below lateral line 7 (7), around caudal peduncle 14 (14).

FAMILY MULLIDAE—LACHNER 35

Body depth 2.9; head length 3.2 (8.0); length of caudal peduncle 3.8 (4.4); length of longest pectoral ray 4.0; all in standard length. Bye 5.3 (5.1); length of upper jaw 2.6 (2.3); depth of caudal peduncle 2.3 (2.8); length of snout 1.8 (1.7); length of barbel 1.5 (1.5); width of interorbital 3.2 (3.5); all in length of head. Eye in snout 2.9 (2.9). Interorbital width in eye 6.1 (6.8). Gill rakers 0, 8+1+26, 2 (0, 9+1+27, 2), totaling 37 (39); longest raker about 0.9 length of longest filament.

Teeth of upper and lower jaws uniserial, blunt, stout and widely spaced; no teeth on vomer or palatines; scales on caudal fin, absent on dorsal and anal; cheeks and opercles scaled; barbels reach vertical touching posterior margin of preopercle; first dorsal spine small, about one-seventh length of second spine; second dorsal spine about four- fifths length of third, tip not flexible; peritoneum transparent; lateral line complete; scales ctenoid, with 4 to 9 radii in anterior field; last ray of soft dorsal fin slightly elongate, that of anal same length as other rays of fin.

Color in alcohol.—Head and body dusky to dusky tan. A diffuse blackish mark or stripe on each side of snout from middle of eye to middle of upper jaw. Some diffuse brownish coloration on chin, cheek, and opercle. An irregular light tan to brown coloration en- circles spiny and soft dorsal and is broken in various places, forming irregular spots. The pectoral fin is transparent; the spiny dorsal is transparent, tipped with black; the soft dorsal is transparent, with about 6 dusky to blackish horizontal streaks; the 3 streaks near base of fin formed by nearly circular spots; the tip of the soft dorsal bor- dered with blackish color; the caudal is dusky, the outer ray edged in white; the anal fin is similar to soft dorsal except the spots and streaks are not so pronounced; pelvic fin light near base, dusky towards tips, the outer edge of first ray and tips of first three rays edged in white. Barbels dusky near base, light brown near tips. The Johnston Island specimen has a barbel blackish from base to tip.

Remarks.—This species has been considered synonymous with Upeneus luteus Cuvier and Valenciennes (Histoire naturelle des poissons, vol. 7, p. 521, 1831, type locality, Mauritius) by several authors. The problem is partially reviewed by Weber and de Beau- fort (Fishes of the Indo-Australian Archipelago, vol. 6, p. 404, 1931). Certain facts from my data and from the literature lead me to disagree with this contention. First, the characteristic dusky to blackish coloration of the specimens from Bikini and Johnston Island has not been reported by other authors for specimens of luteus; and second, the number of gill rakers in these specimens is much higher than that reported in published accounts of luteus. Weber and de Beaufort (1931, p. 401) found 20 gill rakers, including rudiments, on the lower

36 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

limb of lutews, and merely listed crassilabris, quoting the account of Cuvier and Valenciennes (1831, p. 523), since they had no specimens. Our specimens of crassilabris have 28 and 29 rakers (table 59) on the lower limb. From my study of the variability of gill rakers in various species of Parupeneus, the differences of counts of 20 and 28 to 29 for the lower limb are so great as to warrant the statement that our speci- mens certainly cannot be compared with those reported on by Weber and de Beaufort. Fowler (1933, p. 313) listed luteus but had no specimens from the area studied, and his single specimen of crassilabris was that specimen cited above from Johnston Island.

Day’s illustration of luteus (Fishes of India, p. 125, pl. 31, fig. 2, 1878) resembles our specimens of crassilabris in general body form but the dark margins of the fins, characteristic of our specimens, was not shown. Bleeker’s figure of luteus (Atlas ichthyologique . . ., vol. 9, pl. (4) 394, fig. 1, 1877) shows numerous light spots on the body, arranged in about 8 horizontal rows; these spots are not present on our specimens of crassilabris.

Fowler (Acad. Nat. Sci. Philadelphia Monogr. 2, pp. 124, 201, and 285, 1938) lists the distribution of crassilabris as Johnston, Jarvis, Christmas, and Apataki Islands. It has not been reported from the Hawaiian Islands.

A specimen, USNM 71651, 130 mm. in standard length, taken by the Bureau of Fisheries off Japan, might represent P. luteus: Its head and body are brownish and the fins dusky; the scales have a dark brown spot located near the center and slightly smaller than the pupil; and these spots, which form about 8 to 9 horizontal lines on the side of the body, are more evident on the caudal peduncle. Otherwise, this specimen resembles the darker specimens of P. cyclostomus in our collection. The following counts and measurements were also taken: Vertical scale rows 28; scales around caudal peduncle 14; dorsal rays VIII-i,8; anal rays 1,1,6; pectoral rays 1,14; head length 3.0, length of caudal peduncle 4.0, both in standard length; eye 6.0; length of upper jaw 2.9, length of snout 1.7, interorbital width 4.3, length of barbel 1.1, all in length of head. Gill rakers 1,5+1-+-18,5, totaling 30.

Genus MULLOIDICHTHYS Whitley

Mulloidichthys WuitLEy, Rec. Australian Mus., vol. 17, No. 3, p. 122, 1929 (type species, Mullus flavolineatus Lacepéde). (Mulloidichthys Whitley, proposed to replace Afulloides Bleeker.)

Mulloides (not Richardson, Rep. Fifteenth Meet. British Assoc. Adv. Sci., p. 16, 1846) BLeEKerR, Verh. Bataviaasch Gen. vol. 22, p. 6, 1849 (type, Mullus flavolineatus Lacepéde).

The characters that best define the genus are associated with the

teeth: dentition incomplete; teeth in the jaws in villiform bands, widest anteriorly, where they are in several irregular rows, and

FAMILY MULLIDAE—LACHNER 37

tapering posteriorly to one or two more or less irregular rows; no vomerine or palatine teeth.

The following characters were common to all species of Mulloi- dichthys studied and apply to some of the other genera in the family: Scales on caudal fin, absent on dorsal and anal fins; dorsal fin rays VIII-i,8 the first spine minute, pectoral fin rays 16 to 18; scales above lateral line 3, below lateral line 6 or 7, around narrow portion of caudal peduncle 15 or 16; first elongate spine of spinous dorsal flexible near tip.

This genus inhabits the tropical and subtropical littoral marine waters of the Indo-Pacific, Eastern Pacific, and Western Atlantic regions. At least four species are known from Oceania, three from the Eastern Pacific and one from the Western Atlantic regions. Three species were taken in the Marshall Islands and one from the Marianas.

Characters useful in distinguishing the species are summarized in tables 79 and 80.

No appreciable population divergence was found when comparing various meristic counts, measurements, color, or color pattern in those species where sufficient specimens were available for study from various subfaunal areas. The data comparing the total number of gill rakers showed a tendency toward an East to West gradient when a few specimens were compared, but this was insignificant when larger samples (see descriptions of samoensis and auriflamma) were available.

Several species reported from Oceania are so little known that their generic allocation is even doubtful. Pseudupeneus eutaematus Fowler (Proc. Acad. Nat. Sci. Philadelphia, vol. 96, p. 170, fig. 21, 1944, type locality, New Hebrides) may represent a species of Mul- loidichthys, for his statement, “Teeth minute, uniform close-set, form an apparent row in each jaw, none on palate,’”’ comes closer to describing the dentition of Mulloidichthys. He may have examined only the villiform tooth patch, which tapers to a row or two of small teeth posteriorly in both jaws. Otherwise eutaeniatus could only be referred to the genus Parupeneus. His count for the total number of gill rakers (18) is considerably lower than for any species known from Oceania, other than certain Upeneus where the lowest count was found to be 19. This may suggest that eutaeniatus may represent a species of Upeneus. The small palatal teeth in this genus can be easily overlooked. Pscudupeneus eutaeniatus Fowler, is known only by the type, which was not available for this study.

Another species which may be referred to the genus Mulloidichthys is Upeneus bilineatus Fowler (Bernice P. Bishop Mus., Bull. 38, p. 17, fig. 2, 1927, Palmyra Island). Fowler listed the dentition as,

38 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

“teeth small, simple, conic, uniform, form two rows above anteriorly and five below anteriorly; none on palate or tongue.”’ Herre (Field Mus. Nat. Hist., Zool. Ser., publ. 353, vol. 21, p. 213, fig. 11, 1936) reported it from Nuku Hiva Island, Marquesas Islands, as Paru- peneus bilineatus. He commented on his generic allocation (p. 211) referring bilineatus to species with teeth arranged uniserially in the jaws. Fowler (Mem. Bishop Mus., vol. 10, p. 233, fig. 47, 1928) cites the original description of bilineatus by Cuvier and Valenciennes (Histoire naturelle des poissons, vol. 7, p. 394, 1831, type locality, Amboina), pointing out their reference to the uniserial conic jaw teeth and smooth palate, and erroneously places his Palmyra record with it without mention of his previous statement on the dentition.

TaBLE 79.—Number of gill rakers and vertical scale rows of Mulloidichthys from various Indo-Pacific localities

Totai number of gill rakers Vertical scale rows Species : pe | A eee 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 33 | 34 | 35 | 36 | 37 | 38 | 39 samoensis.—=22--_ 1 fe Sie Pi es) less ed te Ul De? 27a We Sa TA SAT) PR | 1 7} 15} 30 | 12 2s DUTACOLETIS I Sane ena | ee 2 ae hat DTN | s Sull eel ee all ee | ee Sale Sale | aaa 1 DAUGETR rae ee eae ee eee 1 1 | es SSS Ee | eer et eee ees |e ed | ee ees 25 gee CUPL Fos oe he) eae eee Oe |e 1 4 818s | 2251200), 16i|e eee 6 |S Say l2i heb e2ee

TABLE 80.—Measurements of several characters in four species of Muiloidichthys

In percent of standard length

Species Length of head Depth of body

26-27 | 28-29|30-31 |32-23) 34-35] 36-37) 18-19) 20-21) 22-23) 24-25} 26-27 | 28-29] 30-31| 32-33

SAIMOeNSIS! 22) eee ad 1 15] 2 4 1h) tes 9 3 9 Dae se ee 1 Vanicolensis22— 22.4 3-2 23 9 Dj fcsk | ee eee 2 11 ais see EE see ee eee DAU ger ea ae ee ele Bi Se a ee | ee ee ee 2 Lo Bere Se aes gurifiammsa = oan Bee eee 2 14 14 2 | ee eee 2 2, 1 11 6 De s=25

In percent of head length

Species Length of barbel Interorbita! width

SAINIOCTISIS! Soe nae eo: ae | |e cee 2 mek i Bianco ee 10] 138 4a see 1 VANICOIENSIS2 eee 2 16 Gxizeee hee alee ci Sarah eee eel ee See A 10 12 1 Pin eerie s oe we oe Pe ee eee a 1 2 eases ee | | 1) |e

auriflamima: = ose eet eee eae ae 1 8 11 9 4 eneee | hae ae 1

FAMILY MULLIDAE—LACHNER 39

TABLE 80.— Measurements of several characters in four species of Mulloidichthys— Continued

In percent of head length

Species Length of upper jaw Diameter of orbit

26-27 28-29130-31 32-33/34-35 36-37|38-29] 20-21 | 22-23 | 24-25) 26-27| 28-29|30-31/32-33

SAMOECNSIS2 2 Ja 4 220-- 2 eo 1 9 10 PS ieee eee 1 3 5 13 5 Dien Vvanicolensis2a-22-22-255—.4 1 10 LO ip =F | ae i ee 1 7 14 2 ese ee SEA CES er ae eee eee ee Wee 2 1 1 dee ee ae eee eres SUniamma ee oe ee ce eae oes 1 9} 15 5 1p fo e S 1 Seal 10 10

Species 30-33 34-37 38-41 42-45 46-49 50-53 54-57 58-61 Samoensis:© 3545) ee ee aa eee ee 8 8 19 5 1 1 VaEMiCOlORSISS s=-o 2-220 epee sen 15 ORE ee SER C he eae See aR ee a oe pilueeriea-- =e Aes eee RHE ees A etal ele on 2 Abe Nir a ie SSeS | or arcs aurifiammavsss-22-22-5— 20 1 3 16 10 5

KEY TO THE SPECIES OF MULLOIDICHTHYS FROM OCEANIA

la. Peritoneum dark, dark brown to black.

2a. Black spot on side of body below spinous dorsal fin sometimes present; black blotch or spot on inner side of operculum bordering pseudo- branchiae usually present; fins pale or transparent; total number of gill rakers range from 24 to 31, modally 28; snout long and moderately rounded, 38 to 58, eye small 21 to 30, barbel intermediate in length 55 to 72, all in percent of head length; head length 29 to 36 in percent of standard length; vertical line touching end of jaw clearly in front of MAreins.OF ONOMb Les 2 ah M. samoensis (Giinther)

2b. No black spot on body nor on inner side of operculum; fins pale to trans- parent; total number of gill rakers range from 29 to 35, modally 33; snout short and round, 33 to 48, eye large, 25 to 33, barbel long, 65 to 80, all in percent of head length; head 28 to 34 in percent of standard length; vertical line touching end of jaw nearly touches orbit. M. auriflamma (Forsk&l)

2c. No black spot on body nor on inner side of operculum; striations on soft dorsal and anal fins usually blackish, creating a net-like appearance;

total number of gill rakers 27 to 30, modally 29; snout short and pointed, 35 to 39, eye 23 to 28, barbel short, 48 to 55, all in percent

of head length; head short, 26 to 29 in percent of standard length.

M. vanicolensis (Cuvier and Valenciennes)

1b. Peritoneum light, transparent to silvery-white; body and fins with uniform coloration, lacking prominent pigmentation; total number of gill rakers

422142—60—-4

40 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

27 to 29; snout 43 to 46, eye 21 to 24, barbel 64 to 67, all in percent of head length; head length; 30 to 31 in percent of standard length; plate (oe, Ce emeeere eae ee ee M. pflugeri (Steindachner)

MULLOIDICHTHYS SAMOENSIS (Giinther)

PuaTE 78,A

Mulloides samoensis GUNTHER, Journ. Mus. Godeffroy, vol. 2 (Fische der Stidsee, pt. 3), p. 57, pl. 43, fig. B, 1873 (type locality, Apia, Samoa). Upeneus preorbitalis Smirn and Swatrn, Proc. U.S. Nat. Mus., vol. 5, p. 132, 1882 (type locality, Johnston Island; holotype USNM 29662 examined). SPECIMENS STUDIED

Bikini Atoll: 29 stations, 188 specimens, 77 to 238 mm. in standard length. Eniwetok Atoll: 8 stations, 81 specimens, 74 to 222 mm.

Rongelap Atoll: 11 stations, 171 specimens, 75 to 210 mm.

Rongerik Atoll: 6 stations, 53 specimens, 80 to 190 mm.

Guam: 7 lots, 18 specimens, 72 to 240 mm.

In addition, 63 lots totaling 283 specimens were examined from the following localities: Red Sea, 2 lots, 3 specimens; East Indies and Philippines, 22 lots, 34 specimens; New Guinea, 1 specimen; Bonin Islands, 2 lots, 2 specimens; Fiji Islands, 1 specimen; Phoenix and Samoan Islands, 14 lots, 76 specimens; Society Islands, 4 lots, 19 specimens; Fanning Islands, 1 specimen; Johnston Island, 3 lots 23 specimens; Hawaiian Islands, 13 lots, 123 specimens,

Description.—Counts and measurements were taken from 10 speci- mens, all from the Marshall and Marianas Islands unless indicated otherwise, ranging in length from 77 to 242 mm. (additional counts and measurements, including specimens from other localities, are con- tained in tables 80-82): Dorsal rays VIII-i,8; anal rays I,1,6; pectoral rays 16 or 17; vertical scale rows (67 specimens), 33 to 38; scale rows above lateral line 3, below lateral line 6 or 7, around caudal peduncle 16.

Body depth 3.6 to 4.7 (23 specimens); head length 2.8 to 3.5 (44 specimens); length of caudal peduncle 4.1 to 4.3; length of longest pectoral ray 4.7 to 5.0; all in standard length. Eye, 3.3 to 4.5 (28 specimens); length of upper jaw, 2.7 to 3.6 (22 specimens); depth of

17 Mulloides pflugeri Steindachner, Denkschr. Akad. Wiss. Wien, vol. 70, p. 485, pl. 3, fig. 4, 1901 (type lo- cality, Honolulu).—Jordan and Evermann, Bull. U. 8. Bur. Fish, vol. 23 (1903), pt. 1, p. 251, 1905.—Jordan and Snyder, Bull. U. S. Bur. Fish., vol. 26 (1906), p. 214, 1907.

Mulloides flammeus Jordan and Evermann, Bull. U. 8S. Fish. Comm., vol. 22 (1992), p. 186, 1903 (type local- ity, Kailua).—Jordan and Evermann, Bull. U.S. Fish. Comm., vol. 23 (1903), pt. 1, p. 251, fig. 104, 1905.

Our collections total 3 specimens, USNM 55516, Hawaii; USNM 151522, Honolulu; and USNM 188601, Awmboina, East Indies. The latter specimen is tentatively referred to this species. Although it agrees closely with the Hawaiian specimens in respect to counts, measurements and coloration, there is a possi- bility it may represent a distinct form. Several species in the family which appear to be similar in alcohol have characteristically different life colorations. M. pflugeri has been reported only from the Hawaiian Islands.

FAMILY MULLIDAE—LACHNER Al

caudal peduncle 3.4 to 4.0; length of snout, 1.7 to 2.6 (42 specimens) ; interorbital width 3.0 to 4.3 (28 specimens); length of barbel 1.4 to 1.8 (40 specimens); all in length of head. Eye in snout 1.3 to 2.1. Interorbital width in eye 1.0 to 1.4.

Gill rakers 7 to 9+1+16 to 21, total 24 to 31 (150 specimens); longest raker about 1.4 to 2.0 in longest filament; peritoneum black; second dorsal spine flexible near tip; barbels extend posterior of verti- cal through eye but rarely reaching preopercular margin.

Color in alcohol—Head and body pale, darker above and usually light silvery below. Faint traces of a light median horizontal stripe from eye to base of caudal fin. A black spot or blotch on side of body below spiny dorsal fin sometimes present, often faint or obscure. Fins pale or transparent. A black blotch on inner side of operculum bordering pseudobranchiae usually present, sometimes faded.

Color in life-—These color notes were taken from a kodachrome loaned by Dr. Strasburg, U.S. Fish and Wildlife Service, Honolulu: The specimen was 5% inches in total length. Cheek, operculum and dorsal portion of head blackish; snout diffuse orange-red; remainder of head silvery; body light silvery below, becoming dark above; spinous dorsal, soft dorsal, and caudal fins dusky to orange; pectoral fins reddish orange; pelvic fins dusky to black near base, orange on outer portion; anal fin with black spot near base, remainder reddish orange; barbels light blue at base, remainder light orange; iris orange to silvery; pupil black.

Range.—The extensive geographic range of M. samoensis parallels that of M. auriflamma, occurring from the Red Sea eastward through the East Indies, Philippines, and the islands of Oceania through the Hawaiian Islands.

Remarks.—This species is very closely related to Mf. auriflamma with which it can be easily confused, especially the smaller specimens or those poorly preserved. The black blotch present only on the inner side of the operculum of samoensis is a good character to dis- tinguish between most specimens of the two species but it is sometimes faded. The black spot on the midside of the body is often completely faded. The diagnostic characters most useful in differentiating these species are the presence of these two spots in combinations with differ- ences in the number of gill rakers, length of barbel, size of eye, the length of snout.

No significant differences were found in an analysis of populations from several localities throughout the Indo-Pacific on the basis of comparing color, meristic counts, and proportional measurements.

42 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

TaBLE 81.—Number of gill rakers in Mulloidichthys samoensis, by locality

Upper arch! Lower arch ! Total Locality dil neat

7 8 9 | 16] 17] 18] 19 | 20 | 21 | 24 | 25 | 26 | 27 | 28 | 29 | 30] 31 Red Stal ste areloe eee ee Sold e le Tea Meo es sole Oe fae, Bic cea LA 23 =2ee eee a ae | ee Basti Indilesit-c222s-ceseee Le ABs cer eae 1 G5 Cease | A ee ole oe 2a) oa eee ee Philippines:- 3.222252 3 | 18 PF Sea eet ALS eee el a ees 5 | 11 Sere se ade (eet New; Guinea <2. - 222 = se Sele See Dieses eee Gabe 2) ca ee a a See Mey aictee [lee Bonin=s 2222s. se ee |e 1 he eee aoe 1 U1)| psec ae | ee | ee 1 ee ees Marianas Islands_-_------- Sq ones eeee (eel 4! 3 te] 2 Poor Lay Ss 1 5 1 1 Py eee Marshall Islands_.--__-_-- 4] 15 5 Je eset 44) 16 2 aes ined 1 4/14 2 2) eae GS eee ae ee ease le a1 | rata | ee es | U8 | eee ce | | a | ee | TA eesa| es sal eeoe Phoenix and Samoan Is-

Jandss) Fis eek 3 | 16 Beie===1. 55 6 | 12 Di free ees 2 4 7 6 4 Wot Society Islands_-___...---- 37 |e Salen sass ese De Salet oat | ee | eee | ee 25 \CoL ol Soa 1 Hanning Tslands. 2-22 4 ee Le See ae ees DA Oa] AS eee | ES ee Ee TSE ate gohnston Island #=.2- 222-2 Hy -13) | Se eeealy La POs SSS Bish eee ees Dy Lule dail we Seaheaten| eee Hawaiian Islands-...------ Qh | a0 | pees ace OE L251 OU, 2 6 | seen | Sees | aoee Sul Loy eee 1

1 Gill raker at the angle of the arch was included in the total count only.

TABLE 82.—Change in length of snout with increase in body length in specimens of Mulloidichthys samoensis from 81 to 290 mm. in length

Length of snout in percent of head length Standard length (mm.)

38-41 42-45 46-49 50-53 54-57 58-61 essithans100f22- es cee eee 8 5 Bul cees sees 2 eon ees ee | ee eee MOO-}4 Oe oo am ee ote cas eee Se eesee 3 4 Lae. 2 | See T5O=1 99 Ue oat 2 I ee eee FOS se 12 2. sete eee 1 200=200 she 2 2s 2h) Seas ee Ae Se | ee 4 Daa aed

MULLOIDICHTHYS AURIFLAMMA (Forskal)

PuatE 78,B

Mullus auriflamma Forsk&u, Descriptiones animalium ... , p. 30, 1775 (type locality, Djedda, Red Sea).

SPECIMENS STUDIED

Bikini Atoll: 9 stations, 41 specimens, 89 to 225 mm. in standard length.

Eniwetok Atoll: 1 specimen, 170 mm.

Rongelap Atoll: 2 stations, 21 specimens, 93 to 117 mm.

Rongerik Atoll: 1 station, 3 specimens, 99 to 112 mm.

Thirty lots totaling 93 specimens were examined from the following localities: East Indies Islands, 6 lots, 14 specimens; Philippines, 1 specimen; New Guinea, 1 specimen; Bonin Islands, 1 specimen; Phoenix and Samoan Islands, 14 lots, 67 specimens; Tuamotus, 1 specimen; Johnston Island, 2 lots, 2 specimens; Hawaiian Islands, 4 lots, 6 specimens.

Description.—Counts and measurements were taken from 10 speci- mens, all from the Marshall Islands unless indicated otherwise, ranging in length from 89 to 225 mm. (additional counts and measure-

FAMILY MULLIDAE—LACHNER 43

ments including specimens from other localities are contained in tables 80 and 83): Dorsal rays VIII-i,8; anal rays I,i,6; pectoral rays 16 or 17 (15 specimens); vertical scale rows, 35 to 38 (30 specimens) ; scale rows above lateral line 3, below lateral line 7, around caudal peduncle 15 and 16.

Body depth 3.3 to3.8 (23 specimens) ; head length 3.0 to 3.6 (32 speci- mens) ;length of caudal peduncle 4.3 to 4.6 ; length of longest pectoral ray 4.5 to 4.8; all in standard length. Eye 3.1 to 3.9 (385 specimens); length of upper jaw 2.6 to 3.3 (31 specimens) ; depth of caudal] peduncle 3.0 to 3.7; length of snout 2.1 to 2.8 (35 specimens); interorbital width, 3.4 to 4.0 (22 specimens); length of barbel 1.3 to 1.6 (83 speci- mens); all in length of head. Eye in snout 1.4 to 1.6. Interorbital width in eye 1.0 to 1.3.

Gill rakers 7 to 10-++-1+20 to 25, total 29 to 35 (79 specimens); longest raker about 1.2 to 1.6 in longest filament; peritoneum deep black; second dorsal spine fairly flexible near tip; barbels extend to preopercular margin or slightly beyond.

Color in alcohol.—Head and body pale, or light to slightly golden brown, often dusky dorsally. Head and body in some specimens with irregular dark blotches. Fins pale to transparent. Our specimens show no remains on the body of the median yellow horizontal stripe present in life.

Range.—This species has an extensive geographic range. Originally described from the Red Sea, it has been found from the East African coast (Smith, Sea fishes of Southern Africa, p. 231, 1949) eastward through the East Indies, Philippines, Islands of Oceania, and the Hawaiian Islands.

Remarks.—The color and morphological characters of populations of this species in various geographic localities appear homogeneous.

Taste 83.—Number of gill rakers in Mulloidichthys auriflamma, by locality

Upper arch ! Lower arch 1! Total Locality ee SS ee a 718 19 | 10 | 20 | 21 | 22 | 23 | 24 | 25 | 29 | 30 | 31 | 32 | 33 | 34 | 35 Mast Mdiesssoe-o ees ceee eae SL Genas 2 OMe On aces =e ace TOA econ acon eae | ete IPHilippinestsss. ses ee alee eeee Ms ee DRA AE T8 ee Hs eee a eee i ee Se FIs ee New, Guinea oi... S. 2] oe SAE of eC I es Pe a oe oe OL a | all a AESOTIMPES LAT CLS sere aera eran | eee ea fae Hs | ees ae ce Ea ese | errs | ees eee [eee [bers 1 Marshall Islands-_-_------- TP SH QE, KMGK See Sess Balezouenre LEO) Peete ae: SAR Be. 5 Phoenix and Samoan | Wslandst 2222s ese eet a ass 4 8 1 1 2 6 | 2 Se Foak 1 2 2 4 2 Muamotuesland assess === | pee |= LG | es | ee | ee ee 15] DR Ree hae eel eee SEES eT 1 Johnnstonwsland=s =e) 22) se ee eee Ss a a te oe aN aa eS 2 Hawaiian Islands_...------|---- De a a ee Zales ee eee Qe | ode ly, ok 1

ee a rrnrannn TEETER EEEEREEEEEEREEIEEEREETEEEEERED

1 Gill’raker at the angle of the arch was included in the total count only.

44 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

MULLOIDICHTHYS VANICOLENSIS (Cuvier and Valenciennes)

Upeneus vanicolensis Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 7, p. 521, 1831 (type locality, Vanicolo, Santa Cruz Islands). SPECIMENS STUDIED

Bikini Atoll: 8 stations, 26 specimens 77 to 96 mm. in standard length.

Eniwetok Atoll: 4 stations, 53 specimens, 77 to 92 mm.

Rongelap Atoll: 3 stations, 15 specimens, 79 to 93 mm.

Description.—Counts and measurements were taken from 7 speci- mens all from the Marshall Islands, unless indicated otherwise, rang- ing in length from 84 to 95 mm. Dorsal rays VITI-i,8; anal rays I,1,6; pectoral rays 16 or 17; vertical scale rows 35 to 39 (21 speci- mens) ; scale rows above lateral line 3, below lateral line 6 or 7, around caudal peduncle 15 or 16.

Body relatively elongate and slender, body depth 4.5 to 5.2; head length 3.5 to 3.9; length of caudal peduncle 3.5 to 3.9; length of longest pectoral ray 5.8 to 6.5; all in standard length. Eye, 3.5 to 4.2; length of upper jaw 3.4 to 3.7; depth of caudal peduncle 3.0 to 3.2; length of snout 2.6 to 3.0; interorbital width 3.6 to 3.9; length of barbel 1.8 to 2.1; all in length of head. Eye in snout 1.1 to 1.3; interorbital width in eye 1.0 to 1.2.

Gill rakers 7 or 8+1+18 to 21, total 27 to 30 (25 specimens) ; longest raker about 1.7 to 2.0 in longest filament. Second dorsal spine flexible near tip; barbels short almost reach vertical through posterior margin of eye.

Color in aleohol—Head and body dusky to dark brown above and silvery below, lacking any conspicuous color or color mark, some specimens more uniform light brown or tan; peritoneum black; fins transparent, caudal fin and to a lesser extent soft dorsal more con- spicuous in having dusky striations in most specimens.

Color when alive —Schultz recalls that many of the above specimens, especially those collected at night, had a reddish brown coloration when taken from the water. None showed any evidence of stripes or spots as is found on samoensis or auriflamma.

Range.—This species has been reported from the East Indies, the Philippines, and various islands of Oceania. The references are listed by Fowler (Mem. Bishop Mus., vol. 10, p. 234, 1928) and Herre and Montalban (Philippine Journ. Sci., vol. 36, No. 1, p. 135, 1928). I suspect that more than one species was involved in these reports and one should be cautious in the application of these data.

Remarks.—The following arguments were considered in the use of the name vanicolensis:

1. It seemed most logical to apply some available name in order that the specimens may be referred to and found by future workers, although I am not confident of the accuracy of this application. Of

FAMILY MULLIDAE—LACHNER 45

the names available, the description of Upeneus vanicolensis Cuvier and Valenciennes best fits my specimens. The type locality is Vanicolo Island, Santa Cruz Islands, a locality not faunistically different from the Marshall Islands in respect to the Mullidae. Characters listed by subsequent authors for this species agree fairly closely with those of mine, although I have no positive assurance of this, since the material studied by them was often not listed nor was it available for examination.

2. It is conceivable that the material may represent the young or juvenile form of some well known species. All were of a juvenile size, 77 to 96 mm. in standard length. On the basis of the material collected, the species was most active at night, for 73 specimens were taken in 11 collections at night with the aid of lights and only 4 collections totaling 21 specimens were taken during the daylight hours when most of the collecting was done. Specimens of M. samoensis and M. auriflamma were taken with vanicolensis in the same net hauls. The greater number of gill rakers, larger eye, and longer barbels of auriflamma and the light peritoneum of pjflugeri eliminate any possibility of associating these species with my speci- mens of vanicolensis.

On the basis of certain meristic counts and measurements, samoensis is the species most closely related (see table 80) to specimens of vanicolensis; however, my specimens have these pronounced differences from samoensis: Most specimens were reddish brown in life and in preservation the striations of the caudal fin are dusky, and no specimen had a dark spot or yellow stripe on the body as in samoensis; at com- parable sizes the body is slenderer, the snout more pointed, and the mouth more horizontal; certain morphological structures, such as the shorter head, wider interorbital, and shorter snout in vanicolensts, show differences that do not appear to be associated with young or with a transforming stage. The barbel is significantly shorter, although this difference might be associated with transformation, as the young may be pelagic and may not require the barbels as do the bottom-feeding adults.

In several species of Parupeneus, the adults of which equal or exceed in total length that attained by any species of Mulloidichthys, the young have acquired the definitive adult characters at a much smaller size, 40 to 50 mm., than my specimens of vanicolensis. The absence of intermediate specimens lead me to drop the theory that my specimens represent an early stage. Life history studies are needed.

3. The last possibility is that these specimens represent a new species. In view of the fact that we do not have a clear understanding just what form Upeneus vanicolensis Cuvier and Valenciennes may

46 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

represent, and also that we know so very little of the life history, habits, and life coloration of most of the species of Mullidae (items quite important to the taxonomic understanding of the family), I hesitated to proceed with the viewpoint that these specimens are new.

Family POMACENTRIDAE: Damselfishes

By Loren P. Woops and Lreonarp P. Scuuttz

No attempt was made to revise this large complicated family consist- ing of very numerous species, some of which are closely related, of small size, and generally difficult to identify. This latter is especially true if few species are available for comparison.

The family is characterized by having II anal spines, X to XIV dorsal spines; teeth small, conical to compressed, mostly adapted to feeding on algae; a single nostril each side of snout.

Since the pelvic rays normally are I,5 and branched caudal fin rays 7 or 8+6 or 7, usually 7+6, these are omitted in descriptions.

KEY TO THE GENERA OF POMACENTRIDAE FROM THE NORTHERN MARSHALL AND MARIANAS ISLANDS la. Transverse scale rows more than 50; suborbital and preopercle margins serrate; opercle, subopercle, and interopercle with radiating striae and strong serrations; snout naked; teeth of jaws in single row; dorsal spines X, Tare ly Me to hae ea ede aS ed Amphiprion Bloch and Schneider 1b. Transverse scale rows less than 40; dorsal spines XII to XIV. 2a. Teeth of jaws conical, biserial, outer row larger, more widely spaced than teeth of inner row, which are often villiform, usually forming a. band near front of jaws; 2 or 3 short free spines at base of caudal fin above and below. 3a. Suborbital margin, preopercle, and subopercle serrate; body ovate, its greatest depth 1.40 to 1.55 in standard length; angle of upper profile with lengthwise axis of body 60 to 80 degrees___-_Dascyllus Cuvier 3b. Suborbital margin entire when free, usually poorly defined and sometimes not evident; subopercle entire, preopercle entire, or crenulate, some- times finely serrate; body elliptical, its greatest depth 1.70 to 2.0; angle of anterior upper profile with lengthwide axis of body 40 to 50 CLE PTOCS re aoe Rated ep 2 fa ome aire nape gee een ae Meee CUS Ena) Chromis Cuvier 2b. Teeth of jaws not conical but compressed at tips, incisiform (at least at front of jaws) or long and slender with tips pointed or rounded, in one or two rows but second row close against outer and not forming a band; base of caudal without any free spines. 4a. Preopercular margin serrate; suborbital serrate or entire. Pomacentrus Lacepéde 4b. Preopercular margin entire, suborbital margin always entire. | Abudefduf Forsk&l

Genus AMPHIPRION Bloch and Schneider

Amphiprion BLocu and ScHNEIDER, Systema ichthyologiae. . . , p. 200, 1801 (type species, Lutjanus ephippium Bloch).—Schultz, Proc. U. S. Nat. Mus., vol. 103, pp. 187-201, pls. 9 and 10, 1953.

47

48 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

Prochilus (on Klein, 1775) BuerKer, Nat. Verh. Holl. Maatsch., No. 6, ser. 3, vol. 2, p. 20, 1877 (type species, Lutjanus ephippium Bloch).

Actinicola Fow.er, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 12, p. 533, 1904 (type species, Lutjanus percula Lacepéde).

Phalerebus WuittEY, Mem. Queensland Mus., vol. 9, pt. 3, p. 216, 1929 (type species, Prochilus akallopisos Bleeker).

Paramphiprion Wane, Contrib. Biol. Lab. Sci. Soe. China, vol. 15, zool. ser., No. 6, p. 89, 1941 (type species, Paramphiprion hainanensis Wang = A. polymnus (Linnaeus)).

Descriptions and analyses of species referable to the genus Amphi- prion have been based on so few specimens, usually only one or two, that the problem of variability or constancy of the color pattern has been neglected. For most of the few hundred species, among more than 50 families studied in detail by Leonard P. Schultz, the basic color pattern has been observed to be fairly constant. It is of utmost importance in recognizing species, especially in the genus Amphiprion. Weber and de Beaufort (Fishes of the Indo-Australian Archipelago, vol. 8, pp. 330-348, 1940) recognized 8 species, whereas we have dis- tinguished 15 species in the tropical Indo-Pacific, and there may be others recognizable when larger series are compared and additional characters studied.

During December 1953 Schultz had the good fortune to be able to study specimens of Amphiprion in the British Museum. As the result of these studies certain nomenclatorial changes are necessary in regard to his (1953) revision of this genus. Especially important was the discovery that his new A. mauritiensis is a synonym of A. fusciventer Bennett.

Fin ray counts were made on various species, including those ex- amined by Schultz in the British Museum, are recorded in table 84.

Except for original descriptions no attempt was made to include ail references to species referable to the genus Amphiprion. Whenever figures of species were found it has been possible to include these species in the synonymy, but most species descriptions are not in sui- ficient detail for them to be assigned without an examination of the specimen on which the records were based. The members of this genus normally live a commensal life in sea anemone.

In the revision by Schultz (1953) of Amphiprion, A. letezonatus Waite (pl. 79,A) was omitted. This valid species has been added to the following key after Schultz (1953, p. 189) modified in sections 6a and 6b.

KEY TO THE SPECIES OF AMPHIPRION

la. A white band (sometimes indistinct) along middorsal line from snout to dorsal origin or beyond along base of dorsal fin; total pectoral rays 17 to

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 49

19; next to last dorsal spine about 1.3 to 1.5 in longest dorsal spine; no notable emargination in dorsal fin; scales on dorsal surface of head extend forward to a line between front of orbits.

2a. No vertical pale bars; dorsal rays about X,18 or 19; anal about II,12 or 13;

DECLORAL Tl OTA) ery aes eae ee ee ee A. akallopisos"® Bleeker

2b. A single vertical pale bar about 2 scales wide from nape to subopercle; dorsal rays about X,16; anal about II,12 or 13; pectoral 17.

A. perideraion Bleeker

1b. No white band along middorsal line.

3a. Caudal fin with pale or dusky roundish center posteriorly edged with black; outer edges of caudal fin white; second pale or white bar from rear of spiny dorsal fin to anus with a triangular anterior projection under depressed pectoral fin; tips of pelvics black; a broad white color bar on head and on caudal peduncle always present. Dorsal fin deeply indented at rear of spiny part, next to last dorsal spine contained three times in longest dorsal spine; scales on dorsal surface of head do not extend forward of nape; plate 81,D. A. percula ” (Lacepéde)

3b. Color not as in A. percula.

4a. Central part of caudal fin black; outer edges of caudal fin broadly or narrowly edged with white; second pale bar, if present, without any projection anteriorly; pale bar on head present; next to last dorsal spine contained 1.2 to 2.0 times in longest dorsal spine; no notable emargination in dorsal fin.

5a. Second pale bar represented dorsally on body by an ovate white area that continues anterodistally on soft dorsal fin; this white area does not extend below midlengthwise axis of body and never to anus; first pale bar 10 to 12 scales wide; anal fin black, except distally edged with white; spiny dorsal black; pelvics black; pectoral pale, except basally it is blackish; next to last dorsal spine contained about 1.2 to 1.5 in longest dorsal spine;

18 4mphiprion akallopisos Bleeker, Nat. Tijdschr. Nederl.-Indié, vol. 4, p. 281, 1853 (type locality, Sumatra, type examined BM1862.2.28.68).—Schultz, Proce. U. S. Nat. Mus., vol. 103, pp. 190-191, pl. 9,A, 1953. Prochilus akallopisus Bleeker, Atlas ichthyologique . . ., pl. 400, fig. 1, 1878.

19 Lutjanus percula Lacepéde, Histoire naturelle des poissons, vol. 4, pp. 194, 239, 240, 1802 (type locality, New Britain).

Amphiprion tunicatus Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 399, pl. 182, fig.

2, 1830 (type locality, Vanicolo).—Lesson, Voyage . . . la Coguwille . . . , zoologie, vol. 2, pt. 1, p. 192, pl. 25, fig. 3, 1830 (Port Praslin, New Ireland; Doreh, New Guinea). Prochilus percula Bleeker, Atlas ichthyologique . .. , vol. 9, pl. 400, fig. 2, 1878.

Anthias polymna var. (non Linnaeus) Bloch, Naturgeschichte der ausliindischen Fische, vol. 6, p. 103, pl. 316, fig. 3, 1792.

Amphiprion percula Giinther, Journ. Mus. Godeftroy, vol. 15 (Fische der Stidsee, pt.7), pl. 124, fig. A, 1881 (Samoan Islands).—Day, Fishes of India, vol. 2, p. 379, pl. 80, fig. 4, 1878 (Andamans).—Montalban, Bur. Sci. Manila Monog. 24, p. 14, pl. 2, fig. 2, 1928 (Philippine Islands).—Schultz, Proc. U. 8. Nat. Mus., vol. 108, p. 191, pl. 9,C, 1953.

Actinicola percula Aoyagi, Coral fishes, Tokyo, pl. 37, fig. 2, 1943 (Kakure-Kumanomi); Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 175, pl. 9, fig. 2, 1941 (Japan).

Amphiprion bicolor Castelnau, Proc. Zool. Acclim. Soe. Victoria, p. 92, 1873 (Port Darwin).

Actinicola bicolor (Castelnau) Whitley, Mem. Queensland Mus., vol. 9, pt. 3, p. 215, pl. 27, fig. 2, 1929 (Port Darwin).

50 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

no notable emargination in dorsal fin; scales on dorsal surface of head extend forward to a line between rear of orbits. A. laticlavius 2° Cuvier and Valenciennes 5b. Second pale bar continuous from dorasl part of body to region of anus.

6a. (see also 6b and 6c) Second pale bar very broad about 16 to 18 scales wide at level of lateral line (its width there greater than width of second black bar at level of lateral line) from whence it continues ventrally becoming much broader, including the region from just behind the pelvic bases to the beginning of the last third of the base of anal fin; white bar across caudal peduncle present; basal 34 of caudal fin blackish, distal }4 whitish. A. latezonatus 2! Waite 6b. Second pale color bar broad, about 7 to 14 scales wide at level of lateral line (its width there contained about 2 times in width of second black bar at level of lateral line) from whence it continues posterodorsally on soft dorsal fin, also ventrally it is about same width as at lateral line; pelvies dusky to blackish; spiny dorsal blackish; anal blackish at least basally; scales on dorsal surface of head extend forward to a line between rear of orbits. 7a. Caudal peduncle with broad white bar; pectoral fin pale distally, basally dusky_-_-_---- A. chrysogaster Cuvier and Valenciennes 7b. Caudal peduncle black; no white bar; black coloration of posterior part of body continuous on central part of caudal fin; pectoral finiduskyseou0 oe fee oe Sees A. polymnus ” (Linnaeus) 6c. Second pale bar narrow, about 2 to 6 scales wide at level of lateral line, its width there contained 5 or more times in width of second black bar at level of lateral line; caudal fin black, narrowly edged with white; anal fin pale to blackish; scales on dorsal surface of head extend forward to over rear half of pupil; pelvics and pec- torals pale; soft dorsal black, edged with white. 8a. Second and third (peduncular) white bars about 2 or 3 scales wide, their width at level of lateral line contained 7 to 10 times in width of third black bar at level of lateral line; second white bar not extending to distal edge of dorsal fin, ending on basal half of last dorsal spine; posterodorsal part of body blackish; anterior and ventral parts of body pale; spiny dorsal light dusky. A. tricinctus Schultz and Welander

20 Amphiprion laticlavius Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 394, pl. 132, fig. 1, 1830 (type locality, New Guinea).—Schultz, Proc. U. 8. Nat. Mus., vol. 103, p. 193, pl. 9,D, 1953. Amphiprion bifasciatus (non Bloch) Montalban, Bur. Sci. Manila Monog. 24, p. 15, pl. 3, fig. 1, 1928 (Philip- pine Islands).

Amphiprion bifasciatus annamensis Chevy, Travaux Inst. Oceanogr. Indochine, Mem, 4, pt. 1, Poissons, p. 99, pl. 39, 1932 (type locality, Sud-Annam).

Prochilus bifasciatus (non Bloch) Bleeker, Atlas ichthyologique . . . , pl. 400, fig. 4, 1878.

Amphiprion polymnus (non Linnaeus) Aoyagi, Coral fishes, Tokyo, pl. 36, fig. 2, 1943 (Toaki-Kumanomi); Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 178, pl. 12, fig. 4, 1941 (Japan).

Amphiprion unimaculatus (non Meuschen) Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soe. Japan, vol. 3, No. 2, p. 202, fig. 28, 1939 (Itoman, Okinawa).

The following specimens were examined in the British Museum: BM1860.3.14.884, 2 specimens from India; BM1933.3.11.426 or 427, 1 specimen from the Philippines.

21 Amphiprion latezonatus Waite, Rec. Australian Mus., vol. 3, No. 7, p. 201, pl. 34, 1900 (type locality, Lord Howe Island).

22 Perca polymna Linnaeus, Systema naturae, ed. 10, p. 291, 1758 (type locality, ‘‘Indees’’).

Anthias bifasciatus Bloch, Naturgeschichte auslindischen Fische, vol. 6, p. 103, pl. 316, fig. 2, 1792.

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 51

8b. Second and third (peduncular) white bars 4 to 6 scales wide, their

widths at level of lateral line contained from 4 to 6.5 times in

width of third black bar at level of lateral line; second white

bar extends into distal half of dorsal but not quite to the edge

of that fin; dorsal and posterior half of body blackish; antero-

ventral part of body pale; spiny dorsal dusky to blackish; plate

OAT RRR OIE eRe) 2 CARNE TE ee ane eee A. fusciventer # Bennett

4b. Caudal fin plain pale or plain dusky, no black central blotch edged with white posteriorly; first pale bar on head present.

9a. Second pale bar broad, about 7 to 13 scales wide at level of lateral line,

and continuing to distal edge of spiny dorsal fin, thence posteriorly

along distal edge of soft dorsal; caudal fin pale; caudal peduncle

black, without the third pale bar, at least on adults; spiny dorsal,

anal, and pelvic fins black; pectoral fin pale; next to last dorsal

spine contained about 1.5 in longest dorsal spine; scales on dorsal

surface of head extend forward to a line between rear of orbits.

A. sebae Bleeker

9b. Second pale bar, if present, not continuing to distal edge of dorsal fin;

next to last dorsal spine contained 0.8 to 1.2 in longest dorsal

spine; scales on dorsal surface of head extend forward to lines between rear edge of orbits to center of pupil. 10a. First two pale bars typically present on adults.

lla. First two pale bars broad, each about 5 to 7 scales wide at level

of lateral line; third (peduncular) pale bar represented by a

white bar, posteriorly edged with darkish, or the dark body

color on anterior part of caudal peduncle may end abruptly,

the white continuing on caudal fin; spiny dorsal black,

dusky, or pale; soft dorsal and anal fins pale to black; pelvics

pale to black, sometimes edged with black anteriorly;

pectoral pale; next to last dorsal spine about 1.2 in longest

Prochilus bifasciatus Bleeker, Atlas ichthyologique . . ., vol. 9, pl. 400, fig. 5, 1878.

? Amphiprion ocellaris Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 399, 1830 (type locality, Sumatra).

? Amphiprion melanurus Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 400, 1830 (type locality, Sumatra).

Lutjanus jourdin Lacepéde, Histoire naturelle des poissons, vol. 4, p. 191, 235, 1802 (type locality ,Amboina).

Coracinus seu Sciaena wnimaculata Meuschen, Index Zoophyl. Gronow, Pisces, No. 227, 1781 (based on Gronow 1763).

Coracinus vittatus Gray, Catalogue of Fish collected and described by L. T. Gronow, p. 57, 1854 (on Gronow No. 227).

Amphiprion intermedius Schlegel and Muller, Verh. Nat. Gesch. Ned. Overz. Bezitt. Zool., p. 18, 1839-1841

(ref. copied).

Paramphiprion hainanensis Wang, Contr. Biol. Lab. Sci. Soe. China, zool. ser., vol. 15, No. 6, p. 89, 1941 (type locality, Hainan Island, China).

Amphiprion polymnus Schultz, Proc. U. S. Nat. Mus., vol. 103, p. 194, pl. 9,J, 1953.

Amphiprion polymnus (Linnaeus) has been confused almost since the day it wasnamed. Weber and de Beaufort (Fishes of the Indo-Australian Archipelago, vol. 8, p. 344, 1940) discuss the confusion between polymnus and bicinctus, the former name having been used for the species currently called bicinctus. The follow- ing were examined in the British Museum: BM1852.4.21.144-144, from Amboina; 1871.7.20.121 from Manado.

23 Amphiprion fusciventer Bennett, Proc. Comm. Zool. Soc. London, vol. 1, p. 165, 1831 (type locality, Mauritius; holotype BM1856.2.15.10).

Amphiprion mauritiensis Schultz, Proc. U. S. Nat. Mus., vol. 103, p. 196, pl. 9,1, 1953 (type locality, Mauri- tius).

52 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

dorsal spine___--_- A. xanthurus * Cuvier and Valenciennes 11b. First two pale bars narrow, second usually narrower than first, the latter 3 to 4.5 scales wide and the first 3 to 6 scales wide at level of lateral line; caudal peduncle blackish, without third pale bar on adults, sometimes pale bar is present on young; dark color of caudal peduncle gradually fading into pale color of caudal fin; spiny dorsal pale to dusky, soft dorsal pale; anal fin pale; pectoral pale; pelvics pale, except anterior edge black; second from last dorsal spine contained from 1.0 to 1.2 times in longest dorsal spine. A. bicinctus Riippell 10b. First pale bar usually present on head, about 4 to 6 scales wide at level of lateral line; second pale bar lacking, except sometimes on small young specimens. 12a. Anal fin pale or partly pale; pelvics with some pale area. 13a. Anal fin pale, except distally edged with fine black line; pelvics pale, except anterior edge black; breast pale; dorsal spines X, soft dorsal rays usually 17; plate 80,A,C. A. ephippium ” (Bloch) 13b. Anal fin pale or distally pale with basal half (or less) dusky to blackish; pelvies pale distally and blackish ventrally, or inner rays pale and outer rays broadly blackish, except distal tips, which are pale; dorsal spines IX or X (more often IX than X), soft dorsal rays usually 17 or 18. A. frenatus 2° Brevoort 12b. Anal and pelvic fins black, no pale areas anywhere; dorsal spines X, soft dorsal rays usually 17. A. melanopus Bleeker

#4 Amphiprion ranthurus Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 402, 1830

(type locality, Ile de France).—Schultz, Proc. U. S. Nat. Mus., vol. 103, p. 198, pl. 9,G, 1953.

Amphiprion clarckii Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 9, p. 504, 1833 (amended spelling on Bennett).

Anthias clarkii Bennett, Fishes found upon the Coast of Ceylon. Ed. 2, p. 29, pl. 29, 1834 (type locality Ceylon).

Sparus milii Bory de St. Vincent, Dictionnaire classique d’histoire naturelle, vol. 17, p. 130, pl. 113, fig. 2, 1831 (type locality, China Sea).

Prochilus polymnus (non Linnaeus) Bleeker, Atlas ichthyologique . . ., vol. 9, pl. 400, figs. 7, 8, 1878.

Anthias polymna (non Linnaeus) Bloch, Naturgeschichte der auslindischen Fische, vol. 9, p. 89, pl. 316, fig. 1, 1792.

Amphiprion bicinctus (non Riippell) Aoyagi, Coral fishes, Tokyo, pl. 37, fig. 1, 1943 (Kumanomi); Biogeog- raphica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 169, pl. 9, fig. 1, 1941 (Japan).

Amphiprion polymnus (non Linnaeus) Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 204, fig. 30, 1939 (Riu Kiu Islands).—Montalban, Bur. Sci. Manila Monog. 24, p. 10, pl. 1, fig. 1, 1928 (Philippine Islands).

Amphiprion chrysopterus Cuvier and Valenciennes, Histoire naturelle des poissons, vol. 5, p. 401, 1830 (no locality given).

Amphiprion japonicus Temminck and Schlegel, Fauna Japonica, Pisces, p. 66, 1843 (type locality, Japan).

Amphiprion chrysargyrus Richardson, Rep. Fifteenth Meet. British Assoc. Adv. Sci., vol. 15, p. 254, 1846 (type locality, Seas of China and Japan).

Amphiprion boholensis Cartier, Verh. Phys.-Med. Ges. Wiirzburg, new ser., vol. 5, p. 96, 1874 (type locality, Bohol).

Amphiprion melanostolus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 390, 1842 (type locality, Depuch Island).

Amphiprion snyderi Ishikawa, Proc. Nat. Hist. Dep. Tokyo Imp. Mus., vol. 1, No. 1, p. 11, pl. 5, 1904 (type locality, Bonin Island).

FAMILY POMACENTRIDAE—-WOODS AND SCHULTZ 53

TaBLe 84.—Fin ray counts recorded for certain species of Amphiprion

Dorsal

Anal Pectoral (total rays)

IX| X | XI| 14] 15 | 16} 17 | 18 | 19 | 2 13 | 14 | 15 | 16 | 17] 18 | 19 | 20} 21

ephippium north Australia, near

(DarwiWes-se 52s |e LOI eeos eel tO Omens eee koalas death nO commas ence Sr LOM eM | eee melanopus

Marshall Is-

} yao pe eee eS TOME eecst wy eee S: eee ete (<3)p| sees WS His eu oe] (a pe Gi biuleses

Samoan, and

Paumoto Is-

jandssOur- . 5832/2 22 Ay ies Se ee Aires EL Sree sk 2 3) eRe [Ese Splaly [ea eelee LOLClOpiUes > eee 3 | Pe fs A | es |e ese 4 1 2 ABP ec seein 1 4 Bhi fonsn| US frenatus

hilippine Is-

Wandss se eene eS Zon elO| aes | eens | eee 2h Oe 2 eG ees TG ence ees |e Sahezenlheei|oses

Okinawa, and

Japanieeeees a2 2h oes ee Es eee seers 1 Te EL es | 2h ee ea ee | ees | | 1 Sole eS ROMO DISOSsea eee = A Aes Ee a eee oles 3 i 4 1 yy) rN Bee te 2) 2 Tee MEridendiOnss 2-2 |= = Ohi] ea pe hy tg eRe te | kn || Cis ea ee a Ges esl | yd fever Pe chrysogaster__-_----|---- Gulinen || eed Dye Db eG: irri [oak Bp. 7 eee see Lee Onl aS Bicimetis a2 Sea Saris | Bees a EA earch [tates | oeesees | en eer Na Oe erin eae Se 1 Mains SAUSCIDETILET wee at tee |e LON heeds 9 Se | ee TN fe ee cx RE, [Es RR LL TAS) potas SAS eS i S| eee 12 GRICTTUCLULS = 32 Soe ee he | es S| ee | See My eee | Se 8) USS Ee ee Bo | eee sees eS LOLEZONOIMIG tooe 2 eee ee 2 eee lee OL ek a ee 2 1 1 eo eo | eee ee CUS ees | eae 2h eee TOTEUT Sos eee |e ALD) eases | aE Sui] Ale Paes | ee. 12/2 Bale DM|| WAR Ses | See Cee eam Mercula: se Ae! ees |Eees Upon eeee Aalto: tooo eer lene alee Olek pa Se - Tig| me Se EAE (EE cea CURE a eee Dil Pes aE 1 A) abe 2) aee 1 Pt ates | Eker ee 2s eee 2

1 Counts recorded by Waite and from a specimen in British Museum,

25 Lutjanus ephippium Bloch, Naturgeschichte der auslindischen fische, vol. 4, p. 121, 1790 (ref. not seen); Ichthyologie, Histoire naturelle des poissons, vol. 7, p. 98, pl. 250, fig. 2, 1792 (East Indies).

Amphiprion ephippium Day, Fishes of India, vol. 2, p. 378, pl. 80, fig. 1, 1878.—Schultz, Proc. U. S. Nat. Mus., vol. 103, pp. 199-200, pl. 10,B,C, 1953.

Amphiprion monofasciatus Thiolliére, in Montrouzier, Essai sur la faune de 1’Ile de Woodlark ou Moiou, Ann. Sci. Physic. Nat. Agri. Indust., vol. 8, p. 476, 1856 (type locality, Woodlark Island).

Amphiprion tricolor Giinther, Catalogue of the fisbes in the British Museum, vol. 4, p. 8, 1862 (type locality, Port Essington, South Australia; types examined, BM1844.1.13, BM1847.7.21.56, 1855.9.19.564-7).

Amphiprion rippelli Castelnau, Proc. Zool. Acclim. Soc. Victoria, vol. 2, p. 91, 1873 (type locality, Port Darwin).

Amphiprion frenatus (non Brevoort) Day, Fishes of India, vol. 2, p. 378, pl. 80, fig. 2, 1878.

Prochilus ephippium Bleeker, Atlas ichthyologique .. . , vol. 9, pl. 401, figs. 1, 9, 1878.

Amphiprion rubrocinctus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 391, 1842 (type locality, Depuch Is- land [probably young]).

28 Amphiprion frenatus Brevoort, Notes on some figures of Japanese fish, in U. S. Japan Exped. Nat. Hist., Washington, p. 263, pl. 6, fig. 4, 1856 (type locality, Lew Chew [Okinawa]).—Jordan and Snyder, Proc. U. S. Nat. Mus., vol. 24, p. 597, 1902.— Montalban, Bur. Sci. Manila Monog. 24, p. 12, pl. 2, fig. 1, 1927 (Philippine Islands).—Aoyagi, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 167, 1941 (Japan).—Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soe. Japan, vol. 3, No. 2, p. 203, fig. 29, 1939 (Riu Kiu Islands).—Schultz, Proc. U. S. Nat. Mus., vol. 103, p. 201, pl. 10,D, 1953.

Prochilus polylepis Bleeker, Versl. Akad. Wet. Amsterdam, ser. 2, vol. 11, p. 185, 1877; Atlas ichthyologique ..., vol. 9, pl. 401, fig. 6, 1878.

Amphiprion polymnus (non Linnaeus) Montalban, Bur. Sci. Manila Monog. 24, p. 10, pl. 1, fig. 1, 1928 (Philippines).

54 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2 AMPHIPRION PERIDERAION Bleeker PuaTE 79,B

Amphiprion perideraion BuerKER, Nat. Tijdschr. Nederl.-Indié, vol. 9, p. 437, 1855 (type locality, Obi Major).—Monvra.pan, Bur. Sci. Manila Monogr. 24, p. 16, pl. 4, fig. 1, 1928 (Philippine Islands).—ScuutrTz, Proc. U. 8. Nat. Mus., vol. 103, p. 191, pl. 9,B, 1953.

Prochilus perideraion BLEEKER, Atlas ichthyologique . . ., pl. 400, fig. 1, 1878.

2?Amphiprion rosenbergi BureKnr, Act. Soc. Sci. Indo-Néerl., vol. 6, p. 16, 1859 (type locality, New Guinea); Atlas ichthyologique . . ., vol. 9, pl. 402, fig. 2, 1877.

Amphiprion nigripes Recan, Trans. Linn. Soc. London, ser. 2, zool., vol. 12, p. 230, pl. 24, fig. 2, 1908 (type locality, Maldives, BM 1908.3.23.101, holotype examined).

SPECIMENS STUDIED

Bikini Atoll: 1 station, 1 specimen, 39.5 mm. in standard length. Rongelap Atoll: 1 station, 1 specimen, 23 mm.

Description.—Dorsal fin rays X,15 or 16; anal II,13; pectorals ii,15; scale rows 50, with 5 or 6 rows between lateral line and base of dorsal, 19 between origin of anal and lateral line; perforated scales in lateral line 35.

Depth of body 2.14; length of head 3.3; both in standard length. Snout 3.43; eye 3.25; preorbital width 7.5; length of upper jaw 2.8; postorbital part of head (hind margin of eye to upper edge of gill opening) 2.8; interorbital width 3.25; depth of caudal peduncle 1.88; length of pectoral fin 1.29; length of pelvic fin 1.41; length of 3rd dorsal spine 2.61; length of middle caudal rays 1.02; all in length of head (tip of snout to tip of longest opercular spine). Angle of snout profile with lengthwise axis of body 61 degrees; snout profile convex.

Body oblong, preorbitals and suborbitals with strong spines, pre- opercle serrate, spines far apart; opercle and subopercle with strong radiating spines. Anterior spines of dorsal distinctly longer than posterior, so dorsal appears notched, soft dorsal and anal low, rounded, caudal rounded.

Color in alcohol.—Head and back light yellowish brown or reddish brown; sides and belly dirty grayish; a vertical chalky white line, narrower than eye but broader than pupil, extending from temporal region across anterior part of opercle, ending at lower margin of subopercle; a dark reddish brown median line from interorbital area to origin of spiny dorsal; all fins pale yellowish.

Remarks.—This species has been reported from the East Indies, Philippine Islands, and Palau Islands but not previously from Micro- nesia. It is best recognized by the narrow white band on the mid- dorsal line of head, beginning between front of eye and extending to dorsal origin, along with a narrow white vertical bar on rear of head behind eye. Four specimens were studied: USNM 141032, 141033

FAMILY POMACENTRIDAE—-WOODS AND SCHULTZ 55

and 147129 from the Marshall Islands and Borneo, and MCZ 33409 from Amboina.

Schultz examined the type of A. nigripes in the British Museum and found the pectoral fin to have 19 rays, which number is 2 more than for western Pacific specimens of perideraion. Thus A. nigripes may be a valid species close to A. perideraion. The type has faded so that now it is light brown with an indistinct pale streak along base of dorsal fin, a broad pale band along midside, and probably one below, from lower edge of pectoral base to ventral edge of caudal peduncle. A series of specimens with distinct color patterns is needed to work out the relationship of these two forms.

AMPHIPRION CHRYSOGASTER Cuvier and Valenciennes PLATE 81,C

Amphiprion chrysogaster CuvieER and VALENCIENNES, Histoire naturelle des poissons, vol. 5, p. 400, 1830 (type locality, Ile de France).—Lrsson, Voyage ... 1a Coquille . . ., zoologie, vol. 2, pt. 1, p. 191, pl. 28, fig. 3, 1830 (Ile de France).—Scuvuutz, Proc. U. S. Nat. Mus., vol. 103, p. 193, pl. 9,E, 1953.

Amphiprion percula (non Lacepéde), Okapa and IkEpa, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, pl. 6, fig. 1, 1939 (Riu Kiu Islands).

Amphiprion trifasciatus Cuvier and VALENCIENNES, Histoire naturelle des poissons, vol. 5, p. 595, 1830 (type locality, Moluccas).

Prochilus bifasciatus (non Bloch) BurEKer, Atlas ichthyologique .. ., vol. 9, pl. 400, fig. 6, 1878.

SPECIMENS STUDIED

Rongelap Atoll: 1 station, 1 specimen, 21.7 mm. in standard length.

Description.—Dorsal rays X,16; anal 11,14; pectorals 11,18; scale rows from upper edge of gill opening to base of middle caudal rays 49, with 5 rows between lateral line and base of dorsal, 19 between lateral line and origin of anal.

Depth of body 2.0; length of head 3.0; both in standard length. Snout 4.24; eye 2.25; preorbital width 12.0; length of upper jaw 2.76; postorbital part of head 2.76; interorbital width 3.3; depth of caudal peduncle 2.05; length of pectoral fin 1.0; length of pelvics 0.76, length of third dorsal spine 1.8; length of middle caudal rays 0.9; all in length of head (tip of snout to tip of longest opercular spine). Depth of caudal peduncle in its length 1.6; angle of snout profile with length- wise axis of body 45 degrees; snout profile convex.

Body oblong, suborbital entire, preopercle serrate on lower half of posterior margin, opercle with two diverging groups of spines, subo- percle and posterior part of interopercle with spines; spiny dorsal fin with 9th spine shortest, so fin is deeply notched though spiny dorsal is connected with soft dorsal; soft dorsal, anal, and caudal rounded; pelvics long, one reaching to and one past origin of anal (perhaps a juvenile characteristic).

42214260 —_5

56 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

Color in alcohol.—Snout and chin pale yellowish white, body pale grayish brown; head and body with three black-bordered white transverse bars as wide as or wider than diameter of eye, first on pos- terior part of head running from nape across opercles just behind eye, second from the 9th dorsal spine and third soft dorsal ray to anus and origin of anal fin, and third, black-bordered on its anterior margin only, across the posterior % of caudal peduncle; second bar straight on its anterior margin, without forward projecting loop or point; spiny dorsal dusky brown except for last 2 spines, which are white; soft dorsal white on first 2 or 3 rays, rest of fin dusky brown; caudal fin white basally and distally, also on outer rays, rest of fin dusky brown; anal dusky blackish; one of pelvic fins pale, the other faintly dusky on tip; pectoral pale.

Remarks.—This species, with three broad white bars, has the cen- tral part of the caudal fin black and the outer edges white; the second white bar is continuous on the distal part of the soft dorsal fin.

We have examined (in USNM) 1 specimen each in lots from Mauri- tius (USNM 61690), the Marshall Islands (USNM 141034), and the Philippines (USNM 147127); (in MCZ) 3 specimens from Zanzibar, Africa, and also 1 from Apiang, Kingsmill Island (Marshalls), col- lected by Andrew Garrett; and (in BM), bearing the name A. inter- medius, 1 specimen each from the following numbers: BM 1852.11.4. 12, Molluccas; BM 1938.12.6.12, Mombassa; and BM 1862.2.25.1.9, from Dr. Bleeker’s collection.

AMPHIPRION TRICINCTUS Schultz and Welander

PuaTE 79,E

Amphiprion tricinctus Scnuttz and WELANDER in Schultz, Proc. U.S. Nat. Mus., vol. 103, p. 195-196, pl. 9,H, 1953 (type locality, Bikini Atoll). Amphiprion ephippium (non Bloch) var. chrysopierus (non Cuvier and Valen- ciennes) Ginruer, Journ. Mus. Godeffroy, vol. 15 (Fische der Sudsee, pt. 7), pp. 224-225, pl. 122, fig. C, 1881 (Kingsmill Island). SPECIMENS STUDIED Holotype USNM 152929, Bikini Atoll, Amen Island, lagoon, August 21, 1947, Univ. Washington, Staff of Applied Fisheries Laboratory, standard length 75 mm.

Description —Dorsal fin rays X,17; anal IJ,14; pectoral i1,17,i— ii,16,ii; vertical scale rows from upper edge of gill opening to base of caudal fin 54, with 5 scales between lateral line and base of first soft dorsal ray, 20 between lateral line and anal origin; 36 pores in lateral line; predorsal scales 19 or 20; gill rakers 5--1+13.

Detailed measurements were made on the holotype and these data are expressed in thousandths of the standard length, 75 mm.: Greatest depth 560; length of head 272; snout 99; eye 95; preorbital width 35;

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 57

length from snout tip to rear edge of maxillary 101; postorbital length of head 147; interorbital width 100; depth of caudal peduncle 160; length of caudal peduncle from base of last anal ray to midbase of caudal fin 192; length of longest ray of pectoral 287, pelvic 313, upper caudal fin 307, lower lobe of caudal fin 300, spiny dorsal 160; length of next to last dorsal spine 160; width of white part of first pale bar, at level of lateral line, 73, of second 40, of last, or peduncular, 20.

Depth of body 1.7; head 3.4; both in standard length. Snout 3.2; eye 3.1; preorbital width 8.0; upper jaw 2.8; postorbital part of head 2.0; interorbital width 3.0; depth of caudal peduncle 1.8. Length of pectoral fin 1.0; pelvic 0.9; second dorsal spine 23; upper caudal rays 0.9 to 1.0; all in length of head. Depth of caudal peduncle in its length 1.2. Angle of upper profile of head with lengthwise axis of body about 50 degrees; profile of head convex.

Teeth in both jaws in a single row, nearly conical, a little compressed forward, pointed; interorbital space scaled forward to a line connect- ing between middle of pupils; 4 or 5 rows of scales on cheeks; gill cover with a few scales; scales occur part way out on all median fins; preorbital with 3 spines, and suborbital with 10 to 12 smaller spines.

Color in alcohol.—Background coloration of body from about fifth dorsal spine posteriorly and dorsally to midlengthwise axis of body blackish, the anteroventral part of body pale light brown, becoming paler ventrally; spiny dorsal dark brown, soft dorsal black, caudal fin black, except edged with white posteriorly; pectoral and pelvic fins pale or very light tan, anal pale or light tan and distally edged with a black line; first white bar from a little in front of dorsal origin just behind eye, ending on lower edge of subopercle; second white bar begins on last dorsal spine and base of first soft ray in lower third of fin, thence extends ventrally, meeting its fellow in narrow space between anal origin and anus; third white bar, about half width of second, or narrower than width of pupil, crosses caudal peduncle just in front of caudal fin base.

Remarks.—This species may be recognized by the narrowness of the three white bars, especially the second and third, the black caudal fin narrowly edged with white posteriorly.

Named tricinctus in reference to the three white bars.

AMPHIPRION SEBAE Bleeker PLATE 81,B

Amphiprion sebae BieEKER, Nat. Tijdsch. Nederl.-Indié, vol. 4, p. 478, 1853; (type locality, Batavia); Atlas ichthyologique . . ., vol. 9, pl. 400, fig. 9, 1878.—Day, Fishes of India, vol. 2, p. 378, pl. 80, fig. 3, 1878 (Andamans).— Oxapa and IKepa, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, fig. 27, 1939 (Isigaki and Irimote Islands).—Scuuutz, Proc. U.S. Nat. Mus. vol. 103, p. 197, pl. 9, F, 1953.

58 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

Amphiprion papuensis MAacuBAyY, Proc. Linnean See. New South Wales, vol. 8, no. 2, p. 271, 1883 (type locality, New Guinea).—Wu1tLEy, Mem. Queens- land Mus., vol. 9, pt. 3, p. 210, pl. 27, fig. 1, 1929 (on holotype).

SPECIMENS STUDIED Arno Atoll: 1 lot, 1 specimen, 16 mm. in standard length.

Description.—See table 84 for fin ray counts.

Color in alcohol_—This species is close to A. zanthurus but differs in having the second white bar continue to the distal edge of the dorsal fin, whereas in A. zanthurus the second white bar is interrupted by a blackish or brownish distal edge on the dorsal fin.

Remarks. As a result of his study of additional specimens of Amphiprion, and his study of the figure of A. papuensis by Whitley (1929), Schultz removes that species from the synonymy of A. bicinctus and now places it as a synonym of A. sebae.

In addition to those listed in his review the following were examined in the British Museum: BM 1844.2.21.54, Sumatra; BM 1887.11.11.265, Muscat, Arabia; BM 1888.11.6.34-35, East Coast of Madras; BM 1901.12.31.80, Maldives.

AMPHIPRION BICINCTUS Riippell

PiatTE 79,C

Amphiprion bicinctus Rtprrny, Atlas zu der Reise im nérdlichen Afrika . . Fische des Rothen Meers, p. 189, pl. 35, fig. 1, 1828 (type locality, Red Sea; 3 types examined BM 1845.10.29.89; and BM 1860.11.9.78-9).— Scuuttz, Proc. U. 8. Nat. Mus., vol. 103, p. 199, pl. 10,A, 1953.

Amphiprion arion De Vis, Proe. Linnean Soc. New South Wales, vol. 8, p. 450, 1884 (type locality, South Seas).

SPECIMENS STUDIED

Bikini Atoll: 4 stations, 4 specimens, 25 to 99 mm. in standard length.

Description.—Dorsal fin rays X,16 or 17; anal IJ,14; pectoral 11,16 to 18; scale rows from upper edge of gill opening to base of middle caudal rays 52 to 55, with 6 or 7 between dorsal base and lateral line, 17 to 23 between lateral line and origin of anal; gill rakers on first arch, 7+1+13, total 21.

Depth of body 1.83 to 1.94; head 2.84 to 3.36; both in standard length. Snout 3.18 to 3.6; eye 2.81 to 3.43; preorbital width 6.57 to 10; length of upper jaw 2.68 to 3.21; postorbital part of head 2.09 to 2.88; interorbital width 2.58 to 3.21; depth of caudal peduncle 1.8 to 2.0; length of pectoral fin 1.1 to 1.17, of pelvic fin 1.1 to 1.8, of third dorsal spine 1.97 to 2.42, of upper caudal rays 0.78 to 0.96; all in length of head (tip of snout to tip of longest opercular spines).

FAMILY POMACENTRIDAE—-WOODS AND SCHULTZ 59

Depth of caudal peduncle in its length 1.02 to 1.16; angle of snout profile with lengthwise axis of body 62 to 72 degrees.

Body deepest between origin of dorsal and insertion of pelvics; preorbital with 1 or 2 large spines, suborbitals serrate, preopercle entire on 99-mm. specimen, a few teeth wide apart on one 78 mm., and serrate on smallest (27 mm.); opercle, subopercle, and interopercle with long spines, free only near tips; dorsal spines subequal, the last not much shorter than second and third; soft dorsal and anal rounded; caudal with rays of upper angle longest pointed, caudal slightly forked, lower angle rounded.

Color in alcohol.—Lips, chin, lower part of cheeks, and breast pale whitish or yellow; interorbital light brown; body to middle of caudal peduncle dark brown; 2 bluish white, black-bordered transverse bands, the first from nape (where it may or may not join its fellow) ventrally across posterior margin of preopercle and anterior half of opercle to posterior tip of interopercle, this band as broad as diameter of eye at its broadest point, the second band only 3 scale rows in width, extending from between bases of 9th and 10th dorsal spines to about 2 scale rows anterior to anus, but not quite reaching anus, narrowing to a point ventrally; in 27-mm. specimen this band is narrow at base of 10th dorsal spine, where it commences, broadest on lateral line thence narrowing ventrally (it is also somewhat curved forward in the middle); caudal peduncle on anterior half light brown color gradually becoming lighter on posterior half; spiny dorsal dusky brown with narrow dark brown margin; soft dorsal dusky brown on basal %, pale on distal %; anal pale yellowish with narrow dark brown margin (in 27-mm. specimen anal entirely dusky brown); caudal dusky yellowish in large specimens, pale in small; pelvics yellowish or whitish in two large specimens, dusky brown in 27-mm. one; pectoral pale yellowish.

Remarks.—This species has the first two white bars but lacks the peduncular one; the anterior edges of the pelvics are black and the caudal fin is pale.

Six lots, containing 7 specimens, were studied: USNM 61679 from Suva, Fiji; 141030 and 141031 from the Marshall Islands; 3 uncata- loged specimens, 1 from the Paumotus and 2 without data; also one from Bikini in the University of Washington collection.

Among these specimens, 4 adults, 82 to 99 mm. in standard length, have a pale anal fin, whereas 2, 26 and 27 mm. long, have a black anal and one, 25 mm. long, has a dusky anal fin. The caudal peduncle varies from pale dusky to black or brownish. The key gives the essential color pattern of this species.

60 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

AMPHIPRION MELANOPUS Bleeker PuiatE 80,D

Amphiprion melanopus BLEEKER, Nat. Tijdschr. Nederl.-Indié, vol. 3, p. 561, 1852 (type locality, Amboina).—Scuuurz, Proc. U. 8. Nat. Mus., vol. 103, p. 201, pl. 10,E, 1953. Prochilus melanopus BuEEKeER, Atlas ichthyologique .. ., pl. 401, fig. 7, 1878. Prochilus macrostomus BLEEKER, Atlas ichthyologique .. ., pl. 401, fig. 5, 1878. Amphiprion ephippium (non Bloch) Ginruur, Journ. Mus. Godeffroy, vol. 15 (Fische der Siidsee, pt. 7), pl. 122, fig. D (var. melanopus on p. 225), 1881. Amphiprion mccullochi Wurrtey, Mem. Queensland Mus., vol. 9, pt. 3, p. 213, 1929 (type locality, Lord Howe Island). Amphiprion macrostoma (non Bleeker) Curvy, Travaux Inst. Oceanogr. Indo- chine, Mem. 4, pt. 1, Poissons, p. 102, pl. 40, 1932 (Annam). SPECIMENS STUDIED Bikini Atoll: 3 stations, 8 specimens, 36 to 67 mm. in standard length. Rongerik Atoll: 3 stations, 10 specimens, 30 to 72 mm.

Description.—Dorsal fin rays X,16 or 17 (rarely XI,15); anal II,13 or 14; pectoral 11,17 (rarely 11,18); scale rows from upper edge of gill opening to base of caudal rays 47 to 54 (usually 49 to 51), from base of dorsal to lateral line 4% plus 2 on sheath, from lateral line to origin of anal 20 to 22; gill rakers 5+1-+138.

Depth of body 1.7 to 1.9; head 3.0 to 3.2; both in standard length. Snout 2.7 to 4.3; eye 2.5 to 3.8; preorbital width 5.3 to 9.4; length of upper jaw 2.7 to 3.0; postorbital part of head 2.6 to 2.8; interorbital width 2.9 to 3.0; depth of caudal peduncle 1.8 to 1.9; length of pectoral fin 1.1 to 1.2; of pelvic fins 1.0 to 1.2, of second or third dorsal spines 2.2 to 2.5, of middle caudal rays 0.96 to 1.15; all in length of head (tip of snout to tip of longest opercular spine). Depth of caudal peduncle in its length 0.9 to 1.2; angle of snout profile with lengthwise axis of body 53 to 60 degrees, snout profile convex.

Body deepest between origin of dorsal and insertion of pelvic fins; preorbital, suborbital, and preopercle all serrate, opercle and sub- opercle with strong radiating spines; interopercle with spines only in posterior angle; spines of dorsal fin subequal, fin not deeply notched, soft dorsal and anal somewhat pointed at angle, caudal truncate, its outer angles rounded; scales on top of head occur forward to middle of interorbital area.

Color in alcohol.—Lips and chin yellowish or light brown, nape and interorbital area usually darker brown; a broad white band, slightly wider than eye diameter, extending from origin of spiny dorsal across anterior part of opercle and posterior part of preopercle to posterior part of interopercle, this band narrowly bordered by black; breast

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 61

light tan; back and sides dark blackish brown, posterior half of caudal peduncle pale yellowish; spiny dorsal dusky, soft dorsal dusky basally, pale on distal half; caudal pale yellowish, anal blackish; pectorals pale whitish or yellowish; pelvics black.

Remarks.—This species is characterized by the single white bar on the head; plain black body; pale soft dorsal, caudal, and pectoral; but black pelvics and anal.

Genus DASCYLLUS Cuvier

Dascyllus Cuvier, Régne animal, ed. 2, vol. 2, p. 179, 1829 (type species, Chaetodon aruanus Linnaeus).

KEY TO SPECIES OF DASCYLLUS FROM THE NORTHERN MARSHALL ISLANDS AND ADJOINING REGIONS

la. Dorsal rays XII,12, the second spine % the length of the third; head and body with 3 broad transverse black bands, first from origin of dorsal fin through eye to chin, second from about middle of spinous dorsal through base of pectorals to pelvics, third from anterior soft dorsal rays to anal rays, spaces between bands whitish or brownish; caudal fin entirely pale or slightly dusky basally; anal rays II,12 or 13; pectoral ii,15 or 16;

gill rakers on first arch 23 or 24___------------ D. aruanus (Linnaeus) 1b. Dorsal rays XII,14 to 16, the second spine subequal to third, pectoral rays 11,18 or 19.

2a. Body with a narrow brown vertical bar extending from origin of dorsal through pectoral base to base of pelvics (young specimens 14 to 16 mm. with a second vertical dark band from soft dorsal to anterior part of anal); general body color light yellowish, each scale with a narrow brown submarginal line; dorsal rays XII,14 to 16 (usually 15); anal

11,13 or 14; gill rakers on first arch 27 or 28. D. reticulatus (Richardson) 2b. Body without dark vertical bar but with a white spot on sides below 7th to 11th dorsal spines; sometimes with a small white nuchal spot (in

young).

3a. White spot on sides small or rarely absent, only slightly larger than pupil (in specimens of all sizes), situated in or just above 11th to 13th lateral line scales; general body color brown; each scale with dark submarginal line; dorsal rays XII,15; anal II,14; gill rakers on first anchi23: tOUl0 he nse oe ee D. trimaculatus (Riippell) 3b. White spot on sides larger than eye (in specimens of all sizes); extending from base of dorsal to below lateral line, or in specimens under 25 mm. white spot usually a wedge-shaped bar narrowing ventrally; indistinct but still evident in large specimens (75 to 85 mm.); general body pattern reticulated in large specimens the scales with pale centers and dark submarginal line, blackish in smallest specimens; head usually very dark brown; dorsal rays XII,16 (rarely 15); anal 11,15 or 16; gill rakers on first arch 25. (Not yet found in Marshall Talands§c2 201 sion LOL RS wal ho ee D. albisella Gill

62 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2 DASCYLLUS ARUANUS (Linnaeus) PuatEe 80,B

Chaetodon aruanus LINNAEUS, Systema naturae, ed. 10, p. 275, 1758 (type locality,

Indies). SPECIMENS STUDIED

Bikini Atoll: 11 stations, 192 specimens, 7 to 35 mm. in standard length.

Eniwetok Atoll: 2 stations, 34 specimens, 23 to 60 mm.

Rongelap Atoll: 9 stations, 147 specimens, 10.5 to 54 mm.

Rongerik Atoll: 2 stations, 2 specimens, 37 to 40 mm.

Likiep Atoll: 1 lot, Univ. of Washington, 37 specimens, 23 to 58 mm.

Guam: 6 lots, 88 specimens, 20 to 53 mm.

Saipan: 2 lots, 30 specimens, 6.5 to 39 mm.

Description.—Dorsal fin rays XII,12; anal rays JI,12 or 13; pectoral rays 11,15 or 16; scale rows from upper edge of gill opening to base of caudal rays 26 to 29, from base of dorsal to lateral line 4, from lateral line to origin of anal 10; gill rakers 6 or 7+1+15 to 17, totaling 23 or 24,

Depth of body 1.5 to 1.7; head 2.6 to 4.0; both in standard length. Snout 3.15 to 5.2; eye 2.4 to 3.0; preorbital width 6.6 to 6.7; upper jaw 3.2 to 3.4; postorbital part of head (hind margin of eye to upper edge of gill opening) 2.75 to 3.4; interorbital width 2.6 to 2.95; depth of caudal peduncle 1.9 to 2.1; length of pectoral fin 1.1 to 1.2, of pelvics 0.8 to 0.9, of 3rd dorsal spine 1.4 to 1.9, of upper caudal ray 0.8 to 1.0; all in length of head (tip of snout to posterior tip opercular spine). Depth of caudal peduncle in its length 1.0 to 1.9; angle of snout profile with lengthwise axis of body 63 to 67 degrees, upper profile convex.

Shape of body nearly round in young, oval in adults; preorbital, suborbital, preopercle, and subopercle serrate, opercle serrate near upper edge of gill opening and with a, flat triangular spine; scales on snout anterior to nostrils; anterior rays of dorsal and anal longest, these fins angular; caudal with two free spines near base above and below, tips of lobes pointed, fin slightly forked.

Color in alcohol.—Head and body with three broad transverse black bands, the first from origin of dorsal through eye to chin, covering anterior part of head, except for yellowish interorbital area, lips blackish; the second from about middle of spinous dorsal through base of pectoral to pelvics; the third from anterior soft dorsal rays to anal rays; interspaces whitish, scales with white centers, brown margins, or entirely brown; anus black; spiny dorsal membranes black; tips of soft dorsal rays white, posterior rays of soft anal with white tips; caudal peduncle and caudal fin pale; pectoral rays pale, pelvics black.

Color when alive.—As described for preserved specimens, except that interorbital area and lips are grayish, spaces between first and second black bars silvery, base of caudal dusky.

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 63

Remarks.—D. aruanus should not be confused with D. melanurus Bleeker, which is illustrated in plate 81,E.

DASCYLLUS RETICULATUS (Richardson)

PuaTE 82,B,C

Heliases reticulatus Ricnarpson, Rep. Fifteenth Meet. British Assoc. Adv. Sci., p. 245, 1846 (type locality, China, Japan). Dascyllus marginatus FowLEeR and Brean (not Riippell) U. 8S. Nat. Mus. Bull. 100, vol. 7, p. 17, 1928 (part, see remarks). SPECIMENS STUDIED

Bikini Atoll: 2 stations, 11 specimens, 14 to 50 mm. in standard length.

Rongelap Atoll: 1 station, 1 specimen, 16 mm.

Likiep Atoll: 1 lot, Univ. Washington, 1 specimen, 25 mm.

Description.—Dorsal fin rays XIJ,14 to°16 (usually 15), anal II,13 or 14 (usually 14); pectoral ii, 18 or 19; scale rows from upper edge of gill opening to base of caudal rays 27 or 28, with 4 between base of dorsal and lateral line and 11 between lateral line and origin of anal; gill rakers 7 or 8-++-1+19 totaling 27 or 28.

Depth of body 1.5 to 1.6; head 2.7 to 3.3; both in standard length. Snout 3.5 to 3.9; eye 2.0 to 2.8; preorbital width 6.5 to 8.4; length of upper jaw 2.3 to 3.3; postorbital part of head (hind margin of eye to upper edge of gill opening) 2.5 to 3.0; interorbital width 2.5 to 3.9; depth of caudal peduncle 1.8 to 2.0; length of pectoral fin 0.97 to 1.1, of pelvics 0.8 to 0.9, of second dorsal spine 1.1 to 1.3, of upper caudal ray 0.8 to 0.9; all in length of head (tip of snout to hind margin of opercular membrane). Depth of caudal peduncle in its length 1.2 to 1.4; angle of snout profile with lengthwise axis of body 60 (young) to 76 (adult) degrees.

Shape of body oval-elliptical, preorbital and suborbital, preopercle finely serrated, opercle entire except a flat rounded spine on upper half, scales on snout to tip; first dorsal spine about ¥% length of second, second spine somewhat curved, subequal with 3rd; teeth in outer row large, conical, widely spaced, inner row numerous fine, close-set.

Color wn alcohol.——Ground color light yellowish brown, margins of scales darker brown on specimens 14 to 16 mm. with dark brown vertical bar, slightly broader than pupil running from origin of dorsal through base of pectorals to base of pelvics, a second vertical bar from soft dorsal to anterior part of anal; in specimens 20 to 50 mm. the posterior bar disappears leaving only anterior one which becomes paler and narrower; in 60 mm. specimens the anterior bar has also disap- peared leaving only the reticulated pattern of the scales; in specimens of all sizes the spiny dorsal membrane is dark brown or blackish, base of soft dorsal dusky; anal dark brown on basal \ or entirely dark brown or black; caudal dusky brown on basal %; pelvics dark brown or

64 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

black; pectorals pale; small round intensely black spot on upper edge of pectoral base not extending below the base of third ray (more dis- tinct in specimens of 40 to 60 mm. than in smaller ones).

Remarks.—This species has been considered a synonym of D. marginatus (Riippell, Atlas zu der Reise im nérdlichen Afrika . Fische des rothen Meers, p. 38, pl. 8, fig. 2, 1828) by many authors (see Fowler, Mem. Bishop Mus., vol. 10, p. 306, 1928, Fowler and Bean, 1928, Montalban, Bur. Sci. Manila Monogr. 24, p. 24, 1927, Weber and Beaufort, Fishes of the Indo-Australian Archipelago, vol. 8, p. 465, 1940), but we conclude it is distinct (see pl. 81,A). We have carefully compared specimens from the Marshall and Philip- pine Islands with those of comparable size from Zanzibar, as well as with Riippell’s figure, and find them to be distinctly different in coloration. D. marginatus does not possess a dark vertical bar from dorsal origin through pectoral base to pelvic base, but instead has a large dark area (this may be lacking in some preserved specimens) covering entire nape and upper anterior sides downward and back- ward, fading behind pectoral, as clearly shown in Rippell’s figure though not in that of Cuvier and Valenciennes (Histoire naturelle des poissons, vol. 5, p. 437, pl. 133, 1830); pectoral base blackish except opposite lower 2 or 3 rays; furthermore, the general ground color of our specimens from Zanzibar is dark greenish brown as stated by Cuvier and Valenciennes and these specimens have small brown dots in middle of scale bases, the scales are not dark margined as in D. reticulatus.

Giinther (Catalogue of the fishes in the British Museum, vol. 4, p. 14, 1862), who had specimens of marginatus from Riippell’s collec- tion, noticed these differences and separated the two species, discussing one under D. zanthosoma Bleeker and including a discussion of Heliases reticulatus Richardson pointing out that the type specimens of the latter appear to be lost.

DASCYLLUS TRIMACULATUS (Riippell) PLATE 82,A,E

Pomacentrus trimaculatus Ri prevy, Atlas zu der Reise im nordlichen Afrika .. . Fische des rothen Meers, p. 39, pl. 8, fig. 3, 1828 (type locality, Massaua, Red Sea).

Dascyllus trimaculatus KENDALL and GoLpsBoroucH, Mem. Mus. Comp. Zool. vol. 26, p. 298, 1911 (Jaluit, Marshall Islands).

SPECIMENS STUDIED

Rongelap Atoll: 1 station, 1 specimen, 57 mm. standard length. Jaluit Atoll: Albatross, 1 specimen, 93 mm.

Arno Atoll: 1 lot, 1 specimen, 44 mm.

Guam: 1 lot, 1 specimen.

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 65

Description—Dorsal fin rays XIJ,15; anal rays II,14; pectoral rays ii,18 or 19; scale rows, upper edge of gill opening to base of caudal rays 27 or 28, base of dorsal to lateral line 4; lateral line to origin of anal 10 or 11; gill rakers 7+1+-15 to 17 totaling 23 to 25.

Depth of body 1.5; head 3.3; both in standard length. Snout 3.2 to 3.7; eye 2.4 to 2.9; preorbital width 6.2 to 7.6; upper jaw 3.0 to 3.3; postorbital part of head (hind margin of eye to upper edge of gill opening) 3.1 to 4.0; interorbital width 2.6 to 2.7; depth of caudal peduncle 1.6 to 1.8; length of pectoral fin 0.9 to 1.0, of pelvics 0.7 to 0.8, of second dorsal spine 1.2 to 1.3, of upper caudal rays 0.9; all in length of head (tip of snout to hind margin opercle). Depth of caudal peduncle in its length 1.0 to 1.15; angle of snout profile with lengthwise axis of body 78 to 80 degrees, snout profile slightly concave, interorbital convex, upper profile convex.

Shape of body oval in young, anterior profile much steeper in adults; preorbital, suborbital, and preopercle very finely serrated; opercle with broad flat triangular spine, its apex somewhat rounded, lower portion of opercle serrated; scales on snout to tip; first dorsal spine short, about half the length of second, second and third subequal, fifth or sixth dorsal and anal soft rays longest, soft dorsal outline some- what pointed, anal less so; upper caudal lobe longer than lower, caudal fin slightly forked, caudal with two free spines, above and below near its base.

Color in alcohol.—Dark reddish brown to brownish black in large specimens, lips blackish, a small white spot about size of pupil on and just above eleventh to thirteenth lateral-line scales, this spot usually fainter and sometimes absent in large specimens; a white nuchal spot about same size as lateral spot sometimes present; spiny dorsal mem- branes and spines blackish; soft dorsal black where covered with scales, naked portion pale; pelvics, anal, and caudal entirely black; pectorals black basally, usually with small, very black spot just above upper edge of base, rays light tan, membranes hyaline.

Remarks.—Small specimens from the Philippines (19 to 39 mm.) were found to resemble the adults in having the white spot on the sides no larger than the pupil. The general body color was somewhat lighter, the scales having pale centers and dark brown margins, lips dark brown, fins as described in adults. These young of D. trimacu- latus are not at all like the 3 specimens of comparable size in the type series of D. albisella Gill from Honolulu (USNM 6274), which have the head and breast paler than the back and the sides with a large white spot or bar 6 to 8 scales wide (much larger than the eye), and in which the scale margins are pale with bases dark.

Specimens of trimaculatus from the Marshall, Gilbert, Phoenix, and Samoan Islands have the naked part of the soft dorsal fin white or

66 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

pale, whereas that area is dark brown or blackish in specimens from the East Indies to the Red Sea and Mauritius. We believe the speci- mens from the central tropical Pacific may represent a distinct sub- species. D. albisella from Honolulu, plate 82,D, is a distinct species. The detailed relationships of this group of species and subspecies re- main to be worked out.

Genus CHROMIS Cuvier

Chromis Cuvier, Mem. Mus. Hist. Nat. Paris, vol. 1, p. 393, 1815 (type species, Sparus chromis Linnaeus).

KEY TO SPECIES OF CHROMIS FROM THE NORTHERN MARSHALL ISLANDS

la. Preopercle finely serrate; scales of nape and back with small basal auxiliary scales; lower margin of suborbital partly free; upper parts dark brownish; sides and belly lighter; pelvics dusky, anterior portions soft dorsal and anal dark, posterior rays pale; outer caudal rays dusky, median rays pale; dorsal rays XII,11 or 12; anal II,11; pectoral ii,16. C. lepidolepis Bleeker 1b. Preopercle entire or vrenulate; no basal auxiliary scales on nape or back. 2a. Soft dorsal and anal bases naked, outer row of teeth of lower jaw directed outward near symphysis; margin of suborbital obscure, hidden by scales; soft dorsal and anal membranes hyaline; outer caudal rays dusky; pelvies pale or slightly dusky; dorsal rays XII,10 (sometimes 9); anal II,10 (rarely IT,9). 3a. Pectoral rays usually ii,15 or 16 (occasionally 17); pectoral axil dusky but

not completely black._..__ C. caeruleus (Cuvier and Valenciennes) 3b. Pectoral rays usually ii,17 or 18 (occasionally 16 or 19); pectoral axil blgckie2 4. ees Sees C. atripectoralis Welander and Schultz

2b. Soft dorsal and anal bases scaled; outer row of teeth of lower jaw not directed outward.

4a. Pectoral base with a large black spot, pectoral axil black; suborbital margin free, distinct; caudal peduncle and caudal fin pale yellowish;

pelvics black. 5a. Head and body dark purplish brown; caudal peduncle abruptly pale, soft dorsal dark brown on bases of first 2 or 3 rays, the rest entirely pale or white; dorsal rays XII,12; anal II,12; pectorals ii,15; tubular scales in upper part of lateral line 17 or 18; gill rakers CSTE rh eae US a> pe RN ea ae a NS ee a C. dimidiatus (Klunzinger) 5b. Head light yellowish brown, back, sides, and caudal peduncle, light purplish brown; soft dorsal with first four rays entirely brown, the rest dark basally, pale distally; dorsal rays XII,13; anal II,12 to 14 (usually 13); pectorals ii,14 or 15 (usually 15); tubular scales in upper part of lateral line 15 or 16 (usually 15); gill rakers 8 or 9-+1-+-21sor 2231 oF 322 26 asst See C. leucurus Gilbert 4b. Pectoral base without large black spot; pectoral axil pale; suborbital adnate, its margin not evident; broad distinct brown lengthwise band on upper and lower caudal rays; pelvics pale; dorsal rays XII,11; anal IT,11; pectorals ii,15 or 16__..C. ternatensis (Bleeker)

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 67

CHROMIS LEPIDOLEPIS Bleeker PLATE 83,A

Chromis lepidolepis BLEEKER, Versl. Akad. Wet. Amsterdam, ser. 2, vol. 10 (1876), p. 389, 1877 (type locality, Timor).

Dascyllus pomacentroides KenpALL and GoLtpsporoucH, Mem. Mus. Comp. Zool., vol. 26, p. 298, pl. 5, fig. 1, 1911 (type locality, Taritari, Gilbert Islands, holotype USNM 65812 studied).

Serrichromis pomacentroides Fowuer, U.S. Nat. Mus. Bull. 100, vol. 14, part 2,

pp. 77, 78, 1943. SPECIMENS STUDIED

Bikini Atoll: 1 station, 2 specimens, 50 to 55 mm. in standard length; Univ. of Washington, 1 lot, 1 specimen, 46 mm.

Description.—Dorsal fin rays XII,11 or 12; anal rays II,11; pectoral rays ii,16; transverse scale rows from upper edge of gill opening to base of caudal rays 27, with 2 between lateral line and origin of dorsal, 9 between lateral line and origin of anal; upper part of lateral line with 16 or 17 tubular scales; gill rakers 8+-1+ 20, totaling 29.

Depth of body 2.1; length of head 3.3 to 3.4; both in standard length. Snout 3.9 to 4.1; eye 2.7; least preorbital width 12.3 to 12.5; length of upper jaw 2.9 to 3.0; postorbital part of head (hind margin of eye to upper edge of gill opening) 2.8 to 3.0; interorbital width 2.8 to 3.0; depth of caudal peduncle 2.0 to 2.1; length of pectoral fin 1.1 to 1.2, of pelvic fin 0.8 to 0.9, of 3rd dorsal spine 1.8 to 2.0, of upper caudal rays 0.7 to 0.80, of lower caudal rays 0.8 to 0.9; all in length of head (tip of snout to posterior margin of opercular membrane). Depth of caudal peduncle in its length 1.4 to 1.5; angle of upper pro- file with lengthwise axis of body 43 degrees.

Teeth of jaws in 2 rows, those of outer row heavy, conical, widely spaced, those near median line enlarged, inner row teeth smaller; snout short, scaled to a point just anterior to nostrils; preorbital and suborbital scaled, the latter with lower margin partly free; preopercle with posterior margin finely serrate, lower margin entire; some scales of body particularly of nape and upper anterior sides with small basal scales, bases of soft dorsal and anal scaled, soft part of these fins angular; caudal lobes pointed.

Color in alcohol—Lips dusky, upper part of head dark reddish brown, lower part paler; back and upper sides reddish brown, lower sides paler yellowish brown with faint lengthwise reddish brown lines following scale rows; spiny dorsal dusky, edges of membranes black; anterior dorsal rays blackish, posterior rays dusky, membranes clear ; soft anal with anterior half black, posterior half pale; outer caudal rays dark brown or blackish; middle caudal rays lighter but dusky;

68 U.S. NATIONAL MUSEUM BULLETIN 202 Vol. 2

pelvic rays grayish, membranes clear, pectoral similarly colored, base of pectoral dark reddish brown, minute black spot on base of upper ray.

Remarks.—A careful examination of the type of Dascyllus pomacen- troides reveals 2 or 3 scales, with small basal scales. Most of the scales are missing from the nape, back and anterior sides. The type has the same number of fin rays as specimens of Chromis lepidolepis from the Philippines and as two specimens from Bikini; also, it has similar jaw teeth, serrated preopercle, body and fin shape, and color- ation (although faded).

Fowler has made Dascyllus pomacentroides the type of a new genus Serrichromis, ‘distinguished by its serrated preopercle, strong den- tition in combination with other characters.’ These characters, held in common with C. lepidolepis, certainly serve to separate this species from others of the genus Chromis, but Fowler has already proposed the subgeneric name Lepidochromis Fowler and Bean (U.S. Nat. Mus. Bull. 100, vol. 7, p. 58, 1928), for this species.

We propose, in spite of the differences noted above, to leave this species as a subgenus of Chromis, at least for the present, because its jaw teeth, though heavier, are definitely of the same kind as the genus Chromis, as are its oblique mouth, with its long upper jaw, and its general body shape.

CHROMIS CAERULEUS (Cuvier and Valenciennes)

PiaTE 84,B

Heliases caeruleus CUvIER and VALENCIENNES, Histoire naturelle des poissons, vol. 5, p. 497, 1830 (type locality, New Guinea; Ulea).

Heliases lepisurus Cuvier and VALENCIENNES, Histoire naturelle des poissons, vol. 5, p. 498, 1830 (type locality, New Guinea).

Heliases frenatus CuvieER and VALENCIENNES, Histoire naturelle des poissons, vol. 5, p. 498, 1830 (type locality, Guam).—Savuvace, Histoire naturelle des

poissons, in Grandidier, Histoire . . . de Madagascar vol. 16, p. 436, pl. 28, fig. 1, 1887 (Madagascar). Chromis lepisurus BLEEKER, Atlas ichthyologique ..., vol. 9, pl. 403, fig. 7,

1877; Nat. Verh. Holl. Maatsch. Wetensch., vol. 2, No. 6, p. 164, 1877 (East Indies; Zanzibar; Andamans; Guam; Ulea).

Heliastes lepidurus GUNTHER, Catalogue of the fishes in the British Museum, vol. 4, p. 63, 1862 (type locality, Amboina, emended spelling for H. lepisurus Cuvier and Valenciennes).—Day, Fishes of India, vol. 2, p. 389, pl. 82, fig. 1, 1877 (Andamans).—GtnruEr, Journ. Mus. Godeffroy, vol. 15 (Fische der Siidsee, pt. 7), p. 238 (in part), pl. 128, fig. D, 1881.

Glyphisodon anabatoides Day (non Bleeker), Proc. Zool. Soc. London, p. 696, 1870.

Glyphisodon bandanensis BLEEKER, Nat. Tijdschr. Nederl.-Indié, vol. 2, p. 248, 1851 (type locality, Neira; Banda).

Chromis caeruleus (in part), JonpAN and SEALE, Bull. U. S. Bur. Fish. vol. 25 (1905), p. 290, 1906 (Samoan Islands; in a letter cited on p. 291, to Dr. Jordan from Dr. Vaillant, who examined the types of caeruleus, frenatus, and lepisurus, all three are referred to a single species by him).—Aoyaa1, Bioge-

FAMILY POMACENTRIDAE—WOODS AND SCHULTZ 69

ographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 186, fig. 14, 1941 (Japan). SPECIMENS STUDIED

Bikini Atoll: 8 stations, 132 specimens, 11 to 75 mm. in standard length.

Eniwetok Atoll: 2 stations, 21 specimens, 29 to 63 mm.

Rongerik Atoll: 2 stations, 9 specimens, 36 to 56 mm.

Rongelap Atoll: 4 stations, 117 specimens, 10 to 68 mm.

Kwajalein Atoll: 1 specimen, 34 mm.

Guam: 3 lots, 121 specimens, 28 to 59 mm.

Saipan: 3 lots, 170 specimens, 8 to 22 mm.

The following lots were distributed to other museums before the new species was sorted out, and thus may contain specimens of aéripectoralis:

Bikini Atoll: Reer Island, northwest side, lagoon reef, August 12, S-46-332,