Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi

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1 aqua, International Journal of Ichthyology Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi John E. Randall 1, John L. Earle 1, and Luiz A. Rocha 2 1) Bishop Museum, 1525 Bernice St., Honolulu, HI , USA. jackr@hawaii.rr.com 2) Hawaii Institute of Marine Biology, University of Hawaii, P.O. Box 1346, Kaneohe, HI 96744, USA. Received: 21 March 2008 Accepted: 31 March 2008 Abstract Five species of razorfishes of the genus Xyrichtys are known from the central and western Pacific: X. woodi from the Hawaiian Islands; X. sciistius (formerly a synonym of X. woodi) from Japan and Taiwan; X. pastellus, described as new from two specimens from Lord Howe Island and three from Elizabeth and Middleton Reefs, New South Wales; X. halsteadi from Papua New Guinea to Tahiti; and X. koteamea from Easter Island. The first three form a complex of closely related species that are distinguished mainly by color, but also by DNA. In addition, X. woodi has a lower modal gill-raker count than the other two sister species. Zusammenfassung Fünf Arten der Schermesserfische der Gattung Xyrichthys sind vom zentralen und westlichen Pazifik bekannt: X. woodi von den Hawaii-Inseln; X. sciistius (früher ein Synonym zu X. woodi) von Japan und Taiwan; X. pastellus, neu beschrieben nach zwei Exemplaren von der Insel Lord Howe und drei Exemplaren der Riffe Elizabeth und Middleton, New South Wales; X. halsteadi von Papua- Neuguinea bis Tahiti; sowie X. koteamea von der Osterinsel. Die ersten drei bilden einen Komplex nahe verwandter Arten, die sich hauptsächlich durch ihre Farbe unterscheiden, doch auch nach DNA-Untersuchungen. Bei X. woodi ist außerdem die mittlere Kiemenblätter-Zahl geringer als bei den beiden nahestehenden Arten. Résumé Cinq espèces de poisson rasoirs du genre Xyrichtys sont connues en provenance du Pacifique central et occidental: X. woodii des îles Hawaï; X. sciistius (auparavant synonyme de X. woodii) du Japon et de Taïwan; X. pastellus, décrit comme nouvelle espèce, sur base de deux spécimens de l île Lord Howe et trois de Elizabeth et Middleton Reef, Nouvelle Galles du Sud; X. halsteadi de Papouasie Nouvelle- Guinée jusqu à Tahiti; et X. koteamea de l île de Pâques. Les trois premières forment un complexe d espèces très proches qui se distinguent surtout par la couleur, mais aussi par l ADN. En outre, X. woodi a un nombre moins élevé de branchiospines que les deux autres espèces soeurs. Sommario Cinque specie di pesci rasoio del genere Xyrichtys sono note dal Pacifico centrale e occidentale: X. woodi delle isole Hawai; X. sciistius (precedentemente considerato un sino - nimo di X. woodi) originario di Giappone e Taiwan; X. pastellus, descritta come nuova sulla base di due esemplari originari dell isola Lord Howe Island e tre da Elizabeth e Middleton Reefs, Nuovo Galles del Sud; X. halsteadi diffuso da Papua New Guinea a Tahiti; e X. koteamea presente solo all isola di Pasqua. Le prime tre formano un com - plesso di specie strettamente imparentate che si distinguono principalmente per la colorazione, ma anche per il loro DNA. Inoltre, X. woodi ha un valore modale di ra - strelli branchiali inferiore alle altre due specie sorelle. INTRODUCTION Labrid fishes of the genera Iniistius and Xyrichtys are known by the common name razorfishes, in reference to their compressed bodies and the sharp leading edge of their forehead and snout. These are specializations for quick entry into sand with the approach of a predator, enabling these species to exist over open expanses of sand. Most other benthic inshore fishes require the shelter of reefs, beds of seagrass, heavy algal growth, or an existing burrow. Iniistius Gill, 1862 was usually considered a synonym of Xyrichtys Cuvier, 1815 until Randall & Earle (2002) distinguished the two, following the osteological study of Nguyen (1974). The fishes of these two genera can be distinguished externally by characters of the dorsal fin. The origin of the fin is over or less than half an orbit diameter behind the eye in Iniistius, but more behind the eye in 149 aqua vol. 14 no July 2008

2 Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi Xyrichtys. The first two dorsal spines of Iniistius are flexible, whereas only the first is flexible in Xyrichtys. The space between the second and third dorsal spines in Iniistius is much broader than the space between the first and second spines. These two spaces are about equal in Xyrichtys. Some razorfishes have been classified in the genus Hemipteronotus Lacépède, 1801, as by Randall (1965). As recommended by Randall & Bauchot (1993), this generic name was suppressed by Opinion 1799 of the International Commission on Zoological Nomenclature to preserve the names Naucrates Rafinesque, 1810 and Xyrichtys. Species of the latter, such as X. woodi (Jenkins, 1901), have also been placed in the genera Novaculichthys Bleeker, 1862 and Novaculops Schultz, 1960; however, Novaculichthys now contains only one species, N. taeniourus (Lacépède, 1801), and Novaculops is a synonym of Xyrichtys. Six species of Xyrichtys are known for the Atlantic Ocean, and Victor et al. (2001) reported four for the eastern Pacific, one of which is an undescribed species from the Galápagos Islands. Another possible new species of the genus has been photographed at the Revillagigedo Islands (Victor et al. 2001). Only one species of the genus, X. woodi, was recognized from the central and western Pacific region until Randall & Lobel (2003) described X. halsteadi from Papua New Guinea, Guam, Wake Island, and Tahiti, and Randall & Allen (2004) named X. koteamea from Easter Island. In the present paper, we describe a new species from Lord Howe Island and nearby Elizabeth and Middleton Reefs, New South Wales, and show that X. sciistius Jordan & Thompson, 1914 from Japan and Taiwan is a valid species, not a synonym of X. woodi of the Hawaiian Islands. MATERIALS AND METHODS Type specimens of the new species are deposited in the Australian Museum, Sydney (AMS), the Bernice P. Bishop Museum, Honolulu (BPBM), and the U.S. National Museum of Natural History, Washington, D.C. (USNM). Specimens of Xyrichtys sciistius were obtained on loan from the National Museum of Nature and Science, Tokyo (NSMT). The length of specimens is given as standard length (SL), measured from the median anterior end of the upper lip to the base of the caudal fin (posterior end of the hypural plate); body depth is aqua vol. 14 no July 2008 the greatest depth from the base of the dorsal spines to ventral edge of the abdomen (correcting for any malformation of preservation); body width is measured just posterior to the gill opening; head length is taken from the upper lip to the posterior end of the opercular flap; orbit diameter is the greatest fleshy diameter, and interorbital width the least bony width; snout length is measured from the median anterior point of the upper lip to the nearest fleshy edge of the orbit; upper-jaw length from the same anterior point to the posterior end of the maxilla; caudal-peduncle depth is the least depth, and caudal-peduncle length the horizontal distance between verticals at the rear base of the anal fin and the caudal-fin base; lengths of spines and rays are measured to their extreme bases; caudal-fin and pectoral-fin lengths are the length of the longest ray; pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest soft ray. Morphometric data are presented in Table I as percentages of the standard length. Proportional measurements in the text are rounded to the nearest 0.5. Lateral-line scale counts include the last pored scale that overlaps the end of the hypural plate; scales in transverse series are counted from the origin of the anal fin obliquely upward to the base of the first dorsal fin; the count of gill rakers is made on the first gill arch and includes all rudiments. Meristic and morphometric data in parentheses refer to paratypes. Total genomic DNA was extracted from available tissue samples using the QIAGEN DNeasy tissue kit following the manufacturer s protocol. A fragment of the cytochrome oxidase I (COI) gene from the mitochondrial DNA (mtdna) was amplified for nine specimens, three from Japan, five from Hawaii and one from Australia. Primers used for amplification and sequencing were BOL-F1 (5 TCA ACY AAT CAY AAA GAT ATY GGC AC 3 ) and BOL-R1 (5 ACT TCY GGG TGR CCR AAR AAT CA 3 ), modified from Ward et al. (2005). Detailed PCR and sequencing conditions are the same as those in Rocha (2004). Xyrichtys pastellus n. sp. (Figs 1-3, Tables I-II) Hemipteronotus sp. Allen et al. 1976: 420 (Lord Howe Island). 150

3 John E. Randall, John L. Earle and Luiz A. Rocha Novaculops sp. Francis 1993: 165 (Lord Howe Island). Novaculops sp. Francis & Randall 1993: 122, pl. II, fig. B (Lord Howe Island and Middleton Reef). Holotype: AMS I , mm, presumed female, Lord Howe Island, near Admiralty Islands, estimated S E, hook and line, Darrin Nobbs, 16 March 2006 (viscera removed). Paratypes: BPBM 14758, 66.2 mm, Lord Howe Island, reef west of Mt. Lidgbird, 25 m depth, rotenone, J. E. Randall, G. R. Allen, B. Goldman, B. C. Russell, D. F. Hoese, and G. P. Whitley, 5 February 1973; USNM , 77.7 mm, New South Wales, Elizabeth Reef, north side, 14 m depth, dug from sand by hand, J. L. Earle, 24 February 1992; AMS I , mm, same data as preceding; BPBM 35027, male, mm, New South Wales, Middleton Reef, off entrance to lagoon west of wreck of Japanese longliner, sand channel, 10 m depth, caught by hand, J. L. Earle, 25 February Diagnosis: Dorsal rays IX,12; anal rays III,12; pectoral rays 13; lateral line interrupted, the pored scales ; cheek naked; gill rakers 18-20; body depth in SL; dorsal profile of snout about 50 to horizontal axis of body; first dorsal spine flexible and not long, in head length; sec- ond to ninth dorsal spines stiff and sharp-pointed; space between first and second dorsal spines about equal to space between second and third spines; color of body of presumed female in life pale bluegreen, the scales edged in pale orange or reddish, becoming maroon with vertical orange lines ventrally on abdomen, and blue with maroon edges on scales above anal fin; a broad pale orange zone extending posteriorly from beneath pectoral fin, altering to pale green as it passes above anal fin; axil of pectoral fin deep blue; pores of lateral line white; iris pink and white; spinous portion of dorsal fin blue; remaining median fins translucent graybrown; color of male when fresh pale gray, the edges of scales greenish gray except ventrally; sixth and seventh interspinous membranes of dorsal fin nearly black, with blackish pigment continuing obliquely onto body to lateral line; pores of lateral line white; transverse dusky bands ventrally on head; iris red. Description: Dorsal rays IX,12, the first two rays unbranched, the last branched to base; anal rays III,12, all branched; pectoral rays 13, the first two rays unbranched (first ray very short and closely applied to second); pelvic rays I,5, all soft rays branched; principal caudal rays 12, all branched; upper procurrent caudal rays 5 (5-6); lower procurrent caudal rays 5; lateral-line scales , the last Fig. 1. Holotype of Xyrichtys pastellus, presumed female, AMS I , 106 mm SL, Lord Howe Island. Aquarium photo by G. Kelly. 151 aqua vol. 14 no July 2008

4 Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi Table I. Proportional measurements of type specimens of Xyrichtys pastellus as percentages of standard length. Holotype Paratypes AMS I. BPBM USNM BPBM AMS I Standard length (mm) Body depth Body width Head length Snout length Orbit diameter Interorbital width Upper-jaw length Caudal-peduncle depth Caudal-peduncle length Predorsal length Preanal length Prepelvic length Base of dorsal fin First dorsal spine Second dorsal spine Third dorsal spine Ninth dorsal spine 11.2 broken Longest dorsal ray Base of anal fin First anal spine Second anal spine Third anal spine Longest anal ray Caudal-fin length Pectoral-fin length Pelvic-spine length Pelvic-fin length Mouth moderately large, the maxilla nearly reaching a vertical at anterior edge of orbit, the upperjaw length 3.35 ( ) in head length; mouth slightly oblique, forming an angle of about 10 to horizontal axis of body (nearly 20 in small paratype); a pair of large, recurved, outflaring canine teeth at front of jaws that overlap lips when mouth closed (jaws of holotype seem abnormal and cannot close completely); lower pair of canines medial to upper pair; side of jaws with a series of about 12 strong conical teeth, smaller posteriorly; an inner row of nodular teeth behind front canines and continuing closely medial to teeth of side of jaws. Tongue broadly rounded, set far back in mouth. Lips thin, the lower with a flap along side of mandible that is longer posteriorly. Gill rakers short, the longest on first arch one-half length of longest gill filaments. Posterior edge of preopercle free slightly dorsal to level of lower edge of orbit; ventral edge of preopercle free to a vertical at anterior edge of orbit; a narrow fleshy rim to orbit with a free edge in holoon base of caudal fin; scales above lateral line to origin of dorsal fin 4; scales above lateral line to middle of spinous portion of dorsal fin 2 (uppermost small); scales below lateral line to origin of anal fin 9 (the last small); circumpeduncular scales 16; gill rakers 19 (18-20); branchiostegal rays 5; vertebrae 25. Body moderately deep, the depth 2.75 ( ) in SL; body very compressed, the width 2.85 ( ) in body depth; head length 3.1 ( ) in SL; snout length (as measured from lower edge of orbit to front of upper lip) 2.85 ( ) in head length; dorsal profile of head convex, the profile of snout forming an angle of about 50 to horizontal axis of body; front of snout narrowing to sharp ridge that extends to above eye; leading edge of chin rounded; minimum distance from lower edge of orbit to corner of mouth equal to orbit diameter; orbit diameter 4.5 ( ) in head length; interorbital width 7.25 ( ) in head length; caudal-peduncle depth 2.3 ( ) in head length; caudal-peduncle length 3.6 ( ) in head length. aqua vol. 14 no July

5 John E. Randall, John L. Earle and Luiz A. Rocha type below posterior half of eye. Nostrils very small, the posterior oval with a slight rim, three-fourths pupil diameter in front of upper third of eye; anterior nostril a short tube with a small posterior flap, slightly ventral, the internarial space about onefourth pupil diameter. Cephalic sensory pores very small, 9 in suborbital series, the first below nostrils, the next two at end of short ventral branches, the last behind eye; a series of 7 preopercular pores, continuing anteriorly as 3 mandibular pores. Scales cycloid and very thin; lateral-line scales with a single horizontal tubule, ending posteriorly in a pore; scales on chest about three-fifths height of largest scales on side of body; head naked except for a single small partly embedded scale dorsally on opercle; no scales on dorsal and anal fins; four rows of progressively smaller scales on base of caudal fin (counting obliquely, the last pored scale of lateral line in the first row); no prepelvic scales (two partly embedded scales in paratype); no pelvic axillary Fig. 2. Paratype of Xyrichtys pastellus, male, BPBM 35027, 117 mm SL, Middleton Reef, New South Wales. Photo by J. E. Randall. Fig. 3. Paratype of Xyrichtys pastellus, BPBM 14758, 66.2 mm SL, Lord Howe Island. Photo by J. E. Randall. 153 aqua vol. 14 no July 2008

6 Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi Table II. Gill-raker counts of species of Xyrichtys. Gill rakers X. pastellus X. sciistius X. woodi dorsal spine slender and flexible, the remaining dorsal spines sharp-pointed and stiff; space between first and second dorsal spines about equal to space between second and third spines; membrane not notched between second and third spines; first dorsal spine 3.4 ( ) in head length; ninth dorsal spine longest, 2.9 (3.0) in head length (first spine longest in smallest paratype, 3.0 in head length); fifth to seventh dorsal soft rays longest, 2.4 ( ) in head length; origin of anal fin below base of first dorsal soft ray, the preanal length 1.9 ( ) in SL; third anal spine longest, 3.9 ( ) in head length; sixth and sevscale; a single pointed scale extending posteriorly from between base of pelvic fins, its length about one-third length of pelvic spine. Origin of dorsal fin above upper free end of preopercle (two-thirds orbit diameter posterior to eye), the predorsal length 3.85 ( ) in SL; first Fig. 4. Xyrichtys sciistius, presumed female, Izu Peninsula, Japan. Aquarium photo by J. E. Randall. Fig. 5. Xyrichtys sciistius, male, NSMT-P 33516, 144 mm SL, Ani-jima, Ogasawara Islands. Photo by J. E. Randall. aqua vol. 14 no July

7 John E. Randall, John L. Earle and Luiz A. Rocha enth anal soft rays longest, 2.4 ( ) in head length; caudal fin slightly rounded, 4.75 ( ) in SL; third or fourth pectoral rays longest, reaching to above anus (to above origin of anal fin in smallest paratype), 4.75 (4.7) in SL; pelvic fins reaching anus (reaching anal-fin origin in male paratype), the first soft ray longest, 4.9 ( ) in SL; pelvic spine 3.95 ( ) in head length. Color of holotype in alcohol pale gray, a little paler below and behind pectoral fin, with a prominent black spot in axil of pectoral fin; spinous portion of dorsal fin dusky with a faint blackish spot near base of fourth to eighth membranes; remaining fins translucent pale yellowish. Color of male paratype in alcohol pale purplish gray, the scale edges a little darker dorsally on body; fins translucent yellowish, the sixth and seventh interspinous membranes of dorsal fin dusky with a large blackish spot; scales below these membranes to lateral line a little dusky. Color of small paratype in alcohol light brown; axil of pectoral fin blackish; a large blackish spot on first membrane of dorsal fin, and a spot of lesser intensity on second membrane; fins otherwise translucent pale yellowish. Color of holotype in life shown in Fig. 1. We believe the irregular dusky bars and Fig. 6. Xyrichtys sciistius, BPBM 35061, 62 mm SL, Chichi-jima, Ogasawara Islands. Photo by J. E. Randall. Fig. 7. Xyrichtys sciistius, BPBM 35187, 43 mm SL, Chichi-jima, Ogasawara Islands. Underwater photo by J. E. Randall. 155 aqua vol. 14 no July 2008

8 Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi blotches on the body are a disruptive pattern that appears when the fish is stressed. Color of paratypes when fresh as in Figs 2 and 3. Etymology: The specific name for this razorfish, pastellus, is from the Late Latin word referring to colors that are soft and subdued, as in the holotype. Remarks: The following six nominal species of Xyrichtys from the central and western Pacific have been described: X. entargyreus (Jenkins, 1901) from the Hawaiian Islands; X. halsteadi Randall & Lobel, 2003 from Papua New Guinea, Guam, Wake Island, and Tahiti; X. koteamea Randall & Allen, 2004 from Easter Island; X. sciistius Jordan & Thompson, 1914 from Japan; X. tatoo Seale, 1901 from the Hawaiian Islands, and X. woodi (Jenkins, 1901) from the Hawaiian Islands. Xyrichtys entargyreus, X. tatoo, and X. sciistius have long been considered as synonyms of X. woodi; Fig. 8. Xyrichtys woodi, presumed female, Oahu, Hawaiian Islands. Underwater photo by J. E. Randall. Fig. 9. Xyrichtys woodi, presumed male, Oahu, Hawaiian Islands. Underwater photo by J. P. Hoover. aqua vol. 14 no July

9 John E. Randall, John L. Earle and Luiz A. Rocha however, Randall & Allen (2004) wrote that color differences suggest that X. sciistius is probably a distinct species. All of these species have the same fin-ray and scale counts. Xyrichtys halsteadi is the most diverse, with a more slender body (body depth in SL), small size (largest, 120 mm SL), and very different color pattern. Xyrichtys koteamea is easily separated by its deeper body (depth in SL), shorter pectoral fins, large size ( mm SL), and predominantly deep red color. The remaining three valid species have essentially the same body and fin proportions and share some color features, such as a deep blue or blackish spot on the interspinous membranes of the dorsal fin (also found in X. koteamea), the lateral-line pores in a white spot (at least anteriorly), a red iris, and (in juveniles and females) near-vertical white lines on the abdomen and a white patch beneath the pectoral fin that narrows posteriorly. Xyrichtys pastellus and X. sciistius have a deep blue or black spot in the axil of the pectoral fins, in contrast to the red or dusky axil in X. woodi. Adults of X. sciistius exhibit much more orange or red coloration than is found in adults of X. pastellus or X. woodi. Considerable color variation of X. sciistius may be seen in Masuda & Kobayashi (1994: 289: figs 3-8) and Okamura & Amaoka (1997: 518, lower 5 left figures). Figures 1-3 of X. pastellus, Figs 4-7 of X. sciistius, and Figs 8-10 of X. woodi provide for a comparison of the color patterns of these three species. Table II shows a modal difference in the gill-raker counts of X. woodi from X. sciistius and X. pastellus. For a comparison of the DNA, the third author sequenced tissue samples of Xyrichtys pastellus, X. sciistius, and X. woodi using the barcode CO1 gene. The three differed from one another by an average of 0.7% sequence divergence. While this is relatively small, it is supported by the color difference, as well as the isolation of the populations. From the distributions, X. sciistius would be expected to be genetically closest to the Hawaiian X. woodi. However, these two species have, in addition to color, a clear modal difference in gill-raker counts. The DNA differences have enabled us to more confidently treat these three razorfishes as species. Genetics has been used before to help define species, and divergences of this magnitude between closely related sister species are not uncommon (Rocha et al., 2007). Benjamin C. Victor (pers. comm.) has informed us that he has barcode sequences for many of the razorfishes and found that all but one have less than 0.5% difference. It is of interest to note that the present records of the species of Xyrichtys in the central and western Pacific show an antitropical distribution, except for the one collection of X. halsteadi from Papua New Guinea. We should also mention the probable invalid name Xyrichtys javanicus (Bleeker, 1862). De Beaufort (1940: 66) wrote: Bleeker described this species after a specimen labelled,,java and belonging to the old collection of the Leiden Museum. It shows strong likeness to species from the Atlantic and probably the locality mentioned on the label is wrong. It has never been found again. However, Dor (1970: 22) reported one preserved specimen, Fig. 10. Xyrichtys woodi, BPBM 37044, 43 mm SL, Oahu, Hawaiian Islands. Photo by J. E. Randall. 157 aqua vol. 14 no July 2008

10 Xyrichtys pastellus, a new razorfish from the southwest Pacific, with discussion of the related X. sciistius and X. woodi 106 mm SL, as Hemipteronotus javanicus from Eilat, Gulf of Aqaba. Additional Red Sea specimens should be obtained, and the life color determined. Material Examined of Other Pacific Species of Xyrichtys: Xyrichtys sciistius: Ogasawara Islands, Chichi-jima, BPBM 35061, 62 mm; BPBM 35187, 43 mm. Ani-jima, BPBM 35081, 2: mm; NSMT-P 33516, 144 mm. Ryukyu Islands, Okinawa Group, Nagan nu Island, NSMT-P , 26 mm. Izu Islands, Miyakejima, NSMT-P 31557, 59 mm. Shizuoka Prefecture, Izu Peninsula, NSMT-P , 6: mm. X. woodi: Hawaiian Islands, Oahu, BPBM 4723, 4: mm; BPBM 4724, 125 mm; BPBM 4725, 2: mm; BPBM 11980, 116 mm; BPBM 24531, 93 mm; BPBM 31028, 2: mm; BPBM 37044, 43 mm; BPBM 37118, 31 mm. ACKNOWLEDGEMENTS Geoff Kelly of the Lord Howe Island Marine Park maintained the holotype of Xyrichtys pastellus in his aquarium after it was caught by Darrin Nobbs, and provided the photograph of Fig. 1. He sent the specimen to Mark A. McGrouther of the Australian Museum, who passed it on to us for the description. We are grateful to all for their contributions. Thanks are also due John P. Hoover for his underwater photograph of a presumed male of X. woodi, Loreen R. O Hara of the Bishop Museum for X-rays, Hiroshi Senou for obtaining tissue samples of X. sciistius collected by M. Konno and S. Kato, Gento Shinohara for the loan of specimens of X. sciistius from the National Museum of Nature and Science in Tokyo, and Benjamin C. Victor for his information on barcode sequences of razorfishes. REFERENCES ALLEN, G. R., HOESE, D. F., PAXTON, J. R., RANDALL, J. E., RUSSELL, B. C., STARCK, W. A. II, TALBOT, F. H. & WHITLEY, G.P Annotated checklist of the fishes of Lord Howe Island. Records of the Australian Museum 30: DE BEAUFORT, L. F The fishes of the Indo-Australian Archipelago, vol. 8. xv pp. E. J. Brill, Leiden. DOR, M Nouveaux poisons pour la faune de la Mer Rouge. Contributions to the knowledge of the Red Sea 44: FRANCIS, M. P Checklist of the coastal fishes of Lord Howe, Norfolk, and Kermadec Islands, Southwest Pacific Ocean. Pacific Science 47 (2): FRANCIS, M. P. & RANDALL, J. E Further additions to the fish faunas of Lord Howe and Norfolk Islands, Southwest Pacific Ocean. Pacific Science 47 (2): JORDAN, D. S. & THOMPSON, W. F Record of the fishes obtained in Japan in Memoirs of the Carnegie Museum 6: MASUDA, H. & KOBAYSHI, Y Grand Atlas of Fish Life Modes (Color Variation in Japanese Fish). 465 pp. Tokai University Press, Tokyo (in Japanese). NGUYEN, T Osteological studies on the labrid fishes (Family Labridae) of Japan Morphology, taxonomy and phylogeny. PhD thesis, Ocean Research Institute, University of Tokyo. OKAMURA, O. & AMAOKA, K Sea Fishes of Japan. 783 pp. Yama-kei Publisher, Tokyo (in Japanese). RANDALL, J. E A review of the razorfish genus Hemipteronotus (family Labridae) of the Atlantic Ocean. Copeia 1965 (4): RANDALL, J. E. & ALLEN, G. R Xyrichtys koteamea, a new razorfish (Perciformes: Labridae) from Easter Island. Raffles Bulletin of Zoology 52 (1): RANDALL, J. E. & BAUCHOT, M. L Case Naucrates Rafinesque, 1810 and Xyrichtys Cuvier, 1814 (Osteichthyes, Perciformes): Proposed conservation. Bulletin of Zoological Nomenclature 50 (4): RANDALL, J. E. & EARLE, J. L Review of Hawaiian razorfishes of the genus Iniistius (Perciformes: Labridae). Pacific Science 56 (4): RANDALL, J. E. & LOBEL, P. S Xyrichtys halsteadi, a new labrid fish from the central and western Pacific. Bulletin of Marine Science 72 (3): ROCHA, L. A Mitochondrial DNA and color pattern variation in three western Atlantic Halichoeres (Labridae), with the revalidation of two species. Copeia 2004 (4): ROCHA, L. A., CRAIG, M. T. & BOWEN, B. W Phylogeography and the conservation of coral reef fishes. Coral Reefs 26: VICTOR, B. C., WELLINGTON, G. M. & CALDOW, C A review of the razorfishes (Perciformes: Labridae) of the eastern Pacific Ocean. Revista de Biologia Tropical 49, suppl. 1: WARD, R. D., ZEMLAK, T. S., INNES, B. H., LAST, P. R. & HERBERT, P. D. N Barcoding Australia s fish species. Philosophical Transactions of the Royal Society B: Biological Sciences, no. 360: aqua vol. 14 no July