Hoplolatilus Randalli, a New Species of Sand Tilefish (Pisces: Malacanthidae) from the Tropical Western Pacific with Comments on the Validity of H

aqua, International Journal of Ichthyology

Hoplolatilus randalli, a new species of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus

Gerald R. Allen1, Mark V. Erdmann2 and Alison M. Hamilton3

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986 2) Conservation International Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235, Bali, Indonesia 3) Department of Ecology and Evolutionary Biology, 621 Charles E. Young Drive South, University of California Los Angeles, Los Angeles CA 90095, USA

Received: 11 December 2009 – Accepted: 24 August 2010

Abstract sich: Bei beiden Arten ist die Schwanzflosse eingekerbt, aber Hoplolatilus randalli n. sp. is described on the basis of 13 bei H. randalli n. sp. ist der hintere Rand der beiden Lappen specimens, 46.7-158.2 mm SL, collected at the Banda leicht konvex, bei H. fronticinctus hingegen gerade. Außer- Islands and West Papua in Indonesia, Luzon in northern dem hat H. randalli n. sp. längere harte Rückenflossen- Philippines, Palau and Yap in Micronesia, and the strahlen (letzter Hartstrahl 2,5-3,0 vs. 3,5-4,1 in HL) und Solomon Islands. The species has previously been confused auch längere weiche Rückenflossenstrahlen (längster Weich- with H. fronticinctus Günther, a closely related species that strahl 1,1-1,4 vs. 1,5-1,7 in HL). Weiterhin werden geneti - is restricted to the Indian Ocean from eastern Africa to the sche Daten mitgeteilt, die die Artentrennung bestätigen. East Andaman Sea. The two species can be distinguished Außerdem geben wir zusammenfassend Belege für die by basic colour differences, particularly the predominately Anerkennung der Art H. luteus wieder (morphologische, greenish colour of H. randalli n. sp. and mainly bluish hue genetische und ontogenetische, auf das Farbmuster bezo- of H. fronticinctus. They also differ in caudal fin lobe gene Unterscheidungsmerkmale), die kürzlich erst als jün- shape; both species have an emarginate caudal fin, but the geres Synonym zu H. fourmanoiri eingestuft wurde. posterior edge of each lobe is slightly convex in H. randalli n. sp. and straight in H. fronticinctus. In addition, H. ran- Résumé dalli n. sp. has longer dorsal spines (last spine 2.5-3.0 vs. Hoplolatilus randalli n. sp. est décrit sur base de 13 spéci- 3.5-4.1 in HL) and longer soft dorsal rays (longest ray 1.1- mens, 46,7-158,2 mm de LS, collectés aux îles Banda et en 1.4 vs. 1.5-1.7 in HL). Genetic data are also provided that Papouasie occidentale (Indonésie), à Luzon (Philippines du confirm its validity. Additionally, we include comprehen- nord), à Palau et Yap (Micronésie) et aux îles Salomon. sive evidence (morphological, genetic, and ontogenetic L’espèce a été jadis confondue avec H. fronticinctus Gün- colour pattern differences) for the recognition of H. luteus, ther, une espèce étroitement apparentée, limitée à l’océan which was recently considered a junior synonym of H. Indien, de l’est africain jusqu’à l’est de la mer d’Andaman. fourmanoiri. Les deux espèces peuvent être distinguées par des couleurs de base différentes, notamment la couleur verdâtre pré- Zusammenfassung dominante chez H. randalli n. sp. et principalement bleuâ- Hoplolatilus randalli n. sp. wird auf der Grundlage von 13 tre chez H. fronticinctus. Elles se démarquent aussi par la Exemplaren mit 46,7-158,2 mm SL beschrieben, die bei forme des lobes de la caudale; les deux espèces ont une cau- den Banda-Inseln und West-Papua in Indonesien gefangen dale concave, mais l’angle postérieur de chaque lobe est wurden sowie bei Luzon von den nördlichen Philippinen, légèrement convexe chez H. randalli n. sp. et droit chez H. bei Palau und Yap von Mikronesien und an den Salomon- fronticinctus. En outre, H. randalli n. sp. a de plus longues Inseln. Diese neue Art war bisher mit H. fronticinctus Gün- épines dorsales (2,5-3,0 contre 3,5-4,1 pour la dernière ther verwechselt worden, die nahe verwandt ist, aber épine) et des rayons dorsaux mous plus longs (le rayon le beschränkt auf den Indischen Ozean von Ostafrika bis zur plus long: 1,1-1,4 contre 1,5-1,7). Des données génétiques östlichen Andamanensee. Die beiden Arten lassen sich sont également fournies pour confirmer sa validité. Par durch die Grundfarbe unterscheiden; vor allem hat H. ran- ailleurs, nous ajoutons des preuves détaillées (différences de dalli n. sp. eine vorherrschend grünliche Farbe, H. fron- coloration morphologiques, génétiques et ontogénétiques) ticinctus hingegen im Wesentlichen einen bläulichen pour l’identification de H. luteus qui était considéré il ya Farbton. Auch die Form der Schwanzflossen unterscheidet peu comme un synonyme récent de H. fourmanoiri.

171 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus

Sommario species mentioned above as well as H. luteus Allen Hoplolatilus randalli n. sp. è descritto sulla base di 13 esem- & Kuiter, 1989, from eastern Indonesia. They also plari di 46.7-158.2 mm SL, raccolti in diverse località quali discussed an additional species, H. geo Fricke & le Isole Banda e Papua occidentale in Indonesia, Luzon nelle Kacher, 1982, described from the Red Sea without Filippine settentrionali, Palau e Yap in Micronesia e le Isole Salomone. La nuova specie è stata in passato confusa con H. type specimens on the basis of photographs taken fronticinctus Günther, una specie affine ma confinata al - from a submersible. Coverage of the family was l’Oceano Indiano dall’Africa orientale al mar delle also provided by Dooley (1999), who included a Andamane orientale. Le due specie possono essere distinte synopsis of defining characters and an illustrated principalmente per la colorazione, prevalentemente verda - key to the species occurring in the western Pacific stra quella di H. randalli n. sp. contro una tinta bluastra di with the exception of H. luteus and H. pohle. Allen H. fronticinctus. Esse differiscono anche per la morfologia (2007) described another new species, H. erd- del lobo della pinna caudale; entrambe le specie hanno la manni, from West Papua, Indonesia. In addition, pinna caudale emarginata, ma il margine posteriore di ogni the range of H. pohle was recently expanded west- lobo è leggermente convesso in H. randalli n. sp. mentre quello in H. fronticinctus è retto. In aggiunta, H. randalli n. ward to Triton Bay, West Papua (Allen & Erdmann sp. ha spine dorsali più allungate (ultima spina 2.5-3.0 vs. 2009). Dooley & Jimenez (2008) considered H. 3.5-4.1 in HL) e raggi dorsali molli più lunghi (raggio più luteus as a junior synonym of H. fourmanoiri, but lungo pari a 1.1-1.4 vs. 1.5-1.7 in HL). Sono anche forniti we herein provide morphological, genetic and dati genetici che confermano la validità della nuova specie. colour differences which support these as two sep- In aggiunta, includiamo numerose prove (morfologiche, arate, albeit closely related, species. genetiche e differenze ontogenetiche della colorazione) che The present paper describes the thirteenth known portano a concludere che H. luteus, recentemente conside- member of the genus, a species previously referred rata sinonimo di H. fourmanoiri, sia una specie valida. to as H. fronticinctus. Randall & Dooley (1974) indicated this species was known only on the basis INTRODUCTION of the holotype from Mauritius and an additional The Indo-Pacfic genus Hoplolatilus belongs to the specimen from Madras, India, lodged at the Aus- circumtropical family Malacanthidae and contains tralian Museum, although they provisionally iden- elongate, cigar-shaped fishes that occur over silt, tified South Pacific postlarval Hoplolatilus from sand, or rubble bottoms at depths to at least 85 m. Alepisaurus and Thunnus stomachs as H. fronticinc- The genus, although containing 12 species, has tus. Randall (1981) identified specimens of the largely escaped attention due to its relatively deep new taxon from the Solomon Islands, Palau, and dwelling habits and proclivity for open rubble/sand Philippines as H. fronticinctus. substrates that are largely avoided by divers. Diving colleague and British videographer Nick Randall & Dooley (1974) reviewed the genus, Hope captured video images of what we first con- recognizing five species (general distribution indi- cluded was a new species of Hoplolatilus while div- cated in parentheses): H. cuniculus Randall & ing at the Mergui Archipelago in the East Dooley, 1974 (widespread Indo-west and central Andaman Sea off Myanmar during early 2008. He Pacific), H. fourmanoiri Smith, 1963 (Vietnam to later provided detailed locality information and Solomon Islands), H. fronticinctus (Günther, 1887) shortly afterwards we were presented with an (widespread Indian Ocean), H. oreni Clark & Ben- opportunity to spend several days of diving at Tuvia, 1973 (Red Sea) and H. starcki Randall & Myanmar during mid-February 2009. We suc- Dooley, 1974 (widespread western and central ceeded in collecting three specimens at 56 m Pacific). Randall later (1981) recognised three depth, utilising SCUBA diving equipment. One additional species: H. chlupatyi Klausewitz, year later we had another opportunity to observe McCosker, Randall & Zetzsche, 1978 (Philip- and collect this species at the Andaman Islands. pines), H. marcosi Burgess, 1978 (Philippines and Subsequent investigation revealed that the Myan- Indonesia to Solomon Islands) and H. purpureus mar and Andaman Islands fish were identical to the Burgess, 1978 (Philippines and Indonesia to species originally described as H. fronticinctus and Solomon Islands). More recently, the genus was the closely related species from the western Pacific reviewed by Earle & Pyle (1997), who described an is actually a new taxon, which is described herein. additional species, H. pohle, from south-eastern Papua New Guinea. They provided a table com- MATERIALS AND METHODS paring important diagnostic features for the nine Lengths of specimens are given as standard length aqua vol. 16 no. 4 - 15 October 2010 172 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton

(SL) measured from the anterior edge of the upper States National Museum, Washington, D.C. lip to the base of the caudal fin (posterior edge of (USNM), and Western Australian Museum, Perth hypural plate); head length (HL) is measured from (WAM). We also examined the holotype of Hoplo- the same anterior point to the posterior edge of the latilus fronticinctus at the Natural History opercular flap; head depth is measured at the level Museum, London (BMNH). of the posterior margin of the preopercle; cheek We also conducted a genetic analysis of the new depth is measured vertically from the lower rim of species and its relatives. Samples from 21 individu- the orbit to the lower margin of the preoperculum; als representing five species of Hoplolatilus (H. erd- opercular length is measured from the posterior manni, H. fourmanoiri, H. fronticinctus, H. luteus margin of the preoperculum horizontally to the tip and the new taxon) were included in this study. In of the opercular spine; suborbital depth is mea- addition to the data collected for this project, we sured vertically from the lower rim of the orbit to also included previously published (Dooley & the ventral edge of the head; body depth is the Jimenez 2008) sequence data from the16s riboso- maximum depth measured vertically between the mal RNA region of the mitochondrial genome for pelvic fin base and the base of the dorsal spines; H. fourmanoiri (n=1) and H. luteus (n=1) in our body width is the maximum width just posterior to analysis. the gill opening; snout length is measured from the Genomic DNA was isolated from muscle tissues anterior edge of the upper lip to the anterior edge or fin clips using 10% Chelex (Walsh et al. 1991). of the eye; orbit diameter is the horizontal fleshy Double-stranded polymerase chain reaction (PCR) diameter of the eye and interorbital width the least was used to amplify 598 base pairs (bp) of the 16s fleshy width; upper jaw length is taken from the rRNA region of the mitochondrion using primers front of the upper lip to the posterior end of the 16sAR and 16sBR (Palumbi et al. 1991) following maxilla; caudal peduncle depth is the least depth protocols described in Drew et al. (2008). Double- and caudal peduncle length is the horizontal dis- stranded cycle-sequencing was carried out for each tance between verticals at the rear base of the anal amplicon using a BigDye Terminator cycle- fin and the caudal fin base; caudal fin length is the sequencing kit v3.1 (Applied Biosystems) and horizontal length from the posterior edge of the sequences visualized with an Automated 3730 hypural plate to a vertical at the tip of the longest DNA Analyzer (Applied Biosystems) at the Life ray; pectoral fin length is the length of the longest Sciences Laboratory Core Center at Cornell Uni- ray; pelvic fin length is measured from the base of versity. Forward and reverse sequences were proof- the pelvic spine to the tip of the longest soft ray; read using Sequencher 4.7 (GeneCodes Corp), only the pored scales are counted in the lateral line visually checked for accuracy, then aligned in between the upper edge of the operculum and the MUSCLE (Edgar 2004). No hypervariable regions hypural crease (excludes 2-3 pored scales on the were found in the aligned dataset. caudal fin base); scales in a lateral series above the FindModel (http://hcv.lanl.gov/content/sequence/ lateral line are counted immediately above the lat- findmodel/findmodel.html), an online version of eral line between the upper edge of the operculum jModelTest (Posada 2008) was used to identify the and caudal-fin base; gill raker counts are presented best-fit model of nucleotide substitution. Find- as separate counts for the upper and lower limbs as Model calculates AIC scores using the method of well as a combined count; and the last fin ray ele- Posada & Crandall (2001); AIC ranked the TMV ment of the dorsal and anal fins is branched near (Transversion Model plus Gamma) substitution the base and is counted as a single ray. model as having the best fit to the data (Posada & Counts and proportions appearing in parentheses Buckley 2004). Under this model, Maximum Like- apply to the paratype if different from the holo- lihood (ML) heuristic searches were conducted type. Vertebral counts were obtained from radi- using the hill-climbing algorithm in the program ographs. Proportional measurements of selected GARLI v.0.96 (Zwickl 2006) with a random start- type specimens, expressed as percentage of the ing tree and default settings for the genetic algo- standard length, are provided in Table 1. Type rithm. Identical topologies and nearly identical specimens are deposited at the Australian Museum, likelihood scores were obtained for five separate Sydney (AMS), Bishop Museum, Honolulu GARLI runs and 1,000 bootstrap pseudoreplicates (BPBM), Pusat Penelitian dan Pengembangan were used to assess support for the resulting ML Oseanologi, Jakarta, Indonesia (NCIP), United topology.

173 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus Hoplolatilus randalli n. sp. Malu Group, Solomon Islands, 60 m, spear, W. (Figs 1-2; Table I) Starck, 26 March 1977; BPBM 40832, 133.0, Fais Randall’s Tilefish Island, 9°45’57.56”N, 140°30’34.81”E, Yap, Car- oline Islands, 85 m, hand net, B. Greene, 14 April Holotype: NCIP 6354, 154.1 mm SL Boo Kecil, 2007; NCIP 6355, 2 specimens, 109.1-131.5 mm 2°13.393’S, 130°34.687’E, south-eastern Misool, SL, same collecting data as AMS paratype; USNM Raja Ampat Islands, West Papua, Indonesia, 60 m, 396014, 152.4 mm SL, same collecting data as spear, M. Erdmann, 24 March 2009. holotype; WAM P.33094-001, 142.8 mm SL, same Paratypes: AMS I.45065-001, 134.9 mm SL, Batu collecting data as holotype; WAM P.33083-001, Kapal, 4°29.578’S, 129°55.932’E, Banda Islands, 154.3 mm SL, same collecting data as AMS Indonesia, 64 m, spear, M. Erdmann, 16 March paratype. 2009; BPBM 12454, 46.7 mm SL, Augulpelu Diagnosis: Dorsal rays X,13; anal rays II,12; pec- Reef, 7°16.316’N, 134°31.777’E, Palau Islands, 70 toral rays 17; total gill rakers on first branchial arch m, spear, W. Starck, 4 March 1972; BPBM 21109, 27-29; pored lateral line scales 84-92; scales in lat- 3 specimens, 142.3-158.2 mm SL, north side of eral series above lateral line 98-120; preopercular Maricaban Island, 13°40.367’N, 120°51.030’E, serrae 16-23; greatest body depth 3.4-3.9 in SL; Luzon, Philippines, 30 m, rotenone and spear, J. length of last dorsal spine 2.5-3.0 in HL; longest Randall, W. Einziger and K. Carpenter, 2 Septem- soft dorsal ray 1.1-1.4 in HL; colour in life mainly ber 1977; BPBM 26455, 140.0 mm SL, Malau- greenish (darker dorsally), shading to light blue on paina Island, 10°15.598’S, 161°57.642’E, Olu lower head, thorax, and abdomen; elongate sky-

Fig. 1. Underwater photographs of Hoplolatilus randalli n. sp. (upper), approximately 150 mm SL, Triton Bay, West Papua, Indonesia and H. fronticinctus (lower), approximately 150 mm SL, Campbell Shoal, Andaman Islands. Photos by G. R. Allen. aqua vol. 16 no. 4 - 15 October 2010 174 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton blue saddle-like spot dorsally on caudal peduncle; origin of anal fin 37 (37-38); vertebrae 10 + 14. narrow sky-blue band from front of snout to lower Body moderately elongate, compressed, its great- edge of eye; lips mainly blue; narrow yellow-orange est depth 3.7 (3.4-3.9) in SL; greatest body width band just below blue band, passing just under eye 1.8 (1.7-2.0) in greatest depth; caudal peduncle to posterior margin of preoperculum; dorsal and depth 2.3 (2.1-2.3), caudal peduncle length 1.7 anal fins pale yellow with pink margin; caudal fin (1.5-1.9), both in HL. Head blunt, its length 3.8 reddish with indistinct central pale streak, the (3.6-3.9) in SL; head depth 1.2 (1.0-1.3) in HL; upper and lower margins narrowly light blue to snout length 3.6 (3.3-4.3) in HL; upper jaw length pinkish; pectoral fins mainly translucent except 2.2 (2.1-2.4) in HL; cheek depth 3.8 (3.2-4.7) in upper 4-5 rays blue; pelvic fins bluish white; iris HL; opercular length 3.0 (2.9-3.6) in HL; snout to yellow with bright blue streak dorsally and ven- vertical margin of preopercle 1.4 (1.3-1.5) in HL; trally. orbit diameter 4.2 (3.6-4.5) in HL; suborbital Description: Dorsal rays X,13; anal rays II,12; depth 3.6 (2.8-5.0) in HL; fleshy interorbital pectoral rays 17; principal caudal fin rays 17; gill width 2.4 (2.3-2.8) in HL. rakers 10 + 19 = 29 (8-10 + 18-20 = 27-29); pored Mouth inferior, oblique, extending ventroposteri- lateral line scales 91 (84-92); scales in lateral series orly at approximately 26 (22-29) degree angle above lateral line 115 (98-120); transverse cheek below horizontal axis of body; maxilla reaching scale rows 13 (13-14); transverse opercular scale vertical to about posterior edge of orbit; front of rows 9 (9-11); scales above lateral line to origin of upper jaw with enlarged, curved canine on each dorsal fin 16 (13-17); scales below lateral line to side of symphysis; 3-4 irregular rows of tiny villi-

Fig. 2. Comparisons of freshly collected specimens of Hoplolatilus randalli n. sp. (upper, NCIP 6354, holotype, 154.1 mm SL), Misool, West Papua, Indonesia and H. fronticinctus (lower), 114.3 mm SL, Mergui Archipelago, Myanmar. Note – unusual head and pectoral-fin orientation of H. fronticinctus is due to spear wound on opposite side of body. Photo by G. R. Allen.

175 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus

Table I. Proportional measurements for selected type specimens of Hoplolatilus randalli expressed as percentage of the standard length.

Holotype Paratype Paratype Paratype Paratype Paratype NCIP 6354 BPBM 21109 BPBM 21109 WAM P.33094 AMS I.45065 NCIP 6355 Standard length 154.1 158.2 147.4 142.8 134.9 109.1 Greatest body depth 26.7 29.0 26.8 26.3 25.4 28.5 Greatest body width 14.9 14.7 14.7 15.3 14.1 15.9 Head length 26.4 27.1 27.0 26.1 26.8 25.9 Head depth 22.3 22.8 22.5 21.4 25.9 23.6 Opercular length 8.9 8.2 8.1 8.4 7.3 7.8 Snout to preopercle 19.0 18.9 18.5 18.1 19.0 19.2 Snout length 7.3 6.6 7.1 7.4 7.1 7.8 Orbit diameter 6.3 6.6 6.3 6.5 6.9 7.1 Interorbital width (fleshy) 10.8 9.6 9.9 9.9 9.6 11.4 Upper jaw length 12.2 11.8 11.4 12.5 12.1 12.3 Suborbital depth 7.4 7.1 6.2 7.2 6.1 7.1 Cheek depth 6.9 6.4 7.3 7.8 6.8 8.2 Caudal peduncle depth 11.6 12.4 12.1 11.3 11.8 12.6 Caudal peduncle length 143.7 170.4 150.0 150.0 145.0 147.6 Predorsal length 30.5 29.8 30.8 30.4 30.6 30.6 Snout to origin of anal fin 57.1 57.8 57.3 58.3 57.5 57.3 Snout to origin of pelvic fin 30.6 29.2 29.9 29.7 27.0 30.7 First dorsal spine length 2.1 2.8 2.2 2.5 1.8 2.7 Fourth dorsal spine length 5.8 6.3 6.0 6.2 6.4 7.7 Last dorsal spine length 7.5 8.2 8.0 8.1 8.5 9.3 Longest dorsal soft ray 17.7 18.6 17.4 18.5 19.0 17.8 Dorsal fin base length 56.5 54.0 57.2 56.4 56.6 57.8 First anal spine length 1.9 2.2 2.2 2.0 2.0 2.7 Second anal spine length 2.7 4.6 3.9 3.6 4.3 4.1 Longest anal soft ray 12.5 14.5 12.1 14.1 12.5 13.4 Anal base length 28.0 29.6 29.0 30.1 28.5 28.1 Pectoral fin length 24.6 25.3 25.1 24.9 25.1 23.9 Pectoral fin – upper ray 9.4 9.0 9.0 9.0 9.0 9.3 Pelvic fin length 13.4 14.9 14.6 14.8 14.9 15.0 Pelvic spine length 7.8 9.2 8.2 7.4 8.5 10.4 Caudal fin length 17.0 23.4 18.5 17.4 20.3 22.9 Caudal concavity 3.9 6.6 4.3 3.9 4.1 6.5

form teeth at front of jaws, posterior to canines, (16-23) serrae, including enlarged spine at preop- tapering to single row on side of jaw; posteriormost ercular angle; opercular spine flat, broad-based, tooth much enlarged and directed anteriorly; lower slight double curvature, roughly forming equilat- jaw with 16-18 enlarged conical teeth in outer row eral triangle with concave sides and thickened cen- of anterior portion of jaw and 3-4 rows of tiny vil- tre, less than pupil diameter in length, not extend- liform teeth behind tapering to single row posteri- ing beyond opercular membrane. Scales extending orly with posteriormost tooth enlarged and anteriorly on head to level of about midway directed anteriorly; palatine, vomer, and tongue between posterior rim of orbit and preopercular edentate; pronounced wart-like projection of white margin (extending nearly to posterior rim of orbit skin on inner edge of clavicle under operculum in largest paratype); scales generally ctenoid, except near pectoral fin base. mostly cycloid and smaller on head region; most of Lateral line pores in low arching profile; pores of caudal fin scaled, remaining fins naked except for cephalic system clearly visible, arranged as illus- small scales on pectoral fin base. trated for H. fronticinctus by Randall & Dooley Dorsal fin nearly uniform in height except for (1974), including 4 pores on each dentary, 5 pores lower anterior spinous portion; base of dorsal fin on preopercular margin, 8 pores in area immedi- 1.8 (1.7-1.9) in SL; origin of dorsal fin over upper ately above preoperculum-operculum, 9 circumor- pectoral fin base; predorsal length 3.3 (3.2-3.5) in bital pores, and 4 supraorbital-snout pores; median SL; dorsal spines thin, short, increasing in length; interorbital pore absent. Preoperculum with 17 first spine less than half of fourth spine; last dorsal

aqua vol. 16 no. 4 - 15 October 2010 176 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton spine 3.0 (2.5-3.0) in HL; soft portion of dorsal fin SL; pelvic fin spine slightly more than one-half nearly uniform in height; first soft dorsal ray length of longest pelvic ray; all pelvic-fin rays slightly ahead of level of anal fin origin; all soft dor- branched. Caudal fin emarginate with posterior sal rays branched, becoming increasingly branched margin of each fin lobe slightly convex, its length posteriorly, last ray branched at base; penultimate 5.9 (4.3-5.7) in SL; upper and lowermost principal soft dorsal ray notably longer than adjacent rays, caudal rays unbranched, remaining principal rays length 1.3 (1.1-1.4) in HL. Anal fin nearly uni- branched. form in height, rays slightly shorter than dorsal fin Colour in life from digital photographs (Fig. rays; base of anal fin 3.6 (3.3-3.6) in SL; origin of 1, holotype, upper): upper half of head and most of fin below base of first or second dorsal soft ray; two body greenish (darker on back), shading to light short anal spines, the first 1.4 (1.5-2.1) in length of blue on lower head, thorax, and abdomen; elongate second spine; all soft anal rays branched, becoming sky-blue saddle-like spot dorsally on caudal pedun- increasingly branched posteriorly, last ray branched cle; narrow sky-blue band from front of snout to at base; penultimate soft anal ray notably longer lower edge of eye; lips mainly blue; narrow yellow- than adjacent rays, length 2.1 (1.7-2.2) in HL. Pec- orange band just below blue band, passing under toral fins pointed, reaching a vertical at base of last eye to posterior margin of preoperculum; dorsal few dorsal spines; length of longest pectoral ray 4.1 and anal fins pale yellow with pink margin; caudal (3.9-4.2) in SL; all but uppermost and lowermost fin reddish with indistinct central pale streak, upper pectoral rays branched; stout upper ray about one- and lower margins of fin narrowly light blue to third length of longest pectoral ray. Pelvic fins pinkish; pectoral fins mainly translucent except more or less pointed, their origin slightly anterior upper 4-5 rays blue; pelvic fins bluish white; iris yel- to lower pectoral-fin base, length 7.4 (6.5-7.5) in low with bright blue streak dorsally and ventrally.

Fig. 3. Underwater photograph of Hoplolatilus erdmanni, approximately 160 mm TL, Triton Bay, Irian Jaya Barat Province, Indonesia. Photo by G. R. Allen.

177 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus

Table II. Pairwise genetic distances (for the 16s ribsomal RNA gene region of the mitochondrion) among Hopolatilus species included in this study.

H. erdmanni H. fourmanoiri H. fronticinctus H. luteus H. luteus (Bali) H. randalli H. erdmanni .00000 H. fourmanoiri .12105 .00000 H. fronticinctus .04866 .11939 .00000 H. luteus .12105 .00503 .11938 .00000 H. luteus (Bali) .11937 .01005 .11941 .01508 .00000 H. randalli .04866 .11599 .01678 .11598 .11432 .00000

Colour when fresh (Fig. 2, upper): head chlupatyi, H. erdmanni, H. fronticinctus, H. geo and and body generally bluish grey; elongate dark grey H. pohle. This complex is characterised by rela- saddle-like spot dorsally on caudal peduncle; nar- tively few soft dorsal and anal rays (usually 13 and row dark blue band from front of snout to lower 12 respectively), generally fewer lateral-line scales edge of eye; narrow yellowish band below blue (81-97), fewer preopercular serrae (16-23) and a band, passing just under eye to posterior margin of relatively deep body (3.4-4.1 in SL, except 5.1-5.6 preoperculum; dorsal and anal fins yellowish, more in H. chlupatyi). Members of this group are also vivid on spinous portion of dorsal fin; caudal fin known to construct impressively large mounds at dusky dark grey on upper lobe, dusky yellow the entrance of their burrows, sometimes brown on lower lobe with large central translucent approaching 1 m in height and 2-3 m in diameter. streak; pectoral fins mainly translucent to slightly Hoplolatilus geo is provisionally placed in the group yellowish except upper 4-5 rays dark blue; pelvic on the basis of photographic evidence (no speci- fins bluish white; iris yellow with bright blue streak mens have been collected). dorsally and ventrally. Morphologically H. randalli is most similar to H. Colour in alcohol: recently preserved (less fronticinctus (Figs 1, 2, lower) and H. erdmanni than one year) specimens with head and body grey- (Fig. 3). It differs from H. erdmanni in having ish brown (darker dorsally); elongate dark-brown more lateral-line scales (84-92, modally 88, versus saddle-like spot dorsally on caudal peduncle; lips, 76-84) and also differs in colouration. Most cheek, lower operculum, thorax, and abdomen noticeably it lacks the distinctive pattern of bars on dark grey; narrow dark grey band from front of the side of the body and the large red area that cov- snout to lower edge of eye; tan band below dark ers the central portion of the caudal fin, both typ- grey band, passing under eye to posterior margin of ical of H. erdmanni. Another difference is the pos- preoperculum; dorsal fin brown, darker along base torbital orientation (horizontal in H. randalli and of spinous portion and with narrow whitish mar- oblique in H. erdmanni) of the light-coloured gin; anal fin translucent with narrow whitish mar- stripe that passes below the eye. In addition, the gin; caudal fin dusky brown, darker on upper lobe, blue region on the caudal peduncle forms a distinct with large central translucent streak; pectoral fins dorsal saddle in H. randalli, but in H. erdmanni it translucent to slightly yellowish except upper 4-5 spans the entire tail base. Finally, H. randalli has a rays dark grey to nearly blackish; pelvic fins relatively pale red colouration on the caudal fin translucent; iris yellowish with grey streak dorsally instead of the vivid red colour of H. erdmanni. and ventrally. Older preserved (more than 30 The new species is most closely related to H. fron- years) specimens generally light yellowish brown ticinctus from the Indian Ocean. Although known on head and body; elongate dark-brown saddle-like from only a few localities including Mauritius spot dorsally on caudal peduncle; faint indication (type locality), Maldives (Kuiter 1998, as Hoplo- of dark snout band and light band immediately latilus sp. 1), India (Madras and Andaman Islands) below as described above; fins generally pale yel- and Myanmar (Mergui Archipelago), H. fronticinc- lowish tan to translucent. tus is no doubt more widespread in the region, but Remarks: Allen (2007) reviewed previous work remains unnoticed due to its relatively deep habi- dealing with the genus Hoplolatilus. The new tat. The two species appear to provide yet another species belongs to a group of species comprising H. example of an Indo-Pacific geminate species pair.

aqua vol. 16 no. 4 - 15 October 2010 178 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton

Randall (1998) discussed this phenomenon in blackish submarginal band that is relatively narrow detail, providing numerous examples of such pairs on the spinous portion of the fin, becoming involving 17 families. broader posteriorly (occupies about half of the fin Hoplolatilus randalli and H. fronticinctus have width on the posterior part of the soft dorsal). The similar meristic features including counts for fin anal fin of freshly collected H. fronticinctus is yel- rays, gill rakers, scales, and preopercular serrae. low with a bright pink margin and pair of darker However, there are basic colour pattern differences, blue bands on the basal half of the fin (this feature especially in large adults (compare photographs in not evident in underwater photos, but can be seen Fig. 1). The ground colour of H. randalli is notice- in Fig. 2, lower), compared to the uniform pale yel- ably greenish compared with the predominately low fin of H. randalli. The caudal fin of H. randalli blue colour of H. fronticinctus. The blue coloura- is reddish or reddish brown with a slightly paler tion of the latter species is restricted to the upper lower lobe and large central pale streak in compar- half of the body, while the lower half is uniformly ison to that of H. fronticinctus, which features a and strikingly white. In contrast, the greenish more or less uniform bluish to dusky brown caudal colour of H. randalli, although paler on the lower fin with 3-5 smaller pale streaks (compare pho- side, extends to about the level of the lower edge of tographs in Fig. 2). Moreover, the shape of the pos- the pectoral fin base and, instead of being white as terior margin of each caudal fin lobe differs: in H. fronticinctus, is vivid blue (on lower head) to slightly convex in H. randalli and straight in H. pale blue (on ventralmost part of body). There is fronticinctus. Furthermore, H. randalli differs in also a difference in the colour of the median fins. having longer dorsal spines (last spine 2.5-3.0 vs. The dorsal fin of H. randalli is mainly yellowish 3.5-4.1 in HL), and longer soft dorsal rays (longest compared to the bluish-grey dorsal of H. fron- ray 1.1-1.4 vs 1.5-1.7 in HL). ticinctus, which typically has a dark brown to Comparative material for H. fronticinctus con-

Fig. 4. The single best maximum likelihood topology generated from 598bp of the 16s ribosomal RNA region of mtDNA sequence data for five species of Hoplolatilus. Numbers at the nodes indicate bootstrap support for 1,000 maximum likelihood pseudoreplicates. *Bali indicates the H. luteus sequence data published by Dooley & Jimenez (2008); **Bali indicates the H. fourmanoiri sequence data published by Dooley & Jimenez (2008).

179 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus sisted of three specimens, 72.6-114.3 mm SL, from between these two species and H. erdmanni is the Mergui Archipelago, Myanmar (WAM 4.9% uncorrected sequence divergence (Table II). P.33072-001), eight specimens, 137.2-159.6 mm Although these levels of divergence appear low, 16s SL, from the Andaman Islands (WAM P.33247- ribosomal RNA is highly conserved due to its func- 001 and P.33248-001), and the holotype, 159.4 tional and structural roles and this gene is com- mm SL, of Latilus fronticinctus from Mauritius monly used to infer relationships at deeper phylo- (BMNH 1886.2.5.8). Although more than 120 genetic levels (Weisburg et al. 1991), although it years old, the holotype still retains a trace of the has been used successfully for species delimitation diagnostic dark band on the dorsal fin. in other fish species (Aoyama et al. 2001; Craig et Analysis of 598bp of 16s ribosomal RNA from 21 al. 2001). The 1.7% uncorrected sequence diver- samples representing five species of Hoplolatilus gence between H. fronticinctus and H. randalli is confirmed that H. randalli is distinct from H. fron- congruent with interspecific differences calculated ticinctus and H. erdmanni. Monophyly of each of from mtDNA 16s rRNA reported in Anguillidae these taxa is well-supported in our maximum like- (0.4%; Aoyama et al. 2000) and Mugilidae (1.3%; lihood tree, with 100% bootstrap support for H. Papasotiropoulos et al. 2007). There is 12% uncor- randalli and H. erdmanni and 98% bootstrap sup- rected sequence divergence between the mono- port for H. fronticinctus (Fig. 4). These three phyletic clade consisting of H. randalli, H. fron- species form a well-supported (100% bootstrap ticinctus and H. erdmanni and the monophyletic support) clade, with H. fronticinctus recovered as clade containing H. luteus and H. fourmanoiri. the sister taxon to H. randalli (Fig. 4). A single 16s These data support the recognition of H. randalli rRNA haplotype was recovered for each of these as a distinct species that is most closely related to three species. Uncorrected interspecific sequence H. fronticinctus. divergence between the sister taxa H. fronticinctus Specimens of Hoplolatilus randalli have been col- and H. randalli is 1.7% and interspecific variation lected from relatively few scattered locations in the

Fig. 5. Underwater photograph of Hoplolatilus fourmanoiri, adult and juvenile, approximately 100 and 45 mm SL, Brunei. Photo by G. R. Allen. aqua vol. 16 no. 4 - 15 October 2010 180 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton western Pacific including the Banda Islands and the mound. The Triton Bay area (3°54.905’S, West Papua in Indonesia, Luzon, northern Philip- 133°59.291’E) of West Papua, Indonesia, provides ines, Palau and Yap in Micronesia, and the vast tracts of deep rubble habitat and is home to at Solomon Islands. Kuiter & Tonozuka (2001) illus- least seven species of Hoplolatilus: H. randalli, H. trated specimens from Bali and nearby Nusa erdmanni, H. cuniculus, H. luteus, H. marcosi, H. Penida and we have also observed this species or pohle, and H. purpureus. seen underwater photographs of it from other Etymology: The species is named randalli for Indonesian localities including Triton Bay and reknowned ichthyologist John (“Jack”) Randall in Cenderawasih Bay in West Papua, Alor and Lom- recognition of his numerous valuable contribu- bok. It will certainly be found in other areas with tions to our knowledge of Indo-Pacific fishes and more widespread use of mixed-gas closed circuit particularly his previous work on the genus Hoplo- SCUBA equipment and submersibles that allow latilus. investigation of deeper reef environments. The Comments on the validity of Hoplolatilus luteus: habitat consists of gentle rubble slopes at depths Dooley & Jimenez (2008) presented what they between 30 and at least 85 m. Like other large believed to be sound evidence for considering H. Hoplolatilus, including H. fronticinctus, the new luteus as a junior synonym of H. fourmanoiri. Their species constructs impressive rubble mounds over contention was based on molecular data (compar- its burrows, which range from about 50-70 cm in ing 16s ribosomal RNA sequence data from a sin- height and 1.5-2 m in diameter. Typically, the fish gle individual of H. fourmanoiri and a single indi- are seen hovering a short distance from their home vidual of H. luteus) and aquarium observations of mound, and we have seen up to six individuals specimens purchased from an aquarium dealer, using a single mound. When approached closely or who received the fishes from an exporter based in frightened by spear shots the fish quickly retreat Bali, Indonesia. Previous data on the natural his- into the burrow, which is located near the apex of tory of both species was extremely limited, essen-

Fig. 6. Freshly captured (anesthetized) specimen of Hoplolatilus luteus, adult, 112 mm SL, Andaman Islands. Photo by G. R. Allen.

181 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus tially based on a few scattered underwater observa- large dark spot above the operculum (Fig. 5). As tions of only a few individuals, and therefore cast- the juveniles grow, the dark patch becomes con- ing doubt on the validity of H. luteus. Dooley & fined to the caudal fin, while the dark opercular Jimenez (2008) suggested this species, which is spot becomes evident. While adults are highly vari- mainly yellow, represents the juvenile stage of H. able with regards to the pattern of yellow patches fourmanoiri, undergoing a dramatic colour change on the dorsal part of the head, nape and adjacent at about 90-105 mm SL. Their assertion was diffi- back, at no point in their life history do individu- cult to evaluate given the overall dearth of speci- als of H. fourmanoiri display an all-yellow phase. mens and in situ observations of these two species. By contrast, the young of H. luteus are entirely Fortunately, during the past 12 months (2009- yellow, with neither a dark caudal patch nor a dark 2010) we have made extensive underwater obser- opercular spot. As individuals mature, they develop vations involving more than 50 individuals of each a large blue-edged black spot above the opercle that of these two species, with good representation of is linked by an irregular blue stripe to the upper young and adult fish between about 40-115 mm rim of the eye (Fig. 6). Unlike H. fourmanoiri, the SL. We were also able to collect four additional ground colour of H. luteus remains completely yel- specimens (WAM P.33246-001) of H. luteus, 98.7- low throughout their ontogeny and they never 112.0 mm SL, from the Andaman Islands and six develop a dark caudal patch. additional specimens (WAM P.33132-001) of H. The contention of Dooley & Jimenez (2008) that fourmanoiri, 71.2-100.2 mm SL, from Brunei. H. luteus represents the juvenile phase of H. four- Based upon these in situ observations and morpho- manoiri is also untenable based on the gonadal logical and molecular analysis of our additional development of the specimens we have examined. specimens, we are able to demonstrate below con- All four of our H. luteus specimens are sexually sistent differences in ontogeny of colour pattern, mature individuals including two males, 98.7- genetic sequence data and scale morphology 112.2 mm SL, and two females, 101.1-101.2 mm between H. fourmanoiri and H. luteus, thereby SL. Similarly the six specimens of H. fourmanoiri showing conclusively that H. luteus should be con- were invariably sexually mature, including four sidered a valid species. males, 71.2-100.2 mm SL, and two females, 73.4- In their description of H. luteus, Allen & Kuiter 95.8 mm SL. (1989) concluded that important differences in In addition to examination of ontogenetic colour colour pattern readily separated this species from pattern changes and gonadal development of H. H. fourmanoiri. Though Dooley and Jimenez luteus and H. fourmanoiri, we re-analyzed the (2008) speculated that H. luteus simply represents genetic data presented by Dooley & Jimenez the juvenile stage of H. fourmanoiri, our underwa- (2008) in light of the additional sequence data we ter observations of six juveniles of approximately collected. Our analysis of 598bp of mitochondrial 40-75 mm SL show clearly that the colour pattern data (16s ribosomal RNA) recovers two well-sup- of young H. fourmanoiri is very similar overall to ported monophyletic lineages: H. luteus (four indi- that of adults, except that the dark patch on the viduals from the Andaman Islands) and H. four- caudal fin extends well forward onto the ventral manoiri (three individuals from Brunei). The pre- part of the body and juveniles generally lack the viously published 16s ribosomal RNA sequence data from a single individual of H. fourmanoiri from Bali (Dooley & Jimenez 2008) is the same mtDNA haplotype as the three H. fourmanoiri individuals from Brunei sequenced in this study (Fig. 4), and the monophyly of this taxon is highly supported in our maximum likelihood analysis (95% bootstrap support). Likewise, a maximum likelihood analysis provides strong support (96% bootstrap) for the monophyly of the four H. luteus sequenced in this study. In contrast, the H. luteus Fig. 7. Scales of Hoplolatilus fourmanoiri (left), 100.2 mm sequenced for this study (all four individuals share SL, and H. luteus (right), 101.2 mm SL. Photographed a single mtDNA haplotype) and the previously region is at centre of caudal peduncle along lateral line. published 16s ribosomal RNA mtDNA data for a aqua vol. 16 no. 4 - 15 October 2010 182 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton single H. luteus from Bali (Dooley & Jimenez intraspecific variation of H. fourmanoiri would be 2008) do not share the same mtDNA haplotype, 1.5%, a value comparable to divergences between nor do these five individuals form a monophyletic species in the remainder of this dataset and in other clade (Fig. 5). groups of fish (Aoyama et al. 2000; Papasotiropou- The uncorrected interspecific sequence diver- los et al. 2007). It is more likely that these data sug- gence between H. fourmanoiri and the four H. gest there is unrecognized diversity within this luteus sequenced in this study is 0.5% (Table 2), clade, suggesting that further study of the taxo - greater than the previously reported interspecific nomy in this group is required. divergence for other fish species (Aoyama et al. Dooley & Jimenez (2008) suggest that the “near 2000). Also significant is that a single mtDNA identity” of H. fourmanoiri and H. luteus at two haplotype (0% divergence) was recovered for each slowly evolving loci (the mtDNA region of 16s species in this study, despite geographic variation rRNA and the nuclear gene RAG-2) is evidence for in sampling localities. This supports the view that considering these two lineages as a single species. the variation between H. fourmanoiri and H. luteus The molecular data included in their study consists does not represent intraspecific variation, but of only a single individual from each of the two rather is likely reflective of species boundaries. species in question. To make appropriate inferences Additionally, the single H. luteus sequenced by regarding the recovered variation between these Dooley & Jimenez (2008) is more divergent from two species, this variation must be viewed in the the four H. luteus samples included in our study context of the variation that exists within each (1.5% uncorrected sequence divergence) than it is species. When the sequence divergence between from H. fourmanoiri (1.0% uncorrected sequence the two individuals in their study is compared with divergence). If we were to synonymize H. luteus the intraspecific variation within each species, their and H. fourmanoiri based on these data, the argument for synonymy based on sequence simi-

Fig. 8. Individual scales of Hoplolatilus fourmanoiri (left), 100.2 mm SL, and H. luteus (right), 101.2 mm SL. Drawing mod- ified from micro-photograph. Scales were taken from identical positions one row above lateral line at level of base of last soft dorsal ray.

183 aqua vol. 16 no. 4 - 15 October 2010 Hoplolatilus randalli, a n. sp. of sand tilefish (Pisces: Malacanthidae) from the tropical western Pacific with comments on the validity of H. luteus larility does not hold. Dooley and Jimenez (2008) larger, coarser scales than H. luteus. This is particu- also appear to argue that the high level of support larly notable on the posteriormost portion of the (100% bootstrap) for a sister taxa relationship body and caudal peduncle. Microscopic compari- between H. fourmanoiri and H. luteus is evidence son of two scales (Fig. 8), one from each species, of synonymy. The recovery of these two lineages as taken from identical body positions, clearly illus- sister taxa indicates that they shared their most trate the size difference as well as the difference in recent common ancestor with each other, and are ctenii structure, which are generally longer and more closely related to one another than to other more slender in H. fourmanoiri. species in the recovered phylogenetic tree. Simi- Finally, we note that contrary to the contentions larly, the authors stated that the high level of iden- of Dooley & Jimenez (2008), these two species do tity between the H. fourmanoiri sequence and the not show evidence of sympatry. Though both H. luteus sequence “were not seen in any other tile- species occupy similar habitats (silty sand bottoms fish species-pair” comparisons. As their dataset is in about 30-40 m depth), H. fourmanoiri is known comprised of a single individual of each species and only with certainty from the South China Sea no information regarding the geographic origin of region (type locality in Vietnam, Brunei as the selected samples or the degree of intraspecific reported in the current study and Luzon as variation occurring within these species, it is diffi- reported in Dooley & Jimenez (2008)), while H. cult to have confidence in inferences made from luteus has been recorded from Maumere Bay in these data beyond a close relationship between H. Flores (type locality in Allen & Kuiter (1989)), Tri- fourmanoiri and H. luteus. One explanation for the ton Bay and Cendrawasih Bay in West Papua higher genetic similarity between this species pair is (Allen & Erdmann 2009) and now from the that they result from a more recent speciation event. Andaman Islands. The two juvenile specimens Morphological differentiation between the two from Guadalcanal, Solomon Islands reported as H. species is more subtle than the colour and genetic fourmanoiri by Randall (1981) and discussed by differences we describe above. Allen & Kuiter Dooley & Jimenez (2008) must also be referred to (1989) initially suggested that H. fourmanoiri and H. luteus given their overall yellow colouration and H. luteus were closely related based on similar the lack of the large black caudal patch which is meristic and morphometric features, and close evident even in small juveniles of H. fourmanoiri comparison of the four specimens (WAM P.33246- (Fig. 5). 001) of H. luteus from the Andaman Islands and As for the specimens of both H. luteus and H. six specimens (WAM P.33132-001) of H. four- fourmanoiri imported from Bali (via the Marine manoiri from Brunei during the present study con- Center, Dallas, Texas) in Dooley & Jimenez firms their close relationship. Counts for dorsal fin (2008), we note that it is misleading and inaccurate rays (X or XI,21-23), anal fin rays (II,18-20), pec- to assign a collection location to specimens toral fin rays (15-17), gill rakers on the first obtained from aquarium dealers. Bali is one of the branchial arch (4-6 + 11-12), pored lateral line largest consolidation and transhipment centers for scales (95-106), scales in lateral series (115-130), marine ornamental fishes in southeast Asia, pro- predorsal scales (33-40) and preopercle serrae (29- cessing and exporting fishes that were collected 44) were generally similar and overlapping. The throughout Indonesia and neighbouring countries. only notable modal difference that was detected in It is therefore impossible to assign locality data to our relatively small sample size was the presence of the specimens reported in Dooley and Jimenez XI dorsal spines in three of four specimens of H. (2008), though it is likely that the H. luteus were luteus compared with a count of X in five of six collected from eastern Indonesia while the H. four- specimens of H. fourmanoiri. manoiri may have come from the Riau Archipelago We did, however, detect one important difference in the South China Sea, as both of these areas are while handling the specimens. Gently stroking the major collection centers for the exporters based in body from the tail towards the head (i.e. “against Bali (Erdmann, pers. obs.). the grain” of the scales) reveals that H. fourmanoiri Traditional approaches to taxonomy that rely has a rough texture similar to fine sandpaper in solely on morphological characters (such as colour contrast to H. luteus, which is relatively smooth to pattern or morphological traits associated with the touch. Closer comparison (Fig. 7) of similar- feeding) may underestimate the number of species sized individuals shows that H. fourmanoiri has if these characters result from convergent evolution aqua vol. 16 no. 4 - 15 October 2010 184 Gerald R. Allen, Mark V. Erdmann and Alison M. Hamilton

(Yang & Rannala 2010.) Recognition of this mar. We are also grateful to Dexter and Susan potential drawback and the need to identify cryp- Paine and their three children, Mercy, Sam, and tic species promoted the development of barcoding Honor, for inviting us to participate on a Silolona methods for species identification. DNA barcod- cruise to Banda and West Papua, Indonesia, during ing employs the use of a single gene region (typi- March 2009, a cruise which facilitated the collec- cally the mtDNA region CO1, although occasion- tion of the holotype and several paratypes of H. ally multiple gene regions or genes are sequenced) randalli. The Silolona’s excellent crew was especially to delimit species boundaries (Aliabadian et al. helpful, providing invaluable logistic support on 2009). Some studies have applied a threshold level both the Myanmar and Indonesian cruises. Jack of divergence (for example 11% for the cytb region Randall, one of the first collectors of the new of mtDNA) denoting species-level recognition species, generously sent his specimens for inclusion (Bradley & Baker 2001). An emerging approach in in the type series. We also thank Dr. Suharsono assessing species-level boundaries is more integra- and Rianta Pratiwi of the Indonesian Institute of tive and recognizes that morphological traits result Sciences (LIPI) for support and assistance with from multi-locus variation as opposed to the varia- curation of specimens, and Conservation Interna- tion quantified by analysis of a single gene region tional and the Indonesian Department of Nature and therefore may be more reflective of intra-spe- Conservation (PHKA) for continued sponsorship cific variation (Ezard et al. 2010). Additionally, of our marine biodiversity censuses in eastern many morphological characters may reflect adap- Indonesia. tive processes and thus may be more reflective of species boundaries if divergent selection drives speciation (Rundle & Nosil 2005). A combined REFERENCES approach to species delimitation that integrates ALLEN, G. R. 2007. Hoplolatilus erdmanni, a new species of sand tilefish (Pisces: Malacanthidae) from western molecular, morphological and ecological data, such New Guinea. Aqua, International Journal of Ichthyology as presented here, is a more robust approach to the 12 (3): 101-106. delimitation of species boundaries, especially as ALLEN, G. R. & ERDMANN, M.V. 2009. Reef fishes of the they identify processes that operate at different Bird’s Head Peninsula, West Papua, Indonesia. Check List evolutionary time-scales (Leache et al. 2009). Mea- 5 (3): 587-628. sures of differentiation must be viewed in context. ALLEN, G. R. & KUITER, R. H. 1989. 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