AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 1

aqua, International Journal of Ichthyology

Hoplolatilus andamanensis, a new species of sand (Pisces: Malacanthidae) from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

Gerald R. Allen1 and Mark V. Erdmann2,3

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected] 2) Conservation International Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235, Bali, Indonesia. Email: [email protected] 3) California Academy of Sciences, 55 Museum Concourse Drive, San Francisco, CA 94118, USA.

Received: 09 August 2018 – Accepted: 21 October 2018

Keywords die zwei Arten synonym sind. Die neue Art ist nahe ver- Ichthyology, , coral reef fishes, eastern Indian wandt mit H. fourmanoiri, unterscheidet sich aber durch Ocean. Schuppenmerkmale und Farbgebung. Ihre Vertreter be- sitzen gewöhnlich 11 Rückenflossendornen, während es Abstract bei H. fourmanoiri in der Regel nur 10 sind. Nach den Hoplolatilus andamanensis is described on the basis of Ergebnissen der 16s ribosomalen Sequenzanalyse beträgt four specimens, 99.2-110.4 mm SL, collected at the An- die Divergenz zwischen beiden Arten 1,5%. daman Islands in the eastern Indian Ocean. The species was previously misidentified as H. luteus, a species that was Sommario originally described from a single specimen from Flores, Hoplolatilus andamanensis è descritto sulla base di quattro Indonesia. Previous authors have debated the issue of esemplari di 99.2-110.4 mm SL, raccolti nelle isole An- whether H. luteus is a valid species or a junior synonym of damane nell'Oceano Indiano orientale. La nuova specie H. fourmanoiri. Additional evidence in the form of under- era precedentemente erroneamente identificata come H. water observations and collected specimens support the luteus, una specie originariamente descritta da un singolo contention of Dooley & Jimenez that the two species are esemplare proveniente da Flores, in Indonesia. Gli autori synonymous. The new species is closely related to H. four- precedenti hanno discusso la questione se H. luteus sia una manoiri, but differs in scale morphology and coloration. It specie valida o un sinonimo di H. fourmanoiri. Ulteriori also usually has 11 dorsal spines versus a usual count of 10 prove sotto forma di osservazioni subacquee e campioni spines in H. fourmanoiri. There is also a 1.5% divergence raccolti confermano la tesi di Dooley e Jimenez secondo between the two species based on 16s ribosomal RNA se- cui le due specie sono sinonimi. La nuova specie è stretta- quence data. mente imparentata con H. fourmanoiri, ma differisce nella morfologia delle scaglie e nella colorazione. Di solito ha Zusammenfassung anche 11 spine dorsali contro un comune conteggio di 10 spine in H. fourmanoiri. C'è anche una divergenza Hoplolatilus andamanensis wird auf der Grundlage von vi- dell'1,5% tra le due specie sulla base dei dati di sequenza er Exemplaren, 992-110,4 mm SL, beschrieben, die an dell'RNA ribosomiale 16s. den Andaman-Inseln im östlichen Indischen Ozean gefan- gen wurden. Die Art wurde bisher fälschlicherweise als H. luteus bestimmt, eine Art, die ursprünglich nach einem INTRODUCTION einzigen Exemplar von Flores, Indonesien, beschrieben The circumtropical family Malacanthidae, popu- worden war. Frühere Autoren haben die Frage diskutiert, ob H. luteus eine valide Art ist oder ein jüngeres Synonym larly known as , is divisible into two sub- von H. fourmanoiri. Zusätzliche Belege in Form von Un- families: Latilinae and Malacanthinae (Nelson terwasser-Beobachtungen und gesammelten Exemplaren 2006). The latter group contains two genera, unterstützen die Auffassung von Dooley & Jimenez, dass Malacanthus Cuvier and Hoplolatilus Günther,

1 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 2

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

which are common over sand and rubble bottoms hypural plate); head length (HL) is measured from in the vicinity of coral reefs. All of the known the same anterior point to the posterior edge of the species, except for M. plumieri (Bloch) of the At- opercular flap; head depth is measured at the level lantic, are distributed in the Indo-west and central of the posterior margin of the preopercle; cheek Pacific region. The Hoplolatilus largely es- depth is measured vertically from the lower rim of caped attention until the advent of scientific scuba the orbit to the lower margin of the preoperculum; diving due to its relatively deep dwelling habits. opercular length is measured from the posterior These long, slender fishes typically hover above margin of the preoperculum horizontally to the tip burrows, which they construct in soft substrates. of the opercular spine; suborbital depth is mea- There are currently 14 species of Hoplolatilus sured vertically from the lower rim of the orbit to recognised as valid (Eschmeyer et al. 2018): H. the ventral edge of the head; body depth is the chlupatyi Randall et al., 1978 (Philippines), H. cu- maximum depth measured vertically between the niculus Randall & Dooley, 1974 (widespread Indo- pelvic-fin base and the base of the dorsal spines; west and central Pacific), H. erdmanni Allen, 2007 body width is the maximum width just posterior to (West Papua, Indonesia), H. fourmanoiri Smith, the gill opening; snout length is measured from the 1963 (Vietnam to Solomon Islands), H. fronticinc- anterior edge of the upper lip to the anterior edge tus (Günther, 1887) (widespread Indian Ocean), of the eye; orbit diameter is the horizontal fleshy H. geo Fricke & Kacher, 1982 (Red Sea), H. luteus diameter of the eye, and interorbital width the least Allen & Kuiter, 1989 (eastern Indonesia), H. mar- fleshy width; upper jaw length is taken from the cosi Burgess, 1978 (Philippines and Indonesia to front of the upper lip to the posterior end of the Solomon Islands), H. oreni Clark & Ben-Tuvia, maxilla; caudal peduncle depth is the least depth, 1973 (Red Sea), H. pohle Earle & Pyle, 1997 (Bali, and caudal peduncle length is the horizontal dis- West Papua and Papua New Guinea), H. purpureus tance between verticals at the rear base of the anal Burgess, 1978 (Philippines and Indonesia to fin and the caudal fin base; caudal fin length is the Solomon Islands), H. randalli, Allen et al., 2010 horizontal length from the posterior edge of the (Philippines and Indonesia to Solomon Islands and hypural plate to a vertical at the tip of the longest eastern Micronesia), and H. starcki Randall ray; pectoral fin length is the length of the longest & Dooley, 1974 (widespread western and central ray; pelvic fin length is measured from the base of Pacific). the pelvic spine to the tip of the longest soft ray; The status of H. luteus is controversial. Dooley & only the pored scales are counted in the lateral line Jimenez (2008) suggested this species is a junior between the upper edge of the operculum and the synonym of H. fourmanoiri. However, Allen et al. hypural crease (excludes 2-3 pored scales on the (2010) provided morphological, genetic and caudal fin base); scales in a lateral series above the colour evidence to refute this conclusion, stating it lateral line are counted immediately above the lat- was unequivocally valid. Although H. luteus was eral line between the upper edge of the operculum originally described on the basis of a single speci- and caudal-fin base; gill raker counts are presented men from Flores, Indonesia, the comparative spec- as separate counts for the upper and lower limbs as imens utilized by Allen et al. (2010) were from the well as a combined count; the last fin ray element Andaman Islands. The present study reveals that of the dorsal and anal fins is branched near the base the Andamans material actually represents a new and is counted as a single ray. taxon described herein, and the previous conclu- Counts and proportions appearing in parentheses sion regarding the separate status of these species is apply to the paratype if different from the holo- invalid. We also provide supporting evidence in the type. Counts separated with a backslash (/) were form of underwater observations and several re- taken on both left and right sides of the same indi- cently collected specimens of H. fourmanoiri that vidual. Vertebral counts were obtained from radi- show the two putative species are definitely a single ographs. Proportional measurements of type speci- entity as first proposed by Dooley & Jimenez. mens, expressed as percentage of the standard length, are provided in Table I. Type specimens are MATERIALS AND METHODS deposited at the Western Australian Museum, Lengths of specimens are given as standard length Perth (WAM). (SL) measured from the anterior edge of the upper lip to the base of the caudal fin (posterior edge of

aqua vol. 25 no. 1 - 25 January 2019 2 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 3

Gerald R. Allen and Mark V. Erdmann Hoplolatilus andamanensis n. sp. 50-52 mm SL, Cenderawasih Bay, West Papua Andaman Tilefish Province, Indonesia H. fourmanoiri: P.33132-001, (Figs 1-2, 4 & 7A; Table I) 6 specimens, 70-97 mm SL, Brunei; P.33894-017, 4 specimens, 107-118 mm SL, Lembeh Strait, Holotype: WAM P.33246-001, 110.4 mm SL, North Sulawesi Province, Indonesia; H. fronticinc- “The Wall” dive site, approx. 12°03.297’N, tus: P. 33248-001, 6 specimens, 137–150 mm SL, 92°57.525’E, Havelock Island, Andaman Islands, Campbell Shoal, Andaman Islands; H. marcosi: P. 35-40 m, spear, M. Erdmann, 23 March 2010. 33359-001, 2 specimens, 78-84 mm SL, Cender- Paratypes: WAM P.33246-000, 3 specimens, 99.2- awasih Bay, West Papua Province, Indonesia; H. 100.6 mm SL, collected with holotype. pohle: P. 33070-002, 2 specimens, 130-137 mm Comparative specimens (all WAM): Hoplolatilus SL, Triton Bay, West Papua Province, Indonesia; cuniculus: P.34812-001, 2 specimens, both 85 mm H. purpureus: P. 33070-001, 99 mm SL, Triton SL, Lau Group, Fiji; H. erdmanni: P. 32860-012 Bay, West Papua Province, Indonesia; H. randalli: (paratype), 130 mm SL, Selat Iris, West Papua P. 33083-001 (paratype), 154 mm SL, Banda, Province, Indonesia; P.33051-002, 2 specimens, Maluku Province, Indonesia; P. 33094-001

Fig. 1. Hoplolatilus andamanensis n. sp., underwater photograph of freshly captured (anesthetised) holotype, 110.4 mm SL, Andaman Islands. Photo by G. R. Allen.

Fig. 2. Hoplolatilus andamanensis n. sp., preserved holotype, 110.4 mm SL, Andaman Islands. Photo by G. R. Allen.

3 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 4

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

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

Holotype Paratype Paratype Paratype WAM P.33246 WAM P.33246 WAM P.33246 WAM P.33246 male female female male

Standard length 110.4 100.6 99.8 99.2 Greatest body depth 18.1 20.0 20.8 19.6 Greatest body width 10.6 11.0 12.8 12.4 Head length 26.2 27.8 28.5 27.8 Head depth 16.0 16.6 17.2 16.7 Opercular length 8.9 8.5 8.7 8.4 Snout to preopercle 18.5 20.0 19.2 19.5 Snout length 6.1 6.6 6.6 6.5 Orbit diameter 7.1 7.6 7.4 7.3 Interorbital width (fleshy) 8.2 7.7 8.3 7.7 Upper jaw length 12.3 11.9 12.4 12.1 Suborbital depth 6.0 6.5 6.2 6.4 Cheek depth 5.7 5.6 5.5 6.2 Caudal peduncle depth 11.2 11.1 11.1 11.1 Caudal peduncle length 13.6 12.6 12.0 12.7 Predorsal length 28.4 31.3 31.0 30.3 Snout to origin of anal fin 54.2 54.0 53.0 54.6 Snout to origin of pelvic fin 24.2 24.8 25.7 26.0 First dorsal spine length 3.5 3.6 3.4 3.5 Fourth dorsal spine length 6.6 6.8 7.2 6.6 Last dorsal spine length 6.9 5.9 7.0 6.8 Longest dorsal soft ray 11.3 10.1 13.0 11.5 Dorsal-fin base length 62.0 61.0 64.2 62.7 First anal spine length 1.4 1.4 1.4 1.4 Second anal spine length 2.7 3.0 3.4 2.1 Longest anal soft ray 10.9 11.0 12.0 10.9 Anal-base length 33.0 32.8 33.2 32.6 Pectoral-fin length 21.4 22.2 23.4 22.4 Pectoral fin – upper ray 7.6 7.5 7.7 11.6 Pelvic-fin length 11.0 11.8 11.9 11.0 Pelvic-spine length 8.0 7.8 7.8 6.7 Caudal-fin length 18.6 19.1 20.7 19.4

(paratype), 143 mm SL, Misool, West Papua stripe; fins translucent, except central part of cau- Province, Indonesia; P. 33391-001, 72 mm SL, Ra- dal yellow, widest at base and tapering to middle of ja Ampat Islands, West Papua Province, Indonesia. posterior margin. Diagnosis: Dorsal rays X or XI,21-23; anal rays Description: Dorsal rays XI,21 (one paratype II,17-19; pectoral rays 15-16; total gill rakers on with X,23); anal rays II,17 (one paratype with first branchial arch 16-19; pored lateral-line scales II,18 and two with II,19); pectoral rays 15/16 (one 96-106; scales in lateral series above lateral line paratype each with 15 or 16 on both sides and one 119-135; preopercular serrae 32-40; greatest body with 15/16) ; branched caudal-fin rays 15; gill rak- depth 4.8-5.5 in SL; length of last dorsal spine 2.3- ers 5 + 11 = 16 (5-7 + 11-12 = 16-19); pored lat- 2.8 in HL; longest soft dorsal ray 1.3-1.6 in HL; eral-line scales 106 (96-101); scales in lateral series colour in life mainly yellow, except white on thorax above lateral line 119 (125-135); transverse cheek and lower head; prominent ocellus above opercu- scale rows 11 (9-12); transverse opercular scale lum, larger than pupil, consisting of black spot rows 10 (9-10); scales above lateral line to origin of completely surrounded by narrow, pale blue mar- dorsal fin 14 (13-14); scales below lateral line to gin, and linked to upper rim of eye by pale blue origin of anal fin 33 (31-34); vertebrae 10 +14.

aqua vol. 25 no. 1 - 25 January 2019 4 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 5

Gerald R. Allen and Mark V. Erdmann

Body moderately elongate, compressed, its great- margin of preopercle 1.4 (1.4-1.5) in HL; orbit di- est depth 5.5 (4.9-5.1) in SL; greatest body width ameter 3.7 (3.7-3.9) in HL; suborbital depth 4.3 1.7 (1.6-1.8) in greatest depth; caudal peduncle (4.3-4.6) in HL; fleshy interorbital width 3.2 (3.4- depth 2.3 (2.5-2.6), caudal peduncle length 1.9 3.6) in HL. (2.2-2.4), both in HL. Head blunt, its length 3.8 Mouth inferior, oblique, extending ventroposteri- (3.5-3.6) in SL; head depth 1.6 (1.7) in HL; snout orly at approximately 26° angle below horizontal length 4.3 (4.2-4.3) in HL; upper jaw length 2.1 axis of body; maxilla reaching vertical to nearly (2.3) in HL; cheek depth 4.6 (4.5-5.2) in HL; op- posterior edge of orbit; front of upper and lower ercular length 2.9 (3.3) in HL; snout to vertical jaws with small villiform teeth arranged in multi-

Figs 3A-B. Hoplolatilus fourmanoiri, underwater photographs: A. approx. 100 & 45 mm SL, Brunei. Photo by G. R. Allen. B. “luteus” phase, approx. 50 mm SL, Banda Sea, Indonesia. Photo by M. V. Erdmann.

5 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 6

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

ple, irregular rows, tapering to 1-2 rows posteriorly, scales extending anteriorly on head to level of teeth of outermost row much enlarged and stouter about posterior rim of orbit; scales finely ctenoid compared to inner teeth, including 5-7 teeth ante- (Fig. 4B), except mostly cycloid and smaller on riorly in upper jaw, and 6-7 progressively larger head region, adjacent body anterior to level of dor- teeth posteriorly in lower jaw, posteriormost pair sal-fin origin, and uppermost portion of back be- (situated on middle part of jaw) largest, canine-like low base of spinous portion of dorsal fin; basal half and slanted posteriorly; palatine, vomer, and of caudal fin scaled, remaining fins naked except tongue edentate; tongue rounded with papillate for small scales on pectoral-fin base. dorsal surface. Predorsal length 3.5 (3.2-3.3) in SL; origin of dor- Pored lateral-line scales in low arching profile on sal fin about level with posterior extent of opercular upper back, gradually descending to level slightly spine; base of dorsal fin 1.6 in SL; first dorsal spine above middle of side on caudal peduncle; pores of about two-thirds length of second spine, 7.4 (7.7- cephalic system clearly visible, including 5 pores on 8.3) in HL; dorsal spines then gradually increasing each dentary, 7 pores on preopercular margin, 8 in length to fourth or fifth spine, fourth spine 3.9 pores in area immediately above preoperculum-op- (4.0-4.2) in HL, remaining spines approximately erculum, 9 suborbital/posterior orbital pores, 4 subequal, last spine 3.8 (4.1-4.7) in HL; soft por- supraorbital-snout pores, and 2 pores below nos- tion of dorsal fin slightly taller than spinous portion trils; also single, median interorbital pore. Preoper- with rays approximately subequal, longest (third culum with 35/33 (27-41) serrae, including en- from last) dorsal soft ray 2.3 (2.2-2.7) in HL; all larged spine at preopercular angle; opercular spine soft dorsal rays branched except first, becoming in- flat, relatively broad-based and dagger-shaped, creasingly branched posteriorly, last ray branched at longer than pupil diameter in length, extending base. Anal fin more or less uniform in height, rays slightly beyond opercular membrane. Predorsal about equal to those of dorsal fin rays; base of anal

Fig. 4A-B. Individual scales of Hoplolatilus fourmanoiri (A), 100.2 mm SL, and H. andamanensis (B), 100.6 mm SL. Drawing from micro-photograph. Scales were taken from identical positions one row above lateral line at level of base of last soft dorsal ray.

aqua vol. 25 no. 1 - 25 January 2019 6 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 7

Gerald R. Allen and Mark V. Erdmann

fin 3.0 (3.0-3.1) in SL; origin of fin below base of above operculum, larger than pupil, consisting of fourth dorsal soft ray; two short anal spines, the first black spot completely surrounded by narrow, pale 1.9 (1.6-2.5) in length of second spine; all soft anal blue margin, and linked to upper rim of eye by pale rays branched except first, becoming increasingly blue stripe (sometimes slightly wavy with dorsal branched posteriorly, last ray branched at base; bulge or short side extension); iris of eye (Fig. 7A) longest (penultimate or third from last) soft anal ray mainly brown with blue marking on ventral sec- 2.4 (2.4-2.6) in HL. Pectoral fins pointed, reaching tion and larger yellow marking on anterior section, a vertical just beyond base of last dorsal spine; surrounding scleral rim mainly pale blue; fins length of longest pectoral ray 4.7 (4.3-4.5) in SL; all translucent, except central part of caudal yellow, but uppermost and lowermost pectoral rays the marking widest at base and tapering to middle branched; stout uppermost ray about one-third of posterior margin. length of longest pectoral ray. Pelvic fins short and Colour in alcohol (Fig. 2): Generally pale triangular, their origin slightly anterior to pectoral- tannish, except greyish on interorbital, front of fin base, length 9.1 (8.4-9.1) in SL; pelvic-fin spine snout, and predorsal; large black spot above oper- slightly less than one-half length of longest pelvic culum, linked to upper rim of eye by grey stripe ray; all pelvic-fin rays branched. Caudal fin slightly (sometimes slightly wavy with dorsal bulge or short rounded, its length 5.4 (4.8-5.2) in SL, tip of upper side extension); fins semi-translucent, except cau- lobe slightly protruding; all principal caudal rays dal-fin base tannish. branched. Comparisons: The new species is most closely re- Colour in life (Figs 1 & 7A): Mainly yellow, lated to H. fourmanoiri (Fig. 3) from the East Indi- except bluish grey on interorbital, front of snout, an region (Vietnam to Solomon Islands). It was er- and narrowly on predorsal midline, and white on roneously identified as H. luteus by Allen et al. thorax and lower head; prominent ocellus (Fig. 7A) (2010), which is now recognised as a synonym of

Fig. 5A-D. Hoplolatilus fourmanoiri, underwater photographs, Lembeh Strait, North Sulawesi Province, Indonesia: A) approx. 110 mm SL; B) approx. 50 & 80 mm SL; C) approx. 60 mm SL; and D) approx. 110 mm SL. Photos by M. V. Erdmann.

7 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 8

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

H. fourmanoiri as first suggested by Dooley & lateral-lines scales (92-106), relatively few gill rak- Jimenez (2008). The two species exhibit similar ers on the first arch (16-19), usually 11 precaudal meristic and morphological features and belong to vertebrae, and a slender body (4.9-6.3 in SL). The the complex that also includes H. oreni from the only notable meristic difference that was detected Red Sea. The group is characterised by a high num- in our relatively small sample size was the presence ber of soft dorsal and anal rays (21-23 and 18-20 of XI dorsal spines in three of four specimens of H. respectively), moderately high number of pored andamanensis compared with a count of X spines

Fig. 6. Hoplolatilus fourmanoiri, preserved specimens (WAM P.33894-017), 106.5-117.7 mm SL, Lembeh Strait, North Su- lawesi Province, Indonesia, showing typical adult coloration (upper pair) and transitional individuals. Photo by G. R. Allen.

aqua vol. 25 no. 1 - 25 January 2019 8 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 9

Gerald R. Allen and Mark V. Erdmann

Figs 7A-B. Comparison of head markings of Hoplolatilus: A. H. andamanensis, holotype, 110.4 mm SL. Photo by G. R. Allen; and B. H. fourmanoiri (“luteus” phase), 107.6 mm SL, Lembeh Strait, North Sulawesi. Photo by M. V. Erdmann.

9 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 10

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

in seven of 10 specimens of H. fourmanoiri. How- Dooley & Jimenez (2008) suggested that the yel- ever, there is an important difference in scale mor- low coloration of H. luteus, although different than phology. Gently stroking the body from the tail to- the mainly mauve pattern (with dark caudal patch) wards the head (i.e. “against the grain” of the of H. fourmanoiri, possibly represented the juvenile scales) reveals that H. fourmanoiri has a rough tex- stage of H. fourmanoiri, which then changed ture similar to fine sandpaper in contrast to H. an- colour at about 90-105 mm SL. However, based damanensis, which is relatively smooth to the on field observations of juvenile and adult H. four- touch. Closer comparison (Fig. 4) of similar-sized manoiri in the South China Sea (Brunei) and H. individuals shows that H. fourmanoiri has larger, luteus in eastern Indonesia, Allen et al. (2010) coarser scales than H. andamanesis. This discrepan- maintained that both H. fourmanoiri and H. luteus cy is particularly notable on the posteriormost por- retained their distinctive colour patterns through- tion of the body and caudal peduncle. Microscopic out their respective ontogenies. However, subse- comparison of two scales (Fig. 4), one from each quent observations of a large “colony” of over 60 species, taken from identical body positions, clearly individuals at Lembeh Strait, North Sulawesi by illustrate the size discrepancy as well as the differ- the authors in 2013 lend strong support to the syn- ence in ctenii structure, which are generally shorter onymy of H. luteus and H. fourmanoiri. In stark and stouter in H. andamanensis. Examination of contrast to our previous observations in Brunei, seven additional species of Hoplolatilus at WAM the Banda Sea and West Papua, in Lembeh Strait (see list of comparative specimens above), includ- the two putative species/colour morphs were seen ing H. chlupatyi, H. cuniculus, H. erdmanni, H. together cohabitating burrows, and moreover, we marcosi, H. pohli, H. purpureus, and H. randalli, in- observed several individuals with transitional pat- dicates scale morphology is consistent within each terns (Figs 5 & 6). Despite this finding we are still species, and moreover the scales of these species are unable to explain the seemingly random nature of generally “rougher” (due to stouter ctenii) than the colour variation, which does not appear to be those of H. andamanensis. correlated with ontogeny or sex. Moreover, we note

Fig. 8. The single best maximum likelihood topology generated from 598bp of the 16s ribosomal RNA region of mtDNA sequence data for five species of Hoplolatilus (from Allen et al. 2010). Numbers at the nodes indicate bootstrap support for 1,000 maximum likelihood pseudoreplicates. Bali specimens (* &**) are sequence data for H. fourmanoiri published by Doo- ley & Jimenez (2008).

aqua vol. 25 no. 1 - 25 January 2019 10 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 11

Gerald R. Allen and Mark V. Erdmann

that in our extensive field observations of sand tile- tion origin, it is also a well-documented centre for fish throughout the Coral Triangle region, only the the aquarium trade and Bali exporters accumulate Lembeh Strait population shows both colour specimens from numerous localities across the In- morphs of H. fourmanoiri; in the South China Sea donesian Archipelago. However Bali, and especial- we only observed the mottled mauve and yellow ly Sumatra cannot be discounted as the collection “fourmanoiri” phase with the dark caudal patch, locality. We have documented the occurrence of while in Bali, Flores, the Banda Sea, and West numerous Indian Ocean species at Bali and nearby Papua, only the yellow “luteus” phase was seen. Nusa Penida (Allen & Erdmann 2012). In addi- Further observations of this enigmatic species may tion, unpublished data collected by GRA reveals eventually reveal a discernible pattern of colour that the reef-fish fauna of Weh Island, off the phase occurrence, but our observations to date sug- northwestern tip of Sumatra is primarily composed gest only random variation. of Indian Ocean species with approximately 85% The new taxon is very similar in general appear- of its species shared with the Maldives. ance to the yellow “luteus” phase of H. fourmanoiri, The habitat of the new species consists of silty but there is an important difference in the config- sand bottoms in an area of periodic strong currents uration of the large dark spot above the operculum at 35-40 m depth. (Fig. 7) that is characteristic of adults. This distinc- tive marking is completely surrounded by a blue ACKNOWLEDGEMENTS margin in H. andamanensis, but only partially sur- We sincerely thank Dexter and Susan Paine and rounded (posteriorly) in H. fourmanoiri. Addition- their three children, Mercy, Sam, and Honor, for ally the blue-edged dark mark of H. andamanensis inviting us to participate on a cruise to the An- is connected to the upper rim of the eye by a blue daman Islands during March 2010, which facilitat- stripe, which is invariably absent in all colour phas- ed the collection of the type specimens of H. an- es of H. fourmanoiri. damanenis. Patti Seery, owner of M. Y. Silolona, Allen et al. (2010) also presented detailed DNA and her excellent crew were especially helpful, pro- evidence for the separation of the Andamans pop- viding invaluable logistic support during this jour- ulation of H. luteus (= H. andamanensis) and H. ney. We are also grateful to Bruce Moore, owner of fourmanoiri . Their analysis of 598bp of mitochon- the Black Sand Dive Retreat at Lembeh, North Su- drial data (16s ribosomal RNA) recovered two lawesi, and his excellent dive staff, for hosting us well-supported monophyletic lineages based on the and providing outstanding dive support during a four type specimens of the new taxon from the An- visit in July 2013 when we made observations and damans and three specimens of H. fourmanoiri collections of H. fourmanoiri. We also thank Con- from Brunei (Fig. 8). Although, the uncorrected servation International’s Indonesia marine pro- interspecific sequence divergence between the two gram for the financial support for this work and es- species is only 0.5%, 16s ribosomal RNA is highly pecially the Paine Family Foundation for their con- conserved due to its functional and structural roles tinued support of our taxonomic research. Mark and this gene is commonly used to infer relation- Allen and Glenn Moore of WAM provided curato- ships at deeper phylogenetic levels (Weisburg et al. rial support, including digital x-rays. 1991). However, it has been used successfully for species delimitation typified by low sequence di- REFERENCES vergence in other fish species (Aoyama et al. 2000, ALLEN, G. R. & ERDMANN, M. V. 2012. Chapter 3. Reef 2001; Papasotiropoulos et al. 2007). Fishes of Bali, Indonesia. In: Bali Marine Rapid Assess- Etymology: The species is named andamanensis ment Program 2011 (First English edition June 2012) (Eds. with reference to the Andaman Islands type locali- P. L. Mustika, I. M. J. Ratha, & S. Purwanto): 15-68, ty. Bali Marine and Fisheries Affairs, South East Asia Center Distribution: The new species is reliably known for Ocean Research and Monitoring, Warmadewa Uni- only from the Andaman Islands. Dooley & versity, Conservation International Indonesia, Denpasar, Jimenez (2008, figure 1, page 78) illustrated a 103 147 pp. ALLEN, G. R., ERDMANN, M. V. & HAMILTON, A. mm SL specimen (as H. luteus) from Bali (listed as M. 2010. Hoplolatilus randalli, a new species of sand tile- Sumatra on page 79 of their publication) that was fish (Pisces: Malacanthidae) from the tropical western Pa- purchased from a New York aquarium dealer. Al- cific with comments on the validity of H. luteus. aqua, though Bali is definitely a possibility for the collec- International Journal of Ichthyology 16 (4): 171-186.

11 aqua vol. 25 no. 1 - 25 January 2019 AQUA 25-1_ok.qxp_AQUA 05/02/19 10:45 Pagina 12

Hoplolatilus andamanensis, a n. sp. of sand tilefish from the Andaman Islands and confirmation of H. luteus as a junior synonym of H. fourmanoiri

AOYAMA, J., ISHIKAWA, S., OTAKE, T., MOCHIOKA, N., on morphological and molecular data. aqua, Internation- SUZUKI, Y., WANTANABE, S., SHINODA, A., INOUE, J., al Journal of Ichthyology 14 (2): 77-84. LOKMAN, P. M., INAGAKI, T., OYA, M., HASUMOTO, H., ESCHMEYER, W. N., FRICKE, R. & VAN DER LAAN, R. KUBOKAWA, K., LEE, T. W., FRICKE, H. & TSUKAMOTO, (eds). 2017. Catalog of Fishes. (http://researcharchive. K. 2001. Molecular approach to species identification of calacademy.org/research/ichthyology/catalog/fishcat- eggs with respect to determination of the spawning site of main.asp). Electronic version accessed 20 October 2017. the Japanese eel, Anguilla japonica. Fisheries Science 67: NELSON, J. S. 2006. Fishes of the World. 4th edition. John 761-763. Wiley & Sons, Hoboken, N.J., 601pp. AOYAMA, J., WANTANABE, S., ISHIKAWA, S., NISHIDA, M. & PAPASOTIROPOULOS, V., KLOSSA-KILIA, E., ALAHIOTIS, S. TSUKAMOTO, K. 2000. Are morphological characters dis- N. & KILIAS, G. 2007. Molecular phylogeny of grey mul- tinctive enough to discriminate between two species of lets (Teleostei: Mugilidae) in Greece: evidence from se- freshwater eels, Anguilla celebesensis and A. interioris? quence analysis of mtDNA segments. Biochemical Genet- Ichthyology Research 47: 157-161. ics 45: 623-636. DOOLEY, J.K. & JIMENEZ, L. 2008. Hoplolatilus luteus WEISBURG, W. G., BARNS, S. M., PELLETIER, D. A. & Allen & Kuiter, 1989, a junior synonym of H. four- LANE, D. J. 1991. 16S ribosomal DNA amplification for manoiri Smith,1964 (: Malacanthidae), based phylogenetic study. Journal of Bacteriology 173: 697-703.

aqua vol. 25 no. 1 - 25 January 2019 12